U) 5 a) = = CDI a Cet = 77 SX

Seno NS WAT 9 N LX S AY ca AI 3 x

+ =

JIYVYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31)

S31#V4911 LIBR

INSTITUTION

NOILALILSNI NVINOSHLINS S3IUVY811 LIBRARIES. SMITHSONIAN INST

NVINOSHLINS S3I4VHAIT LIB

NVINOSHLINS S31YVY917

o É z 2

zZ 6 6 . 5 N

wn a 2 n

: È = zi

5 | 5 a x 3

LIBRARIES SMITHSONIAN. INSTITUTION NOILNLILSNI NVINOSHLIWS S31

= u z u > ob u

4 = +0 = a ui

= = er

= È 3 = < ER =

< x = oc = ec

3 a. ' o) È © u

z — = - = Ag

ISTITUTION NOILOLILSNI NVINOSHLIWS S3IYVUGIT LIBRARIES” SMITHSONIAN” INST

6 TE ö — 6 >. —

== wo N _ ow _ #7 œ

E o \N 5 2 5 “Ag,

E > RIE = = ARE

5 N 5 5 Pr

= ig te SE D Z D

AIUVUGIT LIBRARIES SMITHSONIAN INSTITUTION, NOILMLILSNI- NVINOSHLINS S31)

REA = CAL Ay? Se =

z Hd = OS 4 z 4

3 ; 3 Nè BEE DE 4

E 2 END 2 E 2/7

5 2 5 2 5 2

ISTITUTION NOILNLILSNI SIINVYAIT LIBRARIES SMITHSONIAN INST

v = w =_= wo —

us 5 w 5 u D». 5

a = az = e Non.

< 2 .< = < LS =

x A o S co. pot =

ca o = To) Be 5

Da | = — = el =

318V4g17 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31

n = = = E z

DE y's PD = GMs NT

5 = È E À al:

= a = È 3 5 K

Hi te: Ho = o a -

NOILNLILSNI NYINOSHLINS S31YV#911 LIBRARIES SMITHSONIAN INST

n = v =. ere u =

= = = zh NS = =

2; 2 Æ ZR 2 8 N

2 E = = N 2 E

> = > = ee =

ve n 2 | un mn a

31UVYg1T_ LIBRARI ES SMITHSONIAN _ INSTITUTION, NOILNLILSNI_NVINOSHLINS | Sa |

5 = +5 = 8 NK 2

ra. we. 4% J pa œ a N, R oc

= < } = < zul N <

Cc œ Yi = œ = az

ZA en = m = K m

St a N \ sp & [A 9 Om È <| —& SL GC LA ES ST |

JS AO ad 5 tu < 7 SY x \}

7 3 E Ii ER CER

z “ate — = + z pr

ARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS Saiuvudi7 LIBRA

= E = rm z =

5 È 2 : = :

pol

Ci > = > : 2 >

Ee = = 2 = a

Ls m a m u m

= i v Z a , z u,»

NLILSNI NVINOSHLINS SAIUVHAIT LIBRARIES SMITHSONIAN INSTITUTION

= v = es u > È [2]

< = TL We = < NN =

5 = 6 AN 2 OT 5 SAN =

= 0% EE N 8 = \ Wr

E 2 = x 2 = 2

RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS SIIUYYAIT_

u > dd 5 NCT ” z

en Bi af = pes cox =

ty eel En m.

u MA 4 = per = =

PIZZE KL Cc ol & X =

— Ge O — O == O

ALILSNI _NVINOSHLIWS 53 IUVUA 11 LI BRARI ES_ SMITHSONIAN, INSTITUTION „NollN.

MO ° = © le ©)

> We 5 2 = be. 5

a YI D= 7 = ° 2 =

= N E È F . pi E

ARIES, SMITHSONIAN INSTITUTION, NOILOLILSNI _ NVINOSHLIWS Sa ruvugio_ ti! AR:

= Ca ° Op = AP

= 5 5 : 8 E 5

MT] D . 0 2m n or

oO Es : © x re) =

N 2 = z, = 2, =

=” = 5 2 5 5 D

NLILSNI_NVINOSHLIWS S3 INVYAIT LIBRARIES SMITHSONIAN_INSTITUTION,,

2 - 3 5 i di

= = = = Me “=

= _< <= < 5 | <

“ œ = o. S =

lo) m 3 Mm. = m

ET — = oO

= — z eal = —

RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31y#vVug171 LIBRA

= È = = È i

di > = =: > >

il = e 2 = 2

La m a m n m

z 6 di = Di D“ u

LNLILSNI NVINOSHLINS SIIUVUAII SMITHSONIAN _

ze 2) = (77) z u

= = „Z = < =

z = = a > =

ca Ou = x [e] 55 (©)

= 2 = zZ = 2

RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS S31uvaug11_LIBR:

u 2 u 5 ER u 5

ps IMRE ed = u = ao

397.77 2E = : = =

< ; +2 < <

N N

KA Tier ots |

Ree gr, ef 4 ij en

| a

9 or a

id 450

4 u, is 7 Pr er f he 0

a ef 1A Mi oe &

5 iv lo x | i È | i i!

| py

19 | 4 te se) iw sé

L, GA = . A et i 2 er

hi iv * “> P7 | p À a

A » ore è ai I y

w# 4 Wo Le Ù = >

k a ) , CS È 4 my oy

N rg

à Le

Ù NI, 7 ; ' rahe d

#4 Es he u sg f wir . fi bj

PEN ur

i kerkdeur mR OR

EUERLANDSE ENTOMOLDOGISCHE

pe NR, | i

FR ;

0 Hr

’ Da

maan wii

ay SAY hoy

mth NI iy,

(a ll

1 ap 18

}

PR TE 1

’ 4 4 Vy

ay I}

VEREINE |

= VU\ t

DEEL 123 1980

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

JUL 9 8 1981

> TEC

Tijdschrift voor Entomologie, deel 123, 1980

NEDERLANDSE ENTOMOLOGISCHE VERENIGING

BESTUUR (BOARD)

Voorzitten (Chaim an) ee EEE EEE R. H. Cobben

Mice=Moorzitter(Mice-President) ennn A. K. Minks

Secretans(Secretary) > eee eee RI I R. de Jong

AGGRESSIONI Rijksmuseum van Natuurlijke Historie,

Raamsteeg 2, Leiden 2311 PL

lesPenningmeesten(Mireasùrenl) PER L. H. M. Blommers

AGAR OSS. iti WANNE RE Herenstraat 102, Rhenen 3911 JH

2e Penningmeester (Treasurer Il) . .......... P. Oosterbroek

AGAheSS © tees EN: Baanstraat 2, Edam 1135 CB

Bibliothecanis(WlEipranian)e ne W.N. Ellis

Plantage Middenlaan 64, Amsterdam 1018 DH

Bidi@Memben) SEE RENE ER ees oe ore H. van Oorschot

TIJDSCHRIFT VOOR ENTOMOLOGIE

Redactie (EditorialiBioard) scence ie P. J. van Helsdingen, R. de Jong, J. Krikken,

M.A. Lieftinck, C. van Achterberg

AAARESS erneer LI ER CCR Rijksmuseum van Natuurlijke Historie,

Raamsteeg 2, Leiden 2311 PL

The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea.

Subscription rate: D.Fl. 200,— per year.

Afleveringen 1—3 verschenen 29.11.1980

Afleveringen 4 en 5 verschenen 9.v. 1980

Aflevering 6 verscheen 31.x.1980

ISSN 0040-7496

LEA CA IL

TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

Pe “uy

1] 9

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING ö 1981

* Een sterretje duidt aan een naam nieuw voor de wetenschap

* An asterisk denotes a name new to science

ARANEAE

Agelena 92, 98

Coelotes 93

consociata 92, 98

ferokus 100

lineatus 98

mimosarum 89

Pirata 116

piraticus 116

republicana 98

sarasinorum 89 sqq

simoni 89

Stegodyphus 89 sqq

terrestris 93

Uloborus 100

COLEOPTERA

bicarinatum 38

*ssp. birmanicum

[(B. drescheri) 1, 13, 15, 36

Bolbelasmus 7

Bolboceras |

*Bolbohamatum 1,5 sqq

Calanus 1, 7, 8, 11, 15, 20

cyclops 1, 6, 8, 14, 19

*drescheri 1, 7, 8, 11, 13,

nel 19,88

grande |

*ssp. indosinicum

[(B. drescheri) 1, 13, 14, 36

*kuijteni 1, 8, 12, 14, 32

laevicolle 1, 8, 11, 15, 27

laterale 1, 8, 12, 15, 33

*marginale 1, 8, 11, 12, 14,

[30

*meridionale 1,8, 11, 15,

[29

nigriceps 37, 38

*phallosum 1,8, 11, 15, 21

Odonteus 7

*pseudogrande 1, 8, 15, 23

*pyramidifer 1, 8, 11, 15, 28

*robustum 1, 8, 11, 15, 25

Scarabaeus |, 6

subglobosum I, 19

*syncopator 1, 8, 11, 15, 24

tumidulus 21

validus 6

veter |

DERMAPTERA

Forficula 90

DICTYOPTERA

americana 99

Mantis 98

Periplaneta 99

DIPLOPODA

Antheromorpha 71 sqq

bipulvillata 87

bistriata 71, 79, 85 sqq

bivittata 71, 81, 85 sqq

Brachytropis 71

comotti 71, 83, 85 sqq

coriacea 87

Dajakina 87

Diglossosternum 87

Eudasypeltis 87

festiva 85 sqq

Gigantomorpha 87

harpaga 85, 87

mediovirgata 85 sqq

melanopleuris 71, 77,

[85 sqq

minlana 85 sqq

miranda 71, 72, 76, 85 sqq

Nesorthomorpha 87

orophila 85 sqq

Orthomorpha 76 sqq

pardalis 71, 82, 85 sqq

Piccola 87

uncinata 85 sqq

DIPTERA

brunnipes 105 sqq

chamaeleon 105

conica 116

SSI

dorsata 108, 113, 116, 122

Drosophila 113

Hydromyza 113, 116, 121

Hydrophoria 116

livens 113, 116, 121

Neorhynchocephalus 290

Notiphila 105 sqq

riparia 115

stagnicola 105

tauscheri 290

HYMENOPTERA

acervorum 133, 200, 203

var. acutivalvis

[(M. italica) 309, 310, 314

aegyptiaca 169, 184, 244

[279

aestivalis 203

angustilabris 157, 177, 251

albifrons 132, 134, 139

[147 sqq, 193, 227

ssp. albovaria (M. albi-

[frons) 132, 134, 147, 149,

[196, 197, 200, 203, 204

*alcestis 162, 164, 270

*alecto 162, 164, 186, 273

alluaudi 321

*amanda 161, 182, 266

Andrena 131, 283

*angustilabris 157, 177, 251

Anthophora 131, 132, 133,

[192, 196, 200, 202, 203,

[210, 219, 227, 230, 233,

[251, 286, 289, 290, 321,

[331

Apis 139

armata 193, 194, 196, 202,

[204

assimilis 171, 187, 318

ashabadensis 192, 336

aterrima 204, 206, 208

atra 139, 204, 206

baeri(i) 139, 158, 171, 187,

352 TIIDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, 1980

(210, 211, 215, 243, 260,

[282

bipunctata 204, 206

bituberculata 306

*brevipila 156, 191, 235,

[239

caesareae 275, 287

calabrina 204 sqq

ssp. callura (M. separata)

[132

canariensis 145, 192, 322

candiae 337

*candida 170, 243, 268

canescens 133

caroli 175, 190, 323

chinensis 150, 211, 218

corpulenta 160, 181, 260

var. crassicornis (M. itali-

[ca) 309, 314

crassipes 133

crinipes 202, 203, 219

Crocisa 139, 204, 206

curvispina 173, 189, 320

diacantha 239

duodecimmaculata (Me-

[lecta) 148, 151, 210, 213,

[217

duodecimmaculata

[(Nomada?) 139, 194, 212

ebusana 254, 325

eczmiadzini 228, 270

edwardsii 132

egyptiaca 244

Fumenes 90

Eupavloskia 134, 209, 313

eversmanni 192, 228, 239,

1333

*excelsa 152, 217

fasciculata 204, 227, 233

*festiva 167, 186, 293

*fulgida 158, 178, 219, 226

[nota, 257

fulvitarsis 196, 203

fumipennis (Anthophora)

[251

*fumipennis (Melecta) 158,

(178, 248

funeraria 313

var. gigantea (M. albi-

[frons) 193, 196

glasunowi 261

*gracilipes 175, 176, 332

grandis 165, 185, 286, 309

*guichardi 162, 179, 254

Habropoda 132, 294

hispanica 210, 286, 290

histrionicus 194

*honesta 161, 182, 264

italica 132, 174, 191, 203,

[206, 308, 309

ssp. jakovlewi (M. duode-

{cimmaculata) 151,

[215 sqq

kashmirensis 192, 337

kuschakewiczi 261

lentina 331

leucorhyncha 133, 176,

[192, 239, 308, 325, 330

lindbergi 244

luctuosa 139, 155, 194,

[196, 197, 203, 227, 233,

[286, 330

var. mediterranea (M. al-

[bifrons)196, 197, 313

*megaera 175, 185, 243,

[301

Melecta 129 sqq

var. meridionalis (M. itali-

[ca) 309, 313, 314

var. minima (M. luctuosa)

[196, 197

ssp. nigra (M. albifrons)

[132, 133, 147, 149, 203,

(204, 206, 324

nigripennis 206, 334

nivosa 157, 179, 257

Nomada 131, 139, 143,

(194, 212, 213

notata 205

novellai 196, 197

octomaculata 275, 279,

[334

obscura 134

orbatus 194

oulskii 283

Paracrocisa 261

parietina 202

personata 192, 202

plagiata 202

plurinotata 210, 213, 215

*prophanta 158, 170, 290

pseudoarmata 196, 197

Pseudomelecta 139, 239,

[266, 282

Psithyrus 135

punctata 139, 193, 194,

[196, 204, 227

pygialis 308

quadripunctata 309, 314

quadripustulata 314

quatuordecimpunctata

[210, 233

retusa 227, 233

robusta 290

rufipes 204

rutenica 153, 190, 233

separata 132

sexta 194

sibirica 173, 236, 238

sicheli 331

soederbomi 210, 213

ssp. simulatrix (E. obscu-

(ra) 134

*solivaga 145, 180, 279

ssp. taormina (M. leu-

[corhyncha) 133, 177, 192,

[206, 324, 330

tarsata 294

testaceipes 205, 206, 334

Thyreus 140, 143, 194

tibiopunctata 331

transcaspica 168, 183, 261,

[275, 279

Triepeolus 138

*tuberculata 134, 167, 188,

[289, 300

turkestanica 172, 189, 239

ODONATA

Coenagrion 116

elegans 116

Erythromma 116

Ischnura 116

najas 116

pulchellum 116

ORTHOPTERA

anatolica 52

astyla 41 sqq, 63

beybienkoi 45

ssp. bimucronata (E. cha-

[brieri) 45

ssp. brummeri (E. cha-

[brieri) 45

chabrieri 39, 40 sqq

cretica 41 sqq, 64

danconai 39, 54

epirotica 40, 42, 47, 52

ssp. epirotica (E. cha-

[brieri) 39, 45, 52

Eupholidoptera 39 sqq

festae 59

ssp. festae (E. smyrnensis)

[39

forcipata 41 sqq, 45, 52

ssp. galvagnii (E. cha-

[brieri) 45

ssp. garganica (E. cha-

[brieri) 40, 47, 49

gemellata 41 sqq

Gryllotalpa 90

hesperica 52

*icariensis 39, 41 sqq, 61

ssp. kaltenbachi (E. cha-

[brieri) 39, 45, 47

Karabagia 39, 65

latens 41 sqq

ledereri 52

*leucasi 39, 40, 42, 50

Locusta 40

ssp. magnifica (E. cha-

[brieri) 39, 45

marani 45

megastyla 39, 40, 42,

[47, 54

Olynthoscelis 65

palaestinensis 52

pallipes 41 sqq

Pholidoptera 47

prasina 41 sqq, 60

ssp. schmidti (E. cha-

[brieri) 39, 40, 45, 47

smyrnensis 40, 42, 47

57559

spinigera 41 sqq, 61

Thamnotrizon 47, 49

ssp. usi (E. chabrieri) 45

uvarovi 39, 42, 65

Uvarovistia 39, 42, 65

zebra 65

AMPHIBIA

Rana esculenta 116

AVES

Fulica atra 121

PLANTAE

Acacia 90

Acorus calamus 111,

[122, 124

Anchusa 205

Anchusa azurea 289

Anchusa officinalis 244

Asphodelus 211

Brassica oleracea 310

Caragona arborescens 327

Echium 332

Ficus 90

Glomerella cingulata 113

353

Glyceria maxima 122

Iris pseudacorus 111, 122

Lamium album 233

Lavendula 211

Lithospermum officinale

[233

Mentha aquatica 108, 122

Nepeta cataria 233

Nuphar lutea 105 sqq

Nymphaea alba 105 sqq

Nymphaea candida

[105 sqq

Nymphoides peltata

[105 sqq

Ophrys 133, 134

Ophrys reinholdi 134

Ophrys sphecodes atrata

[134

Phragmites australis 122

Robinia pseudacacia 327

Rosmarinus 211

Rumex hydrolapathum

[108, 111, 122

Salvia 205

Saponaria ocymoides 205

Scirpus lacustris 108, 111

Typha angustifolia 108,

[111, 122, 124

Typha latifolia 125

Zizyphus 90

25 Jif ee

ru A Mon A

77)

re sn di à

can khad

rn ad

Ps Tri med

N terre

prs U AR

en. Aad ha

sot!

LE RIN ee

rT? oe

West) eg ih

ar (hives big)

elkom wists vii

vor i è i ‘

SS. with vances

090. wis dry È

>> ps a ne en

rl nu À

ren) CL

a” Lois!

een veh if pall

IE ie eet e 11

ro) eT aa de. 1

Sine qm

Yi — ASMA 6:

er Sain

mi pro 6) OW eu

paris ER

AREA

beet iP mik fes aan

we Tha N

bet me 9 X

wit ih nj.

paci LL PR:

it) run

si è

q all ar,

2 er

tte sed)

es all? tap 7

IT doté

-- Mr Ba

TNT Pay peo Mat

11 Pin is A

Ni caso ti papetti

af cc PUT li zz TS

DCE." PE

u Lr» |

cinivgnà ER Sorrall)

ré

«4

‘

re,

416 ;

nin à As

Vom ve

i {

’

4 ta è sa

4 “+

po ail) see ola I 6

INHOUD VAN DEEL 123

Bradoo, B. L. — Feeding behaviour and recruitment display in the social spider Stegodyphus

Carasmnorumiarschi(ATANEACMETESI 20) a. eration iene mien 0

Brock, T. C. M., zie Velde, G. J. van der.

Jeekel, C. A. W. — A revision of the Burmese Paradoxosomatidae (Diplopoda, Polydesmida)

in the Museo Civico di Storia Naturale at Genova (Part III) ..............

Krikken, J. — Bolboceratine scarabs of the Oriental genus Bolbohamatum nov. (Coleoptera,

Geotnunidae) RA IT cl cae lo ne nt nn

Lieftinck, M. A. — Prodrome to a monograph of the Palaearctic species of the genus Melecta

Batreillesl8021(EIymenoptera, Anthophornidae) . acs TT oe eee ces:

Velde, G. van der, & T. C. M. Brock. — The life history and habits of Notiphila brunnipes

Robineau-Desvoidy (Diptera, Ephydridae), an autecological study on a fly

assoglatedivuithinymphaeidivegetallonse mei EE sie ee) cee enone

Willemse, F. — Classification and distribution of the species of Eupholidoptera Ramme of

Greecel(OrthopteramliettisoniordeaNDECticinae) RR RR

105

ae)

ager

bj |

"NE

moines oS ew ail en al aval oa

einen di te rial AN

ie |

eects à ii odori, elm TAI

Ved awe £ IN

hua 88 eej mang, ue it] 440 Vo Mara BAT

Bemet vo, ae We ytd peta dit af) ta Arijan en wal are

co dat omis

eer, Aa y soft tee Vinti eue Set — pend w DY

Ai ape the pianti de ar rau (chiesta

na benk i 11116 LAAN

| Vesna egt) breit are

EDEEL 123 AFLEVERING | 1980

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

J. KRIKKEN. — Bolboceratine scarabs of the Oriental genus Bolbohamatum nov.

(Coleoptera, Geotrupidae), p. 1—38, figs. 1—71.

Tijdschrift voor Entomologie, deel 123, afl. 1 Gepubliceerd 29-11-80

BOLBOCERATINE SCARABS OF THE ORIENTAL GENUS

BOLBOHAMATUM NOV. (COLEOPTERA, GEOTRUPIDAE)

J. KRIKKEN

Rijksmuseum van Natuurlijke Historie, Leiden

With 71 text-figures

ABSTRACT

The genus Bolbohamatum nov. is proposed to accommodate four species hitherto combined with

Bolboceras Kirby, and nine new species. The genus, characterized by a spine-bearing lobe separating

the middle coxae, is endemic in the Oriental Region. The 13 species are keyed, illustrated, and their

| distribution is recorded. The following novelties are described: Bolbohamatum drescheri drescheri

| (Java), drescheri indosinicum (Indo-China), drescheri birmanicum (Burma, Laos); B. kuijteni (India),

| marginale (India), meridionale (India, Sri Lanka), phallosum (India), pseudogrande (India), pyramidifer

| (India), robustum (India), syncopator (India). The four old species are B. cyclops (Olivier) (type-species;

synn. nov. are Scarabaeus veter Fabricius, Bolboceras subglobosum Westwood), calanus (Westwood),

laevicolle (Westwood) (syn. nov. Bolboceras grande Westwood), laterale (Westwood). For three names

lectotypes are designated. The phylogeny of the genus is briefly discussed.

INTRODUCTION

The larger Bolboceratini of tropical Asia, i.e. those with a total length between

13 and 23 mm, belong nearly all in one monophyletic group, for which here a new

generic name is proposed: Bolbohamatum. This monophyly is inferred from the

presence of a characteristically shaped, spine-bearing metasternal lobe between

the middle coxae (fig. 56), a feature that seems to be unique within both the tribe

and the family. The new genus comprises four species described more than a

century ago, and nine described below for the first time. Two names possibly

relating to Bolbohamatum species remained dubious, their types having

disappeared.

With some experience the males of most Bolbohamatum species can easily be

identified on the basis of the diagnostic information presented in this paper; in

some cases the examination of the genitalia will prove to be indispensable. The

identification of the females will present greater difficulties, because of their

rather poor, uniform cephalic and pronotal ornamentation, and because they

cannot always be associated with the opposite sex of the same species. The former

factor also renders the identification of the occasional underdeveloped male

troublesome. More work on Bolbohamatum is certainly needed. It seems,

nevertheless, undesirable to delay the publication of the present results, were it

alone for the fact that the collections of this group cannot be expected to grow

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

ae annen

asen

Tom

A gt géné oe

Fig. 1. Bolbohamatum drescheri drescheri, holotype (male) from Java. Length ca. 18.5 mm.

KRIKKEN: The genus Bolbohamatum 3

significantly in the years ahead. Any supplementary information regarding

Bolbohamatum will be included in a proposed review of the Asian bolboceratine

genera. For more about the scope of this series of papers on Bolboceratini, see

Krikken, 1977b.

TECHNICAL REMARKS

Some points have already been discussed in my Bolbogonium paper (1977b). In

this treatment of Bolbohamatum I have kept the descriptions of the novelties rather

concise, and of the other species I give descriptive details only as far as they are

taxonomically important (identification, variation, sexual dimorphism).

The parameres and other parts of the phallus are in Bolbohamatum usually

strongly sclerotized and provide excellent characters for the identification to

species. The homologies of these characters, however, are as yet poorly

understood. The phallus proper moves within a complex structure (*sagulum),

consisting of a (ventral) bottom sledge (*fundus), a pair of lateral flaps, and a roof

(*tectum). The parameres frequently have sclerotized protrusions (*paramerites)

and sacks (*sacculi); the latter seem to be inflatable. The dorsal side of the basal

capsule (in fact consisting of scarcely separated distal and proximal parts) is open,

i.e. covered by a membrane. The median lobe (with the penis proper, the internal

sac) is usually concealed by the aforesaid parts. The positions of these parts (terms

marked with an asterisk proposed ad hoc) are explained in four diagrams (figs.

5—8). The genital characters of the females, as well as the taxonomic value of the

sagulum of the males, need further study.

The arrangement and development of the fossorial protrusions on the outer

(anterior) side of the middle and hind tibiae are more or less characteristic but

difficult to describe. In Bolbohamatum these protrusions are bidentate-bilobate

and decreasingly pronounced proximad. Usually there is one complete anteapical

protrusion (i.e. there are two denticles or lobes connected by a complete crest),

denoted Ic. Proximad there are some paired incomplete protrusions (i.e.

protrusions lacking a complete crest), numbered 2, 3, n, from apex to base; these

are either placed opposite to each other, a situation denoted n po, or shifted

relative to each other, a situation denoted n ps. Near the tibial base the protrusions

become single and/or obsolescent. In the descriptions given below a formula

lc—2ps—3ps—4 then means that from apex to base the outer side of the tibia is

provided with one anteapical pair of protrusions connected by a crest, two pairs

lacking a crest, in both cases the superior protrusions being shifted proximad,

while, finally, there is a single protrusion near the tibial base. The protrusions 2 on

the hind tibia (figs. 54, 55) are used for diagnostic purposes (see synoptic table,

character 15).

The dentation of the fore tibia is described as a + b, in which a is the number of

large distal denticles, b the number of smaller denticles, the size of which

decreases proximad.

Lectotypes are selected if in the original description under a given species-group

name at the same time: (1) no specimen is singled out as “type” (the type) or

holotype; (2a) there are phrases from which it can be inferred that there was more

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Figs. 2, 3. Approximate known distribution of Bolbohamatum species. Numbers refer to species as

numbered in text. 2, species 1—8; 3, species 9—13. Numbers between square brackets refer to

insufficiently detailed locality data.

KRIKKEN: The genus Bolbohamatum 5

than one specimen at hand; or (2b) there are no phrases from which it can be

inferred that there was a single specimen at hand. This rigid formulation seems

necessary because the practice of lectotype selection varies among taxonomists.

Most topographic names have been checked against those occurring in The

Times Atlas of the World and the Gazetteers of the U.S. Board on Geographic

Names. Places not located with a precision better than a hundred km are followed

by an asterisk, and, if mapped, the symbol concerned is placed between square

brackets. The spelling of topographic names as given on the labels is frequently

maintained, in order to enable easy recognition of the specimens examined. The

data of two old species have been condensed by separately giving localities,

months, collections.

In the drawings the scale lines all represent | mm.

The collection abbreviations used are as follows:

BH Zoologisches Museum der Humboldt-Universitat, Berlin D.D.R.

BM British Museum (Natural History), London.

L Rijksmuseum van Natuurlijke Historie, Leiden.

B Museum National d'Histoire naturelle, Paris.

SMT Staatliches Museum für Tierkunde, Dresden.

Other collections are mentioned in full.

Genus Bolbohamatum nov.

Generic diagnosis. — Metasternum with anterior lobe narrowly separating

middle coxae, anteriorly always with a small spiniform protrusion (fig. 56), midline

of lobe more or less raised; metasternal disc rhomboid in outline. Head of males

with pair of tubercles on clypeus or between eye-canthi (figs. 1, 9, etc.); frons of

females (fig. 59) with transverse ridge; surface behind cephalic protrusions of

males frequently impressed-callose. Pronotum of males with median and lateral

protrusions (figs. 1, 9, 10, etc.); surface between paramedian and lateral protru-

sions usually concave; lateral tubercles in some cases obsolete; females with feeble

protrusion(s) on anterior side of pronotal disc. Aedeagus usually strongly

sclerotized, structure complex, including various accessory elements. Proximal

surface on antennal club segment | with glabrous polished area (fig. 57). Pronotum

entirely marginate, though base not always distinctly ridged.

Outline of left mandible in dorsal view variable, from simply arcuate to sinuate-

lobate (figs. 51 —53). Labrum with fine transverse ridge. Outline of clypeus (figs. 1,

9, etc.) approximately trapeziform, with distinct perimarginal ridge. Frontolateral

ridge well pronounced. Eye-canthus (figs. 1, 9, etc.) with straight anterior border,

anterolateral angle more or less angulate. Eyes not completely divided by canthus

and temporal lobe. Dorsally visible area of eye-canthus usually little larger than

area of eye-canthus. Scutellum deltoid (figs. 9, etc.), sides more or less sinuate;

ratio length/width 1—2; base finely ridged. Apex of pronotum with narrow velum.

Elytral base without well-defined ridge. Elytral epipleuron reaching apico-sutural

angle. Elytron with 7 striae between suture and humeral umbone, | reaching side

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

of scutellum (figs. 9 etc.), stria 2 curved and more or less effaced near base; others

reaching base; all striae (except stria 2 in two species) superficially impressed and

interstriae flat or slightly convex. Antennal club large, but not thicker than length

of pedicel and flagellar segments combined. Anterior paramedian costae of

prosternum distinct. Posteromedian part of prosternum simply bulbous. Fore tibia

with 7—10 external denticles, their size decreasing proximad. Fossorial elevations

on middle and hind tibiae (figs. 54, 55) bidentate-bilobate, their size decreasing

proximad. Fore tibial spur long, large, with acute apex. Habitus, fig. 1. Colour

uniformly brown to yellow. Medium-sized to large (total length 13—23 mm).

Type-species. — Scarabaeus cyclops Olivier.

INDIA BURMA &c,

hae) it a

: i UE oO @ +

3 5 Dn 3 È Se Fe

2 ER rt . È

ui Yoon W ba 8 Ot lo) di dii Vi &

a: SO ao DEFo: st sn con ge

a Ord goog Si art ob = U 0:0

oO rt is} =] (00) a ce) u fen Med 4 ‚U

“Di Cd Sm Of Dy © Ed HET

oe (Opens Tats Ui Kr A (= a Sd ae

la Ala a |

l4a a a

5d all à a €

17ai 2 b 2

16a a, a

SSS Ss, = = = = = ee ee = = de om ee oe) en em ee

Fig. 4. Phylogenetic diagram summarizing the present classification of Bolbohamatum, as based on the

stridulatory granules of the pygidium (character 16 in synoptic table, see text), the structure of the

phallus (17), the pronotal protrusions (5), the disposition of the first elytral stria (14), and the shape of

the cephalic tubercles (1). Letters denote character states; species-groups shaded. See also section

Phylogeny in text.

Affinities. — The group around Bolboceroides validus (Klug) (Krikken, 1978)

seems closely affined to Bolbohamatum judged from the strong overall similarity in

external characters. The metasternal spine, however, certainly is a unique

differentiating character of Bolbohamatum. Furthermore, the males of the

Bolboceroides validus group are characterized by the presence of a simple

transverse clypeal ridge, not connected with the margin, whereas Bolbohamatum

KRIKKEN: The genus Bolbohamatum U

males always have a pair of isolated tubercles on either clypeus or the anterior part

of the frons. The only representative of the validus group in the Oriental Region

seems to be Bolboceroides carinicollis (Laporte), the other species being Arabian

and Afrotropical.

Bolboceras Kirby, with which four Bolbohamatum species were combined up till

now, is very remotely related to Bolbohamatum; Bolboceras is a, probably junior,

synonym of Odonteus Samouelle (cf. Krikken, 1978: 301, footnote).

Distribution and composition. — Thirteen species in the Oriental Region, with

transgression of one species into the Palaearctic Region (figs. 2, 3). Only two

species occur outside India, one on Ceylon, the other ranging eastward from

Burma to China and Java. The gap between Java and the continental range is also

found in Bolbelasmus (Krikken, 1977a: 280). The records from Karachi and Taiwan

need confirmation.

Infrageneric dissimilarities. — For the identification to species and subspecies it

is crucial to study well-developed males. In case of doubt, the characters of the

male genitalia are always decisive. On the whole, 17 characters (classified below)

proved to have practical diagnostic value. In the key to the males the genital

features (character 17) are used first to delimit species-groups, the external

features coming second. Three of four species-groups thus delimited are assumed

to be monophyletic, primarily because the structure of their aedeagus is unique.

The three other groups contain only one species each. With the aid of the synoptic

table of characters preceding the analytical key most Bolbohamatum males (those

of phallosum and calanus excepted) should be identifiable on external characters.

As stated above, in the introduction, the identification of females remains very

difficult, and in some cases seems impossible. The key to Bolbohamatum females

given below only provides some guide-lines. Under each species the specific

characters are briefly re-discussed (in the paragraphs headed Identification).

Individual variation. — Most conspicuous is the reduction of pronotal

protrusions and cavities, if this is not already a character of the species. The

topography of these modifications, reduced or not, however, always remains

essentially the same. Such reductions in shape, noticed in all the species available

in good series, have not been described in detail under each species. One distinct

case of geographic variation has been found (B. drescheri).

Bionomics. — Unknown; probably burrowers; collected at light.

Phylogeny. — Evolution in Bolbohamatum can, in my opinion, only be discussed

if four points are accepted: (1) the genus is monophyletic; (2) in the structure of

the phallus there is a trend towards greater complexity (character 17, see synoptic

table and explanation); (3) a pronotal ornamentation like in B.calanus is

plesiomorphous, any reductions and other modifications being, eo ipso,

apomorphous (character 5); (4) some unusual character states (1b and 14b) are

apomorphous. Complexity in the phallus is here understood as development of

paramerites and other elements supplementary to a relatively simple structure like

that in B. calanus (figs. 27, 28).

It could then be argued that B. cyclops is the most strongly derived member of

the genus, standing isolated by its peculiar pronotal ornamentation and by its very

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

peculiarly modified phallus; it seems impossible to link this species with any of the

others. The pseudogrande and laevicolle groups might be cladistic twins, because

the ventral paramerites of the phallus seem homologous, whereas transitions from

one type to the other can easily be imagined. The robust basal capsule of the

phallus of B. marginale, in combination with its pronotal ornamentation, suggests a

link between the laterale and drescheri groups. Both groups lack the row of

stridulatory granules on the pygidium. The position of B. meridionale remains

enigmatic considering its aedeagus, but that species is here placed near pyramidifer

because of the similar cephalic and pronotal ornamentation. Finally, there are the

apparently primitive calanus and phallosum, with a perfect external overall

similarity, the latter, however, having a modified phallus.

These speculations, plus the known distribution of the species, are summarized

in a phylogenetic diagram (fig. 4), which clearly shows with which “‘loose ends” we

are left. The overall situation suggests that Bolbohamatum evolved on the Indian

subcontinent, and spread in a relatively late stage (possibly after India reached

Eurasia) through Burma into Sundaland and China. It is remarkable to see that the

only species occurring East of the mountains on the Indian-Burmese border shows

signs of subspeciation.

SYNOPTIC TABLE OF Bolbohamatum MALES

Characters

i) ne) 3 4 5 6,7 8 9 10 SM A2 IS A See ae

. cyclops

. calanus

. phallosum

. Syncopator

. pseudogrande

. robustum

. laevicolle

. pyramidifer

. meridionale

. marginale

. kuijteni

. laterale

. drescheri

(op (ap (ep

opr ppp p po

io”

NEOTO OPA N RB

—

ep @ pp gag a

ocean» pc pp GB OO

DUIDE SWO) loi EC ho

seer fp pp po p LP pw

rr pp P pp pw

9797840730797 Sis: Bar

ee À JD pp Op pp pp LH

Soos

(ono) enteren eh Toner fom fog Nester

30700020 10.89, 050 0920.00

SS ©. 2.8970 © 0° FG

DET IK Poo O° one 9

OSS aos IS u Cem IRD IN

PD co ca pp pop pp po po so pp pw

ep» pe cp pe TETE pp

Coc pp pp po po pp p mp m

Le)

Explanation of characters and character states (see comparative illustrations):

*1. cephalic tubercles more or less dentiform (a), pyramidiform (b).

2. cephalic tubercles isolated and placed simply on clypeal disc (a), together on

an elevation (b), on or against lateral margin (c); placed between eye-canthi

(d).

3. paraocular ridges straight (a), curved laterad along eye (b).

dorsal outline of left mandible sinuate-lobate (a), simply arcuate (b).

*5, paramedian tubercles on pronotum placed close together on common

KRIKKEN: The genus Bolbohamatum 9

protrusion (a), well separated (b); paramedian tubercles poorly developed or

absent (c); pronotum with large, laterally angulate median protrusion (d).

6. lateral tubercles of pronotum well developed (a), very poorly pronounced or

absent (b).

7. (related with 5:) paramedian tubercles on pronotum separated by less than/

equal to interocular distance (a), separated by more than interocular distance

(b).

93 Tisweied

paramerite

sacculus

culus

aiaweied

sac

paramere

flap

fundus

Figs. 5—8. Some terms relating to male genitalia. S—7, diagram of phallus, dorsal (5), ventral (6), and

lateral views (7); 8, sagulum, unfolded. Paramerites black (also in figs. 25—41).

=17.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

(related with 6:) lateral tubercles on pronotum separated by less than/equal to

distance between anterolateral angles (a), separated by more than distance

between anterolateral angles of pronotum (b).

. (related with 5:) median cavity of pronotal disc deep, very distinct (a),

shallow (b), obsolescent or absent (c).

. lateral impression of pronotum (adjacent to (para)median protrusion(s))

deep, very distinct (a), shallow (b), obsolescent or absent (c).

. primary punctation of pronotal disc behind (para)median protrusion(s)

sparse (punctures separated by more than 5 times their diameters) or

indistinct ( x 25) (a); primary punctation abundant or dense (b).

. secondary punctation of pronotal disc sparse (punctures separated by more

than 5 times their diameter) (a), abundant or dense (b).

. elytra distinctly opaque, due to microreticulation (a), shiny (b).

#14:

elytral stria | scarcely, virtually equally impressed over its entire length, and

equally punctate over its entire length (a); stria 1 distally more strongly

impressed and more strongly punctate (b).

. distal pair of unconnected protrusions on (right) hind tibia placed more or

less opposite to each other, 2po (a), superior protrusion distinctly shifted

proximad, 2ps (b).

. pygidium with distinct transverse row of stridulatory granules at some

distance from apex, elytral file distinct (a); “stridulatory” (strongly

infuscated) granules scattered near apex, not transversely arranged, elytral

file poorly developed or absent (b) !).

phallus of: cyclops type, with spatuliform protrusions (a); pseudogrande type,

with sledge- or collar-like paramerites (b); /aevicolle type, with crossed

paramerites (c); laterale type, with short, simply reflexed paramerites (d);

drescheri type, with reduced parameres and enlarged basal capsule (e);

different from preceding alternatives (z).

Between parentheses (in table above): character state poorly pronounced;

oblique dash: both states occur (subspecies); dash: transitional state occurs; zero:

not applicable; asterisk: used for phylogenetic discussion.

KEY TO SPECIES OF Bolbohamatum?)

Males, with pair of protrusions between eye-canthi or on clypeus ...... 2

Females, with transverse frontal ridge, no pair of isolated tubercles; pronotal

!) Whereas the stridulatory capacities of state (b) remain doubtful, those of (a) can be demonstrated

clearly by gently rubbing the abdomen (after relaxation) in its natural position against the elytral tips.

The elytral file is a series of closely arranged fine transverse ridges (distinct at x 50 and more) on the

juxtasutural costa on the inside of the elytral tips. In the laterale group of species the elytral files

seem fully effaced (magnification x 50), in drescheri they are small but distinct.

?) For more characters, see synoptic table, and paragraphs under each species headed Identification.

KRIKKEN: The genus Bolbohamatum 11

ornamentation poorly developed. TENTATIVE KEY; names between square

brackets: actual females unknown, characters established by inference .. 15

Basal capsule of phallus ventrally with paired spatuliform projection (fig. 26);

aedeagus with similar projection and other characteristic details. Pronotum

with broad, mostly laterally angulate protrusion (fig. 9). B. cyclops

ERRE OIE) hie AG TO sprees SELLER I AM ae i). eal 1. cyclops

EESTE Er ent xh cmt) wee. ARC ett) shall SESIA 3

Aedeagus ventrally with pair of crossed spatuliform paramerites (figs. 35—37).

VENI CO lE ETOU PISTE NAS. Velbon. TOLONE neu Mael eS 10

AREAS tere : .. . : Ne URINA) LOI, Weleda +

Apex of parameres dorsally with short reflexed paramerites (figs. 38—40)

(check carefully in case penis is extruded). B. laterale group ......... [2

EEDE CN EEn EE ee Btn 5

Parameres and other parts of aedeagus relatively small, usually largely hidden

in the greatly enlarged basal capsule of the phallus (fig. 41). B. drescheri

SOD. le oe EEE TEE 13. drescheri, with subspecies, 14

Parameres and basal capsule different. If similar, then elytral stria 1 distally

more strongly impressed and more strongly punctate, see 12 ......... 6

Aedeagus ventrally with sledge- or collar-like, laterally reflexed paramerites

eN 8. pseudogrande group . M... 55... :.. es MONT. 9

enn COMES > 8 VB ORE SEO 9)

Cephalic tubercles pyramidiform (fig. 17). Paramedian tubercles on pronotum

poorly developed or absent (fig. 17), other protrusions and impressions ditto.

Fossorial elevations 2p on hind tibiae shifted. Aedeagus, fig. 32. B. meridionale

COTES) «lasso REESE IR FR 9. meridionale

Cephalic tubercles dentiform (figs. 10, 11) and aedeagus different. Pronotal

protrusions usually well-developed (figs. 10, 11). Fossorial elevations 2p on

hindtibiae (sub)opposite. B>talanusgroup ..:..12. or. weelde. 8

ENE SOS A e A nn ARR he 2 à 2. calanus

PETER ZORIO TEN... en: 3. phallosum

Clypeal tubercles connected at base (fig. 12). Fossorial elevations 2p on hind

tibiaeepposite (fiens4). Aedeagus, fig. 31 tenta 5. pseudogrande

Clypeal tubercles simply separated (fig. 14). Fossorial elevations 2p on hind

timimesnited. Aedeagus fes 33534 .. . Ei i... a 4. syncopator

. Protrusions between eye-canthi pyramidiform (fig. 16). Pronotal ornamen-

tation poorly developed (fig. 16). Aedeagus, fig. 37 ........ 8. pyramidifer

Protrusions on clypeus more or less dentiform (figs. 13, 15). Pronotal orna-

Memanon usually wellideveloped 4... mone A le. 11

. Paramedian tubercles of pronotum closely approximated (fig. 13). Dorsal

outline of left mandible sinuate-lobate (fig. 52). Elevations 2p on hind tibiae

MAMI EMME. 55), Aedeasus, fig. 30.04. bok ed RL ee 6. robustum

Paramedian tubercles of pronotum separated by distinct impression (fig. 15).

Dorsal outline of left mandible simply arcuate (fig. 51). Elevations 2p on hind

MEIC opposites Aedeasus MPS. 39 ret ee oe Gus de. 7. laevicolle

. Elytral stria 1 virtually equally impressed and equally punctate over the entire

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

length. Pronotal disc heavily punctate. Elytra subopaque. Clypeal tubercles

transverse; low:Aedeagussfign38 05M TIA ZERI. 12. laterale

Elytral stria 1 distally more strongly impressed and more strongly punctate

than proximally. Pronotal disc lightly punctate. Elytra shiny. Clypeal tubercles

dentiformi high) moreno SLED NEI. ISSN, WURDEN. eG dare 13

. Paramedian tubercles of pronotum closely approximated, lateral impressions

usually deep, well defined (fig. 20). Phallus (fig. 40) with enlarged basal

eapsulerls «Men AHO? OBR LAAT) Te Daas 10. marginale

Paramedian tubercles of pronotum widely separated, lateral impressions

shallow; ill\defined (fig. 19) Aedeagus, fig. 39 2.5". 91 ORGAAN 11. kuijteni

cyclops calanus

pseudogrande

Figs. 9—12. Bolbohamatum males; dorsal contours of fore-body. 9, cyclops, Nepal; 10, calanus, Sylhet;

11, phallosum, holotype; 12, pseudogrande, holotype. — 10—12, same scale.

KRIKKEN: The genus Bolbohamatum 13

14. Occurring on Java. Ratio distance tips of paramedian pronotal cones/distance

anterolateral angles less than 0.5; axes of cones inclined to midline (fig.

BORAEOUIEN E01 left no vee: i nea She Lire 13a. drescheri drescheri

— Occurring in Burma and Laos. Ratio distance tips of paramedian pronotal

cones/distance anterolateral angles more than 0.6; axes of cones upright or

welnedilatetadi(fie; 22) uns hean.brarsure. dieet. 13c. drescheri birmanicum

— Occurring in Indo-China and China. Ratio variable, in Indo-China usually ca.

0.5; axes of paramedian pronotal cones usually upright (fig. 23). Populations of

Vietnam and Cambodia here named .......... 13b. drescheri indosinicum

robustum

laevicolle pyramidifer

Figs. 13—16. Bolbohamatum males; dorsal contours of fore-body. 13, robustum, holotype; 14,

syncopator, holotype; 15, laevicolle, Bombay; 16, pyramidifer, holotype. — 13, 15, same scale; 14, 16,

same scale.

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

ISO cure onl Savard nn DI Gea 1e ST sada. atk 13a. drescheri drescheri

— Occurring in Indo-China ................. 13b. drescheri indosinicum

— Occurring in India, Sri Lanka, Burma, Laos, China etc. ............ 16

16. Elytral stria 1 distally more strongly impressed and more strongly punctate

than proximally) ten 2.0 BGG! a anton’ 1613, 10. marginale, [1 1. kuijteni]

— Elytral stria | virtually equally impressed and equally punctate over the entire

length 0, 2 0060 „MENUIRES DUES AI DIE RI ISDN EE 17

17. Pronotum and head heavily punctate. Anteromedian ridge on pronotal disc

simply arcuate (fig. 59). Dorsal outline of left mandible sinuate-

RODE arn es eee AL Co SRR LAI IS IT SEES 1. cyclops

— Pronotum and head less heavily punctate; or not agreeing with the other two

characters: SEE i. WGA IONE. EHEM e C ES 18

meridionale

laterale

kuijteni marginale

Figs. 17—20. Bolbohamatum males; dorsal contours of fore-body. 17, meridionale, holotype; 18,

laterale, Nagpur; 19, kuijteni, holotype; 20, marginale, holotype. — All same scale.

KRIKKEN: The genus Bolbohamatum 15

. Superior (stridulatory) granules on pygidium arranged intransverserow . 19

ENE SEALED ICAL ADER | 24. ce 23

. Dorsal outline of left mandible sinuate-lobate (fig. 52, 53) ........... 20

Dorsal outline of left mandible simply arcuate (fig. 51) ............. 22

|| Bossorial elevations 2p on hind tibia shifted (fig.55).....- .@æ. -.......

EE OA el a de . ! [4. syncopator], [6. robustum]

Fossorial elevations 2p on hind tibia opposite (fig. 54) ............. 21

EAD. MENE ee ee ee ee [4. pseudogrande]

EE 2. calanus, (3. phallosum], [9. meridionale]

. Elytral striae superficial, interstriae flat. Pronotal disc anteromedially

bicallose; pronotal derm densely punctate ............... 7. laevicolle

Elytral striae more impressed, interstriae convex ........ [8. pyramidifer]

. Fossorial elevations 2p on hind tibiae shifted. Usually larger (length > 18 mm).

Elytral files distinct. Specimens from Burma and Laos, 13c. d. birmanicum;

OCS AEEA en as 13. drescheri

Fossorial elevations 2p on hind tibiae opposite. Usually smaller (length < 18

mm). Elytral files absent. Apparently restricted to India ...... 12. laterale

d. drescheri

Figs. 21—24. Bolbohamatum drescheri males; dorsal (21) and frontal contours (22—24, pronotum), all

paratypes. 21—22, d. birmanicum, Burma; 23, d. indosinicum, Saigon; 24, d. drescheri, Sukabumi. —

22—24, same scale.

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

cyclops calanus

pseudogrande

meridionale syncopator

Figs. 25—34. Male genitalia of Bolbohamatum (d, dorsal, v, ventral view). 25, cyclops, d, 26, v, apex

only, Dehra Dun; 27, calanus, v, 28, d, Sylhet; 29, phallosum, d, 30, v, holotype; 31, pseudogrande, v.

holotype; 32, meridionale, d, holotype; 33, syncopator, v, 34, d, holotype. — Same scale, except 27.

KRIKKEN: The genus Bolbohamatum

laevicolle robustum pyramidifer

kuijteni

Figs. 35—41. Male genitalia of Bolbohamatum (35—40, ventral, 41, dorsal view). 35, /aevicolle,

Bombay; 36, robustum, holotype; 37, pyramidifer, holotype; 38, laterale, Nagpur; 39, kuijteni, holotype;

40, marginale, holotype; 41, drescheri drescheri, Bandung. — 35—37, 41, same scale; 38—40, same scale.

Figs. 42—44. Ventral paramerites (left) enlarged, full-face. 42, kuijteni; 43, marginale; 44, laterale. All

same scale.

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

| se

= n

deo

Figs. 45—50. Ventral paramerite (left) enlarged, full-face. 45, robustum; 46, pyramidifer; 47, laevicolle;

48, cyclops; 49, pseudogrande; 50, syncopator. All same scale.

Figs. 51—53. Contours of left mandible. 51, /aevicolle, Bombay; 52, robustum, holotype; 53, kuijteni,

holotype. Alle same scale.

Figs. 54—55. Distal-external section of middle tibia to show disposition of protrusions 2p. 54,

pseudogrande, holotype, with situation 2po = paired protrusions (no. 2) opposite; 55, robustum,

holotype, with situation 2ps = paired protrusions shifted. le = complete anteapical crest (no. 1). See

under Technical remarks.

Fig. 56. Metasternal disc and hook-bearing anterior lobe, ventrolateral view, of Bolbohamatum

drescheri drescheri holotype. Fig. 57. Antennal club, view of club segment 1, with some proximal

(flagellar) segments (B. pseudogrande).

KRIKKEN: The genus Bolbohamatum 19

The cyclops group

1. Bolbohamatum cyclops (Olivier) comb. nov.

(figs. 9, 25, 26, 48, 58, 59)

Scarabaeus cyclops Olivier, 1789: 60 (diagnosis, no type-loc. given), pl. 15 fig. 140 (habitus). Westwood,

1848a: 384 (Bolboceras, diagnosis, records) '); 1852: 19 (diagnosis, records), pl. 3 fig. 15 (habitus).

Scarabaeus veter Fabricius, 1792: 33 (diagnosis, type-loc. East India). Boucomont, 1912: 14 (Bolboceras,

in catalogue). Syn. nov.

Bolboceras subglobosus Westwood, 1852 (diagnosis, type-loc. East India), pl. 4 fig. 4 (fore-body). Syn.

nov.

Figs. 58—63. Bolbohamatum males, except 59, female. 58, cyclops, Nepal, 18.5 mm; 59, cyclops, Pusa,

15 mm; 60, calanus, Sylhet, 16 mm; 61, phallosum, holotype, 18 mm; 62, syncopator, holotype, 19.5 mm;

63, pseudogrande, holotype, 20.5 mm.

1) The text of Westwood, 1848b, is identical.

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Identification. — Bolbohamatum cyclops males are easily identifiable by the

characteristic median protrusion on the pronotum. The phallus has a unique

structure. In both sexes the pronotum is heavily punctate. For characters of

female, see below.

Note. — The name cyclops has been applied to some of the other species, e.g. to

drescheri drescheri, because allegedly some material came from Java (see labels of

lectotype). It seems very unlikely that B. cyclops occurs on that island, or

elsewhere outside India.

Variation and sexual dimorphism. — Length ¢ 17—19, 9 17—19 mm. The

median protrusion on the pronotum is simply arcuate in smaller males, whereas

the lateral cavities and adjacent denticles may be reduced as well. The density of

the cephalic punctation varies strongly.

Females are characterized by a very distinct, feebly arcuate transverse ridge

between the obsolescent lateral callosities. The entire pronotum, except for a

small area near the base, is crowded with a well-pronounced double punctation.

On these characters I have here synonymized Bolboceras veter (Fabricius) and

Bolboceras subglobosum Westwood, both based on the female sex, with

Bolbohamatum cyclops, the oldest available name for this species.

Material examined. — 22 males, 14 females.

The male lectotype of Scarabaeus Cyclops Olivier, here designated, with labels

reading “Java” (sic), “cyclops”, “type” (in red circle), “Cyclops Fab./Lees Cabt”,

and a recent Oxford type label “Type COL: 511/Bolboceras/cyclops Oliv./Hope

Dept. Oxford”. The female lectotype of Scarabaeus Veter Fabricius, here

designated, bearing a red type label, and a label reading “Ind: orient: Vahl./Mus. S.

& T: L:/Cyclops./Oliv./Geotrupes/veter. F.” (Zoologiske Museum, Copenhagen).

A female from the Hope Dept. of Entomology, apparently the holotype of

Bolboceras subglobosus Westwood. And material from the following localities,

months, collections:

India: Balaghat; Calcutta; Chapra; Dehra Dun; Moghal Sarai; New Delhi;

Pusa; Simla; Sitapur (W. Almora); Suruwal*; N. India; N.W. India; Nepal;

Himalaya. — Java (mislabelled).

Months vii (7 specimens), viii (2), ix (2), x (2).

Collections BM, P, SMT, Institut Royale des Sciences naturelles de Belgique,

Zoologiske Museum (Copenhagen), Bernice P. Bishop Museum.

The calanus group

2. Bolbohamatum calanus (Westwood) comb. nov.

(figs. 10, 27, 28, 60)

Bolboceras calanus Westwood, 1848a: 385 (diagnosis, type-loc. Bombay); 1852: 21 (diagnosis, &, ? 2),

pl. 3 figs. 19 (habitus), 19a (fore-body), 19b (detail), pl. 4 fig. 6 (fore-body, ? 2).

Bolboceras tumidulus Westwood, 1852: 22 (diagnosis, type-loc. Borhendshukur), pl. 4 fig. 7a, b (fore-

body).

Identification. — Only the characters of the phallus distinguish Bolbohamatum

KRIKKEN: The genus Bolbohamatum 21

| calanus from its relative phallosum. The parameres of calanus have dorsally a

moderately sclerotized, relatively narrow, on the whole poorly developed

paramerite; the ventral side of the parameres is devoid of distinct paramerites. For

_ further characters, see under B. phallosum.

Notes. — Other Bolbohamatum species have frequently been identified as

| calanus Westwood because of their similar pronotal ornamentation. I have not

recovered all the specimens on which Paulian (1945: 41) based his calanus records

for Indo-China, but certainly some of these, if not all, pertain to B. drescheri.

Paulian’s description seems based on drescheri, as the paramedian tubercles of the

pronotum are said to be separated by a rounded foveole. The specimen figured by

Paulian (1945: fig. 25) may, however, indeed belong to calanus.

Variation and sexual dimorphism. — Length g 9 14—18 mm.

A large number of females remain which, judged from the non-sexual characters

and size, may belong to calanus and phallosum. Usually their pronotum has a pair of

feebly developed, slightly transverse median tubercles and an indication of lateral

callosities. The pronotum is abundantly punctate, but never densely punctate

throughout. From the females of the laterale group they can be separated by their

transverse pygidial row of stridulatory granules, and from marginale (and kuijteni)

by their different elytral stria 1. All the females belonging to either calanus or

phallosum that could not be associated with males on the basis of their labels are

recorded below as dubious Bolbohamatum specimens.

Material examined. — 40 males, 10 females.

Male lectotype, here designated, from “India/Boys Sale”, also with label

“calanus Hope”, abdomen missing! Another male “type” in the Hope Dept. of

Entomology, Oxford, is labelled ‘‘Madras/Hope’’, ‘‘Calanus/Hope’’, but that

locality was not mentioned by Westwood (1848a: 385); I extracted the phallus of

this male, and found it to agree with my original interpretation of calanus. In the

BM I saw the holotype of Bolboceras tumidulus Westwood, apparently a female of

either calanus or phallosum. Further material from the following localities, months,

collections:

India: Balasore; Bangalore; Belgaum; Burju* (Bengal); Chickaballapura; Chota

Nagpore; Dallongunj* (Bengal); Dehra Dun; Kanara; Madras; Moiyanala*;

Motinala R.; Mughal Sarai; Namrum* (Bihar); Shimoga; Surada; Tranquebar;

Vellore; Assam; Himalaya; Hindustan; Bangla Desh: Sylhet. — Java

(mislabelled!?).

Months v, vii-viii (only 7 specimens had dates).

Collections BH, BM, DEI, P, SMT, Zoologiske Museum (Copenhagen), Forest

Research Institute (Dehra Dun), Hope Dept. of Entomology, Senckenberg-

Museum, Zoological Survey of India (Calcutta).

3. Bolbohamatum phallosum sp.nov.

(figs. 11, 29, 30, 61)

Holotype (male). — Approximate length 18, width 10.5, height 8 mm. Medium-

brown, shiny; pilosity yellow-brown. Habitus, fig. 61.

22 TUDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Labrum emarginate in front, sides rounded; surface anteriorly limited by

transverse ridge. Cephalic contours, fig. 11. Clypeus with pair of dentiform

tubercles; clypeal margin ridged, genae raised; marginal declivities rugulate-

punctate, horizontal surface contiguously punctate to rugulate-punctate;

clypeofrontal suture effaced. Frontal disc concave; posterior cushion indistinct;

entire frontovertex densely, but vaguely punctate. Eye-canthus rugulate-punctate;

frontolateral ridge extending from genae beyond eye.

Pronotal contours, fig. 11; midline of pronotum impressed; base marginate;

surface behind eyes slightly concave; pronotal punctation double (x 25); primary

punctation coarse, dense on midline impression, on lateral declivities and along

marginal ridges, absent on disc at some distance from base, in lateral cavities and

on tubercles; secondary punctation abundant. Scutellum (fig. 11) subopaque, due

to microreticulation, indistinctly punctate.

Juxtasutural punctures of elytra subobsolete; discal striae shallowly impressed,

finely punctate; punctures separated by 5 + 2 times their diameters; peripunctural

impressions ill pronounced, slightly affecting interstriae. Elytral interstriae very

slightly convex; derm subopaque, due to microreticulation (x 50); punctation

minute, sparse.

Phallus figs. 29, 30.

Fore tibia with 2 + 5 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of non-

apical elevations on right middle tibia 1c—2po—3ps—4, on right hind tibia

le—2po/s—3ps—4ps: complete anteapical crest bidentate. Stridulatory granules

on pygidium arranged in transverse row.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 2.6, maximum width 4.4; distance tips of clypeal tubercles 1.5.

Distance anterolateral angles of pronotum 5.0, tips paramedian tubercles 1.2,

lateral tubercles 6.3; median length 5.6, maximum width 9.9. Median length of

scutellum 1.5, maximum width 1.8. Number of primary punctures on pronotal disc

behind paramedian tubercles 6—7/0.25 sq. mm, their diameters 0.10—0.15 mm.

Variation and sexual dimorphism. — Length ¢ 16—18 mm. There is some

variation in the development of cephalic and pronotal protrusions.

Females of this species probably go unrecognized in the material recorded

further below as dubious specimens.

Identification. — Bolbohamatum phallosum and calanus differ from the other

species by the following combination of characters. Males with closely

approximated paramedian tubercles on the pronotum, lateral protrusions not

shifted to anterolateral corner; clypeal tubercles dentiform. Elytra shiny.

Protrusions 2p on hind tibia more or less opposite to each other. Stridulatory

granules on pygidium arranged in transverse row. B. calanus and phallosum cannot

be separated on external characters, but the phalli of both species are very

different. Dorsally the parameres of phallosum are foliate: ventrally there is a pair

of more or less glider-like paramerites; the basal capsule is in lateral view distally

strongly emarginate.

Material examined. — 9 males.

KRIKKEN: The genus Bolbohamatum 23

Holotype from India: Bombay (Institut Royal des Sciences naturelles de

Belgique, Brussels). Paratypes as follows:

India: Belgaum (1 3, BM); Bombay (1 g, P); ditto, Fontanier (2 g, P); East

India (1 g, BM); East India no “962” (1 g, Senckenberg-Museum); India (1 &,

BM). No data (1 &,P).

The pseudogrande group

4. Bolbohamatum pseudogrande sp. nov.

(figs. 12, 31, 49, 54, 57, 63)

Holotype (male). — Approximate length 20.5, width 13.5, height 9 mm.

Medium-brown, moderately shiny; pilosity yellow-brown. Habitus, fig. 63.

Labrum emarginate in front, sides rounded; surface anteriorly limited by

arcuate ridge. Dorsal outline of left mandible lobate. Cephalic contours, fig. 12.

Clypeus with pair of clypeal tubercles more or less connected at base; clypeal

margin ridged, genae raised; marginal declivities steep, rugulate-punctate;

horizontal surface rugulate-punctate; clypeofrontal suture effaced. Frons with

transversely confluent punctures in front, irregularly punctate behind; posterior

cushion ill pronounced. Eye-canthus rugulate-punctate; frontolateral ridge

strongly pronounced, slightly curved outwards caudad, extending beyond eye.

Pronotal contours, fig. 12. Midline of pronotum slightly impressed; base

marginate; pronotum with closely set subtransverse paramedian tubercles, on

each side separated from high lateral tooth by deep cavity; anterolateral corner

distinctly concave. Pronotal derm with distinct double punctation; primary

punctation coarse, crowded laterally, contiguous along margin, sparse or absent in

anterolateral corner, finer, crowded on anterior declivity just in front of lateral

cavities, secondary punctation (x 12) abundant, evenly distributed. Scutellum (fig.

12) vaguely, crowdedly punctate.

Juxtasutural punctures obsolete (x 50); discal striae shallowly impressed, finely

punctate; punctures separated by 6 + 2 times their diameters; peripunctural

impressions ill pronounced, slightly affecting interstriae. Elytral interstriae

scarcely convex, sparsely, vaguely, minutely punctate (x 50) and microreticulate

(x 50).

Phallus, fig. 31.

Fore tibia with 2 + 6 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to (pairs of) denticles; number of

elevations on right middle tibia Ic—2ps—3ps—4—5, on right hind tibia

1c—2po—3ps—4ps; complete anteapical crests bidentate, with moderately long

setae. Stridulatory granules on pygidium arranged in transverse row.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.5, maximum width 5.3; distance tips clypeal tubercles 1.4. Distance

anterolateral angles of pronotum 5.8, tips of paramedian tubercles 1.2, tips of

lateral teeth 9.1; median length 7.0, maximum width 12.0. Median length of

scutellum 1.7, maximum width 2.3. Number of primary punctures on median

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1. 1980

protrusion of pronotum 15—20/sq. mm, their diameters 0.08 mm and less.

Female sex unknown.

Variation. — The paratype male is 21.5 mm long.

Identification. — Bolbohamatum pseudogrande is a large species, very similar to

syncopator, phallosum, calanus, and robustum. Especially robustum can easily be

confounded with pseudogrande, but the males of the latter have the clypeal

tubercles somewhat connected at their base, whereas the protrusions 2p on the

hind tibia are placed opposite to each other. Further external features

characterizing B. pseudogrande include: Scutellum punctate. Lateral protrusions of

pronotum high, not situated marginally; pronotal cavities very deep. Shape of

ventral paramerites characteristic. This diagnostic information is rather tentative

considering the fact that only two males are at hand.

Materal examined. — Holotype from India: Kangra Valley, vii.1899, Dudgeon,

4500 ft (Howden collection). Male paratype from Assam (BM).

5. Bolbohamatum syncopator sp. nov.

(figs. 14, 33, 34, 62)

Holotype (male). — Approximate length 19.5, width 12.5, height 9.5 mm.

Medium brown, moderately shiny; pilosity yellow-brown. Habitus, fig. 62.

Labrum distinctly emarginate in front, sides rounded; surface anteriorly limited

by arcuate ridge. Dorsal outline of left mandible lobate. Cephalic contours, fig. 14.

Clypeus with pair of dentiform tubercles; clypeal margin ridged, genae raised;

marginal declivities low, rugulate-punctate; horizontal surface rugulate-punctate;

clypeofrontal suture effaced. Frontal disc slightly depressed, posteriorly limited by

pair of transverse callosities; derm densely punctate. Eye-canthus rugulate-

punctate; frontolateral ridge strongly pronounced, slightly curved outwards

caudad, extending beyond eye.

Pronotal contours, fig. 14. Midline of pronotum slightly impressed; base

marginate; pronotum with paramedian and lateral tubercles, surface between

paramedian and lateral tubercles, shallowly impressed; anterolateral corner

shallowly impressed. Pronotal derm with distinct double punctation; primary

punctures very dense on impressed midline between paramedian and lateral

tubercles; rugulate-punctate on lateral declivities and along marginal ridge;

secondary punctation (x 12) very abundant, evenly distributed. Scutellum (fig. 14)

vaguely rugulate-punctate.

Juxtasutural punctures of elytra obsolete (x 50); discal striae shallowly

impressed, finely punctate; punctures separated by 8 + 3 times their diameters;

peripunctural impressions ill pronounced, scarcely affecting interstriae. Elytral

interstriae scarcely convex, densely minutely punctate (x 25) and microreticulate

(x 50).

Phallus, figs. 33, 34.

Fore tibia with 2 + 6 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of elevations

on right middle tibia 1c—2ps—3ps—4ps, on right hind tibia lc—2ps—3ps—4ps;

KRIKKEN: The genus Bolbohamatum 25

complete anteapical crests bidentate, with setae of varying length. Stridulatory

granules on pygidium arranged in transverse row.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 2.9, maximum width 5.0, distance tips of clypeal tubercles 2.0.

Distance anterolateral angles of pronotum 6.0, tips of paramedian tubercles 1.3,

lateral teeth 7.3; median length 6.6, maximum width 10.9. Median length of

scutellum 1.6, maximum width 2.2. Number of primary punctures behind pronotal

impressions 5—10/0.25 sq. mm, their diameters ca. 0.06 mm.

Female sex unknown.

Identification. — The male of Bolbohamatum syncopator looks very much like

those of pseudogrande, phallosum, calanus, and robustum, all these having a more or

less similar cephalic and pronotal ornamentation. B. pseudogrande differs by

having the clypeal teeth more or less connected at base, and by the protrusions 2p

on the hind tibia being placed opposite to each other. Two of the other species

mentioned above have similar protrusions 2p, only robustum having them distinctly

shifted. As the only reliable character to separate syncopator from robustum, |

would suggest the structure of the phallus.

Material examined. — Holotype only, from Bangla Desh: Sylhet (SMT).

The /aevicolle group

6. Bolbohamatum robustum sp. nov.

(figs. 13, 36, 45, 52, 55, 64)

Holotype (male). — Approximate length 21.5, width 14, height 9.5 mm. Light-

brown, subopaque; pilosity yellow-brown. Habitus, fig. 64.

Labrum emarginate in front, sides rounded; surface anteriorly limited by fine

arcuate ridge, dorsal outline left mandible lobate. Cephalic contours, fig. 13.

Clypeus with pair of dentiform tubercles; clypeal margin ridged, genae raised;

marginal declivities rugulate-punctate; horizontal surface contiguously punctate

to rugulate-punctate; clypeofrontal suture effaced. Frontal disc slightly depressed;

posterior cushion indistinct, densely punctate; remainder of frons also abundantly

punctate. Eye-canthus rugulate-punctate; frontolateral ridge extending from

genae beyond eye.

Pronotal contours, fig. 13. Midline of pronotum impressed; base marginate;

surface behind anterolateral angles concave; lateral horns upright; pronotai

punctation double (x 25); primary punctation dense, very dense laterally and

behind vertex; secondary punctation very distinct behind lateral cavity; disc at

some distance from base sparsely, indistinctly punctate, shiny. Scutellum (fig. 13)

superficially, indistinctly punctate.

Juxtasutural punctures of elytra subobsolete; discal striae shallowly impressed,

finely punctate; punctures separated by 6 + 2 times their diameters; peripunctural

impressions ill pronounced, not affecting interstriae. Elytral interstriae flat,

distinctly microreticulate (x 50), abundant minute punctation almost indistinct.

Phallus, fig. 36.

26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Figs. 64— 71. Bolbohamatum males. 64, robustum, holotype, 21.5 mm; 65, laevicolle, Bengal, 19.5 mm;

66, pyramidifer, holotype, 19 mm; 67, meridionale, holotype, 18 mm; 68, marginale, holotype, 16 mm; 69,

kuijteni, holotype, 17 mm; 70, /aterale, Nagpur, 16.5 mm; 71, drescheri indosinicum, holotype, 18 mm.

KRIKKEN: The genus Bolbohamatum 27

| Fore tibia with 2 + 6 external denticles. Middle and hind tibiae with setose

| fossorial elevations, proximally reduced to pairs of denticles; number of non-

apical elevations on right middle tibia Ic—2ps—3ps—4ps, on right hind tibia

| 1c—2ps—3ps—4ps; complete anteapical crest bidentate, with ca. 10 long setae.

| Stridulatory granules on pygidium arranged in transverse row.

| Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.5, maximum width 5.3; distance tips of clypeal tubercles 1.5.

Distance anterolateral angles of pronotum 5.9, tips paramedian tubercles 1.3,

lateral tubercles 9.1; median length 7.6, maximum width 11.6. Median length of

scutellum 1.8, width 2.2. Number of primary punctures on pronotal disc behind

paramedian tubercles 6—10/0.25 sq. mm; punctures slightly elongate, ca. 0.10 x

0.08 mm.

Female sex unknown.

Identification. — Bolbohamatum robustum is externally very similar to

pseudogrande, syncopator, phallosum, and calanus, and therefore, with this single

male at hand, I am reluctant to discuss any slight external differences. The crossed

paramerites of robustum, however, are most distinct, whereas the other characters

mentioned in the synoptic table and the key readily distinguish between the three

species of the laevicolle group.

Material examined. — Holotype only, from the ‘“Himalaja” (Zoologische

Staatssammlung, Munich).

| 7. Bolbohamatum laevicolle (Westwood) comb. nov.

| (figs. 15, 35, 47, 51, 65)

| Bolboceras laevicollis Westwood, 1848a: 385 (diagnosis, type-loc. East India); 1852: 22 (diagnosis), pl. 4

| fig. 8 (fore-body).

| Bolboceras grandis Westwood, 1848a: 384 (diagnosis, type-loc. East India?); 1852: 19 (diagnosis), pl. 4

fig. 3 (fore-body). Syn. nov.

‘Identification. — Externally the males of Bolbohamatum laevicolle are easily

| identifiable by the three usually well-defined, deep cavities separating the four

| pronotal tubercles. Superficially similar is B. drescheri, but its subspecies range

i from Burma to Java, whereas the pronotal cavities are less deep and less defined.

B. robustum males, and those of some species outside the /aevicolle group, are

different by having their paramedian pronotal tubercles closely approximated.

The females of /aevicolle have a transverse ridge on the pronotum (see next

section).

Variation and sexual dimorphism. — Length ¢ 19—20 mm, 9 20—21 mm. The

males at hand are scarcely variant.

On the pronotal disc of the females there is a pair of lateral callosities separated

by a distinct, feebly arcuate transverse ridge. Pronotum with dense to crowded

primary punctatioh, except on basal part of disc.

Because one of the characters distinguishing /aevicolle (males and females) is the

simply arcuate outline of the left mandible, it appears most likely that Bolboceras

grande Westwood, the type of which agrees in this and all other characters, is the

female of laevicolle, and consequently they are synonymized here.

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Material examined. — 6 males, 6 females.

The male holotype of Bolboceras laevicollis Westwood, which has its wings

spread (Hope Dept. of Entomology, Oxford). The female holotype of Bolboceras

grandis Westwood (same museum). And the following specimens:

India: Bombay (2 4, 1 9, BM, I ©, BH); Himalaya (1 g, BH); Kotapad (1 3,

BH); Nagpur (1 ©, BH); Assam (locality illegible) (1 9, SMT); Bengal (1 &,

SMT); no details (1 ©, BH).

8. Bolbohamatum pyramidifer sp. nov.

(figs. 16, 37, 46, 66)

Holotype (male). — Approximate length 19, width 12, height 9 mm. Reddish

yellow-brown, shiny; pilosity yellowish. Habitus, fig. 66.

Labrum emarginate in front, sides rounded; surface anteriorly limited by fine

transverse ridge; dorsal outline of left mandible arcuate. Cephalic contours, fig.

16. Clypeus rugulate-punctate, punctures superficial; margin entirely ridged,

genae raised; declivities with sculpture similar to that of horizontal surface;

clypeofrontal suture effaced. Frons with pair of pyramidiform tubercles between

eye-canthi; posterior cushion and impression poorly pronounced, derm microre-

ticulate (x 50). Eye-canthus coarsely rugulate-punctate; frontolateral ridge

straight, extending from gena beyond eye.

Pronotal contours, fig. 16. Midline om scarcely impressed; top of disc

deplanate, anteriorly limited by arcuate ridge; lateral protrusions represented by

simple callosities; base marginate; pronotal punctation double (x 25); primary

punctures densely set laterally, sparse on subbasal part of disc and on lateral

callosities, elsewhere abundant. Scutellum (fig. 16) irregularly densely punctate.

Juxtasutural punctures of elytra subobsolete; discal striae shallowly impressed,

finely punctate; punctures separated by 7 + 3 times their diameters; peripunctural

impressions ill pronounced, slightly affecting interstriae. Elytral interstriae

scarcely convex; with scarcely distinct, sparse punctures, as well as micro-

reticulation (x 75).

Phallus, fig. 37.

Fore tibia with 2 + 5 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of non-

apical elevations in right middle tibia 1c—2po—3po—4, on right hind tibia

Ic—2po—3po; complete anteapical crests bidentate, with setae of varying length.

Stridulatory granules on pygidium arranged in transverse row.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.1, maximum width 4.5; distance tips of clypeal tubercles 1.6.

Distance anterolateral angles of pronotum 5.5; median length 6.3; maximum width

10.5. Median length of scutellum 1.6, width 2.3. Number of punctures of pronotal

disc 12—17/sq. mm, their diameters ca. 0.1 mm.

Female sex unknown.

Identification. — Bolbohamatum pyramidifer is externally rather similar to

meridionale because of the pyramidiform cephalic tubercles and the apparently

KRIKKEN: The genus Bolbohamatum 29

reduced pronotal ornamentation. B. meridionale, however, has its cephalic

tubercles connected by a distinct, infuscated straight ridge; the same applies to the

paramedian tubercles on its pronotum. The outer margin of the left mandible is in

pyramidifer simply arcuate; the secondary punctures of the pronotal disc near the

base are ocellate. The crossed paramerites place pyramidifer immediately in the

laevicolle group.

Material examined. — Holotype only, from “Ind. or./Kotapad” (BH).

The meridionale group

9. Bolbohamatum meridionale sp. nov.

(figs. 17, 37, 46, 67)

Holotype (male). — Approximate length 18, width 12, height 8 mm. Yellow-

brown, shiny; pilosity yellow-brown. Habitus, fig. 67.

Labrum emarginate in front, sides rounded; surface anteriorly limited by

distinct, slightly curved ridge. Dorsal outline of left mandible lobate. Cephalic

contours, fig. 17. Clypeus short; between eye-canthi a pair of pyramidiform

tubercles distinctly connected by transverse crest; clypeal margin ridged, genae

scarcely raised; clypeofrontal suture effaced; clypeal surface, including declivities,

finely rugulate-punctate. Frons and vertex with apparent double punctation

because some of the punctures (x 18) have an ill-defined peripunctural

impression; frontal cushion and impression poorly pronounced. Eye-canthus

coarsely rugulate-punctate; frontolateral ridge straight, extending from gena

beyond eye.

Pronotal contours, fig. 17; midline of pronotum impressed; base marginate;

discal protrusions small, paramedians connected by feebly, more or less infuscated

ridge; pronotal punctation double ( x 25); primary punctation abundant, except on

basal part of disc, somewhat confluent on shallow impression between paramedian

and lateral tubercles; secondary punctation abundant. Scutellum (fig. 17)

irregularly punctate, slightly wrinkled.

Juxtasutural punctures of elytra subobsolete (x 50); discal striae distinctly

impressed, distinctly punctate; punctures fine, infuscated, separated by 8 + 3

times their diameters, lacking distinct peripunctural impressions. Elytral

interstriae slightly convex, abundantly minutely punctate and microreticulate (x

50).

Phallus, fig. 37.

Fore tibia with 2 + 5 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of elevations

on right middle tibia 1c—2ps—3ps, on right hind tibia 1c—2ps—3ps; complete

anteapical crests bidentate, with setae of varying length. Stridulatory granules on

pygidium arranged in transverse row.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.1, maximum width 4.8; distance terminal tips of transverse ridge 1.6.

Distance anterolateral angles of pronotum 5.5, tips of paramedian protrusions 5.3;

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

maximum length 6.5, maximum width 10.7. Median length of scutellum 1.6,

maximum width 2.2. Total number of primary punctures on pronotum behind

paramedian protrusions 70—80, their diameters ca. 0.07 mm.

Female unknown.

Variation. — Not significant in the three males available.

Identification. — Males of Bolbohamatum meridionale are easily recognized by

their cephalic and pronotal protrusions. B. pyramidifer looks like a poorly

developed meridionale, but in case of doubt the characteristics of the parameres \

are decisive, those of meridionale lacking the strongly sclerotized, crossed |

paramerites. The cephalic pyramids are differently oriented in the two species. —

B. pyramidifer has a simply arcuate outline of the left mandible, which in«

meridionale is sinuate-lobate. The protrusions 2p on the hind tibia are more or less

opposite in pyramidifer, shifted in meridionale.

Material examined. — Holotype from Sri Lanka: Colombo (National Museum,

Colombo). Two male paratypes, one from Sri Lanka: Batticaloa, vi.1915, Baker

(BM), the other from India: Pondicherry, Martin (P).

This is the only species known from Ceylon.

The laterale group

10. Bolbohamatum marginale sp. nov.

(figs. 20, 40, 43, 68)

Holotype (male). — Approximate length 16, width 10.5, height 8 mm. Medium-

brown, shiny; pilosity yellow-brown. Habitus, fig. 68.

Labrum emarginate in front, sides rounded; surface anteriorly limited by

arcuate ridge. Dorsal outline of left mandible lobate. Cephalic contours, fig. 20.

Clypeus with pair of dentiform tubercles, each placed against lateral margin;

clypeal margin weakly ridged, genae raised; marginal declivities rugulate-

punctate; horizontal surface irregularly contiguously punctate; clypeofrontal

suture effaced. Frontal disc slightly depressed; posterior cushion distinct, very

sparsely punctate; remainder of frons with variably abundant punctures of two size

classes. Eye-canthus irregularly contiguously punctate; frontolateral ridge

contiguous with inner border of eye, extending beyond eye, adjacent frontal

surface impressed.

Pronotal contours, fig. 20. Midline of pronotum slightly impressed; base

submarginate, medially virtually emarginate; pronotum with paramedian and

lateral tubercles. Median longitudinal zone and lateral declivities of pronotum

densely, coarsely punctate; secondary punctation (x 25) evenly distributed,

abundant; impression between paramedian and lateral tubercles virtually devoid

of punctures (secondary punctation vague), opaque, due to microreticulation (x

50). Scutellum (fig. 20) virtually impunctate.

Juxtasutural punctures of elytra obsolete (x 50); discal striae shallowly

impressed, finely punctate; punctures separated by 7 + 2 times their diameters:

diameters of punctures in stria | strongly increasing on distal declivity, to 0.1 mm,

KRIKKEN: The genus Bolbohamatum 31

| separated by 2—3 times their diameters; these punctures deep, well defined,

distinctly affecting adjacent interstriae. Elytral interstriae scarcely convex, except

distal section of 1, which is strongly convex; interstriae shiny, sparsely

micropunctate (x 50).

Phallus, fig. 40.

Fore tibia with 2 + 5 or 6 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of non-

apical elevations on right middle tibia Ic—2ps—3, on right hind tibia

lc—2ps—3ps; complete anteapical crest bidentate, with setae of varying length.

Stridulatory granules on pygidium scattered.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.0, maximum width 4.3; distance tips of clypeal tubercles 1.6.

Distance anterolateral angles of pronotum, tips paramedian tubercles 1.3, tips

lateral tubercles 7.2; median length 6.2, maximum width 9.6. Median length of

scutellum 1.3, maximum width 1.7. Number of punctures on anterior declivity of

pronotum 35—40, their diameters ca. 0.1 mm.

Variation and sexual dimorphism. — Length & 14—17, 9 14.5—17 mm.

Ornamention of head and pronotum of smaller males more or less reduced.

A male from Mahé has its paramedian pronotal tubercles wider apart than the

other specimens. A male from Wallardi is rather different from all the others,

particularly because the small clypeal teeth stand exactly between the genal

angles; the pronotum of this male is anteromedially feebly bicallose, whereas the

first elytral stria is distally only slightly impressed and scarcely more strongly

punctate. Length of that specimen ca. 15.5 mm.

Identification. — The males of Bolbohamatum marginale have strongly

approximated paramedian tubercles on the pronotum, and a pair of lateral

tubercles situated almost marginally. By the latter character they are easily

separable from other species with approximated paramedian tubercles; other

species with the lateral protrusions placed far apart lack the strongly impressed,

more coarsely punctate distal section of elytral stria 1. The slight differences in this

character with kuijteni, possibly important for the recognition of individual

females, have to be verified on more material. The basal capsule of the phallus of

marginale is very robust compared to those of laterale and kuijteni, and reminds

one of the drescheri group treated hereafter. The elytral striae 2 etc. of both

marginale and kuijteni are very feebly punctate.

Material examined. — 14 males, 9 females.

Holotype from India: Baihar [sic!]: Balaghat, 21.vii.1927, B.M. Bhatia (BM).

Paratypes as follows:

India: Boria (Jubbulpore), 30.vi.1934, Chatterjee (1 &, Forest Res. Inst. and

Colleges, Dehra Dun); Kodama Hills*, 5.[?].1934 (1 ¢, BH); Mahé (1 g, P);

Mercara (1 &, 1 ©, BH); Motinala River, 13.vi.1927, Chatterjee (1 3, BM); Nilgiri

Hills, Downing (1 &, BM); ditto, Andrewes (4 3, 1 ©, BM); Surada, Babault (1 &,

1 ©, P); Travancore: Wallardi*, 5.ix.1903, Faire (1 4, excluded from type-series,

P); South Mysore (1 3, BM); “India/Orient” (1 ¢, BM).

Six females not associated with males excluded from type-series, from India:

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Anamalais (1 9, BM); Belgaum, 29.vii.1906, Bell (2 9, BM); Mountabu, 1940,

McCann (1 9, Bombay Natural History Society). Pakistan: Karachi, Bell (2 9,

BM). Not mapped.

11. Bolbohamatum kuijteni sp. nov.

(figs. 19, 39, 42, 53, 69)

Holotype (male). — Approximate length 17, width 10.5, height 8 mm. Yellow-

brown, shiny; pilosity yellow-brown. Habitus, fig. 69.

Labrum emarginate in front, sides rounded; surface anteriorly limited by fine

arcuate ridge. Dorsal outline of left mandible lobate. Cephalic contours, fig. 19.

Clypeus bituberculate, tubercles high, posteriorly with 2 vague carinulae running

from tip to base; tubercles contiguous with clypeal margin; clypeal margin ridged,

genae raised; marginal declivities rugulate-punctate; clypeofrontal suture effaced.

Clypeofrons with double punctation (x 25); primary punctation abundant, fine,

density decreasing caudad; vertex with similar punctation, but primary punctures

larger and restricted to lateral parts; posterior cushion arcuate, more or less

costiform. Eye-canthus coarsely, irregularly punctate; frontolateral ridge

extending from gena beyond eye.

Pronotal contours, fig. 19. Midline of pronotum impressed; transition from disc

to anterior declivity gradual; base submarginate, medially virtually emarginate.

Pronotal declivity in front of paramedian tubercles abundantly, distinctly

punctate, punctures separated by one to several times their diameters; punctation

sparse elsewhere; secondary punctation abundant, but scarcely distinct (x 50).

Scutellum (fig. 19) virtually impunctate.

Juxtasutural punctures of elytra subobsolete; discal striae shallowly impressed,

finely punctate; punctures separated by 10 + 5 times their diameters; diameters of

punctures in stria | strongly increasing on distal declivity to 0.15 mm, separated by

2—3 times their diameters; these punctures well defined, deep, distinctly affecting

adjacent interstriae. Elytral interstriae scarcely convex, except distal section of 1,

which is strongly convex; interstriae microreticulate (x 75), punctures scarcely

distinct.

Phallus, fig. 39.

Fore tibia with 2 + 6 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of non-apical

elevations on right middle tibia 1c—2ps—3, on right hind tibia Ic—2po—3po;

complete anteapical crest bidentate, with setae of varying length. Stridulatory

granules on pygidium scattered.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.0, maximum width 4.2, distance tips of clypeal tubercles 1.7.

Distance anterolateral angles of pronotum 4.8, tips paramedian tubercles 4.7, tips

lateral tubercles 7.3; median length 6.3, maximum width 9.30. Median length of

scutellum 1.2, maximum width 1.9. Number of punctures on anterior declivity of

pronotum ca. 15/sq. mm, their diameters 0.07—0.10 mm.

Female sex unknown.

KRIKKEN: The genus Bolbohamatum 33

Variation. — Length & 14.5—17 mm. Cephalic and pronotal ornamentation

somewhat reduced in the small specimens.

Identification. — Bolbohamatum kuijteni males may be confounded with those of

drescheri birmanicum, which also have their paramedian pronotal tubercles wide

apart. The latter, as well as B. laterale, however, has a much denser pronotal

punctation. Compared to most other species kuijteni is very shiny dorsally. With

B. marginale the species agrees in having a strongly impressed, coarsely punctate

distal section of elytral stria 1, but the pronotal ornamentation of the males is very

different. The aforesaid impression and punctation of stria | seems stronger in

kuijteni than in marginale, a character that could be important for the recognition

of individual females. The basal capsule of the phallus of kuijteni is narrow, rather

slender compared to that of marginale, whereas the paramerites have a

characteristic shape.

Material examined. — 8 males.

Holotype from India: Gersappa, 19.vi.1907 (Bombay Natural History Society,

Bombay). Paratypes as follows:

India: Gersappa, 28.vi.1907 (1 g, Bombay Natural History Society, Bombay);

ditto, no date (1 ¢, BM, | &, Bombay); Igatpuri, 200 ft (1 3, BM); Kanara (1 g,

BM, 1 g, Zoologiske Museum, Copenhagen). No data (1 G, Senckenberg-

Museum, Frankfurt).

Note. — This species is dedicated to my friend P. J. Kuijten who, many years

ago, introduced me to the fascinating world of scarabs.

12. Bolbohamatum laterale (Westwood) comb. nov.

(figs. 18, 38, 44, 70)

Bolboceras lateralis Westwood, 1848a: 385 (diagnosis, type-loc. Gogo); 1852: 22 (diagnosis), pl. 4 fig. 10

(fore-body).

Identification. — Bolbohamatum laterale males are very distinct by their

pronotal protrusions and by the heavy double punctation of virtually the entire

pronotum. The two other species in the laterale group have two pairs of pronotal

protrusions instead of one, while their first elytral stria is distally more strongly

impressed and more coarsely punctate than proximally. Female sex not known for

certain, but four females with a heavily punctate pronotum, lacking the transverse

row of pygidial granules, and with a medially dentate frontal ridge, may indeed

belong to laterale.

Variation. — Length g 17—18 mm. Further characters, e.g. anterolateral

impressions of pronotum, slightly variant.

Material examined. — 5 males, 4 females.

The male holotype, which, in addition to “lateralis Westw.”, bears a label

reading “tuberculatus/Hope Gogo”*, plus the usual labels of the Hope Dept. of

Entomology, Oxford. Further specimens as follows:

India: Belgaum, at light (1 &, BM); Darjeeling (2 9, P); Mhow (1 9, BM);

Nagpur, 31.vii.1916, d’Abreu, 1000 ft (1 g, BM); Sagoda (Purna R.), 3.ix.1929,

Chatterjee (1 ©, BM); Kashmir, Hügel (1 g, SMT).

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

The drescheri group

13. Bolbohamatum drescheri sp. nov.

(figs. 1, 21—24, 41, 56, 71)

Holotype (male). — Approximate length 18.5, width 12.5, height 8 mm. Yellow-

brown; subopaque, pronotal disc shiny; tips ridges, margins, sutures more or less

infuscated; pilosity yellowish. Habitus, fig. 1.

Labrum emarginate in front, sides rounded; surface with fine arcuate ridge.

Dorsal outline of left mandible lobate. Cephalic contours, fig. 1. Clypeus

bidentate; surface rugulate-punctate, punctures superficial; margin entirely

ridged, genae raised; marginal declivities high, sculpture similar to that of

horizontal surface; clypeofrontal suture indistinct. Frons abundantly, irregularly

punctate. Vertex with transverse cushion, posteriorly limited by transverse

impression. Eye-canthus rugulate-punctate; frontolateral ridge extending from

genae beyond eye.

Pronotal contours, fig. 1. Midline of pronotum impressed; base submarginate;

pronotal punctation double (x 25); primary punctures densely set, except in lat-

eral cavities and on disc at some distance along base (where primary punctures are

most distinct). Scutellum (fig. 1) irregularly densely punctate.

Juxtasutural punctures of elytra subobsolete; discal striae shallowly impressed,

finely punctate; punctures separated by 10 + 5 times their diameters;

peripunctural impressions ill pronounced, scarcely affecting interstriae. Elytral

interstriae scarcely convex, sparsely, minutely punctate as well as microreticulate

(x 75); punctation simple, punctures separated by more than 5 times their

diameter.

Phallus, as paratype, fig. 41.

Fore tibia with 2 + 6 or 7 external denticles. Middle and hind tibiae with setose

fossorial elevations, proximally reduced to pairs of denticles; number of non-

apical elevations on right middle tibia lc—2ps—3, on right hind tibia

lc—2ps—3ps—4; complete anteapical crest bidentate, with setae of varying

length. Stridulatory granules on pygidium scattered; elytral files present.

Some measurements in mm. Cephalic maximum length (exclusive of labrum and

mandibles) 3.1, maximum width 5.0; distance tips of clypeal tubercles 1.5. Dis-

tance anterolateral angles of pronotum 5.8, tips paramedian tubercles 2.4, lateral

tubercles 7.2; median length 6.6, maximum width 11.3. Median length of scutellum

1.5, width 2.0. Number of punctures between paramedian tubercles on pronotum

6—10/0.25 sq. mm, their diameters ca. 0.1 mm.

Variation and subspecies. — Within Bolbohamatum drescheri there is a notable

geographic variation in the distance of the paramedian pronotal protrusions and

the direction of the axes of these (figs. 22—24). The populations of Java are re-

markably constant in these male characters, the distance of the tubercular tips

being always less than half the distance between the anterolateral angles of the

pronotum, whereas the axes of the cones are usually inclined to the pronotal

midline. The Burmese specimens always have their pronotal cones much wider

apart, up to slightly over 0.75 of the distance between the anterolateral angles,

| KRIKKEN: The genus Bolbohamatum 35

whereas the cones stand outward or upright. The Indo-Chinese and Chinese

populations seem more variable, but the paramedian tubercles of the majority of

‘the males from Vietnam and Cambodia appear to have the same distance as in

those of Java; the axes of the cones, however, are not inclined. The larger males

from Burma and Indo-China have a distinct transverse costa between the

paramedian pronotal tubercles.

On the basis of the characters mentioned, the populations of Java, Indo-China

‘and Burma are here considered to belong to three different subspecies,

‘d. drescheri, d. indosinicum and d. birmanicum. The few Chinese specimens could

‚not be placed. One male and three females from Laos (3 localities) are assigned to

| birmanicum'). In addition to this geographic variation the usual reduction of the

i pronotal ornamentation is observed in smaller males. I have one doubtful male

from Palone (Burma), only with very feeble paramedian pronotal protrusions,

| placed under drescheri because of its stridulation apparatus and its phallus. This

male is recorded under d. birmanicum.

Total length 15.5—21 mm.

Identification. — Bolbohamatum drescheri males have a most characteristic

phallus, its basal capsule being very large compared to the aedeagus. The lateral

tubercles on the pronotum stand very wide apart, their distance exceeding that of

the anterolateral angles (see measurements of holotype of d. drescheri).

B. marginale and kuijteni are similar in this character, but differ in that their clypeal

tubercles are situated directly against the marginal ridge. Contrary to drescheri, is

the elytral stria 1 in marginale and kuijteni distally strongly impressed and strongly

punctate. There are no records of drescheri from localities west of Burma.

| Material examined. — Not allocated to subspecies mentioned below: 3 males, 2

| females, all paratypes, as follows:

China: Hongkong: Taipokan, 10.vi.1964, Voss & Wai Ming Hui (1 ©, Bishop

Museum, Honolulu); H. Kung [? Hongkong] (1 3, BM); Tonglok [? North of

Hongkong] (1 &, BH); Hupe: Ichang (1 9, BH; northernmost locality, province

-correct?); China (1 g, BM).

The above specimen description pertains to the holotype of the nominate

subspecies, mentioned hereafter.

Note. — This species is named after F. C. Drescher (1875—1957), ardent

collector of Indonesian beetles.

13a. Bolbohamatum drescheri subsp. drescheri nov.

(figs. 1, 24, 41)

Material examined. — 32 males, 31 females.

Holotype & from Java: Tjideres, 24.xii.1935, F. C. Drescher, 100 m (Museum

Zoologicum Bogoriense). Paratypes as follows:

!) In principle a good series representing a local population is needed to establish the subspecific iden-

tity of individuals (see e.g. Mayr, 1963: point 3 on p. 349), but this is scarcely ever practicable due to

the usual paucity of material in taxonomic collections. The features used here for the delimitation of

subspecies seem constant enough to justify the present treatment.

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 1, 1980

Java: Ardja Sari (1 &, 2 9, L), ditto, 1923 (1 9, L); Bandung, 23.vi.1930,)

Drescher, at light (1 &, P), ditto, 12.vi.1936, Drescher, 750 m (1 &, Bogor);

Bantam, L. de Vos (1 &, 3 9, L); Bantam (1 g, 1 9, L); Batavia [= Jakarta], De:

Gavere (1 9, L), ditto, xi.1815 (2 9, Zoologiske Museum, Copenhagen); Bogor:

Gunung Batu, 25.xii.1936, Van der Vecht, at light (1 &, L); Sukabumi (1 ¢, SMT);:

Gunung Tjerimai, 30.vi.1926 (1 9), 4.viii.1926 (1 3), Drescher (Bogor); Tjideres,,

24.xii.1935 (1 9), 10.1.1936 (3 9), i.1936 (1 g), vi.1936 (1 GF), vii.1936 (1 9),,

28.x.1936 (1 &), xi.1936 (3 4,4 9), 24.xii.1936 (2 J, 1 9), 27.xii.1936 (1 g), ili. 1937)

(1 9), vii.1940 (1 3g), xii.1940 (2 3), vi.1941 (1 9), xii.1941 (1 9), 1.1942 (1 I

ii.1945 (1 &), xii.1946 (2 9), iv.1948 (1 9), all Drescher, 35—100 m (Bogor, excepti

1 9, P); Gunung Muria: Tjolo, ii.1934 (1 &), 20.ii1.1934 (1 3), 30.1.1934 (1 3, Il

9), Van Doesburg (all L); Preanger (1 8, 3 9, BH); Java, Aulie (1 3, L); Java,,

one, xii.1815 (3, BM, Copenhagen, Institut Royal des Sciences naturelles de:

Belgique); no data at all (1 3, in last-mentioned museum, placed sub calanus by

Gillet).

13b. Bolbohamatum drescheri subsp. indosinicum nov.

(fig. 235710)

Material examined. — 3 males, 5 females.

Holotype male from Vietnam: Saigon (L, donated by Y. Cambefort). Paratypes

as follows:

Vietnam: Mont de Chaudoc, 1877, Harmand (1 9, P); Dong-Dang (Quang-Si

frontier), 1901, Lucas (1 ©, P); Lang-Son, 1904, Neau (1 3, P); Luc-Nam region,

Blaise (1 &, 1 9, P); Phu Tho, Duport (1 g, P); Saigon (1 g, Institut Royal des

Sciences naturelles de Belgique); Tonkin (1 g, P); Cochinchine, 1898, Amiral

Vignes (1 9, P); Cochinchine, Amaus (1 g, P); Cochinchine (1 &, P). —

Cambodia, no details (1 ©, SMT).

Some of these specimens were apparently seen by Paulian, and recorded as

calanus (1945: 41).

13c. Bolbohamatum drescheri subsp. birmanicum nov.

(figs. 21, 22)

Material examined. — 12 males, 15 females.

Holotype ¢ from Burma: Rangoon (P). Paratypes as follows:

Burma: Maymyo, Downing (1 &, 1 9, BM); Myitkyina, 30.viii-1.ix.1914 (1 ©,

BM); Rangoon, 8.vi.1898 (1 9, BM), ditto, v.1886, Fea (1 ¢, Genoa), ditto,

Meggitt (1 3, 1 9, U.S. National Museum), ditto, v.vi.1927 (1 Q, same museum),

ditto, 1933—34 (1 g, BM); S. Shan States: Taunggyi, 1.viii-22.ix.1934, Malaise,

1500 m (1 9, Naturhistoriska Riksmuseet, Stockholm); Tharrawaddy (1 9, BM);

Toungao (1 9, BM); N. Chin Hills (1 g, BM); Tenasserim (1 9, SMT); Lower

Burma (2 ©, BM); North/Upper Burma (1 g, 1 ©, BM); Burma (3 g, BM, P). —

Laos: Houei Sai, 2.vi.1918, Vitalis de Salvaza (1 9, BM); Paklay, 1963 (1 g, 1 9,

Zoologische Staatssammlung, Munich); Luang Prabang, ix.1917, Vitalis de Salvaza

(1 9, BM).

KRIKKEN: The genus Bolbohamatum 37

I have excluded from the type-series a small male (long 16 mm) with feeble

paramedian protrusions and subopaque elytra, with labels reading “Inde”,

“Palone/6.87”, “Ex Museo/Bonvoiloir”, from the Boucomont collection in P. This

may indeed be a representative of birmanicum, a geographical confirmation being

found in Boucomont & Gillet (1921: 70, “Palon, Birmanie’’).

Notes. — This subspecies of drescheri was by several workers, including G. J.

Arrow, identified as Bolboceras nigriceps Westwood, but I can find no reason to

accept this, the type, apparently a female, not having been recognized beyond

doubt in the material at hand. The present females of d. birmanicum do not agree

with Westwood’s drawing (1852: pl. 4 fig. 17).

Although the females from Burma and Laos listed above are slightly variable in

size, shape of frontal ridge and development of tibial protrusions, the material as a

whole seems homogeneous. Therefore these females, associated with males or not,

have all been labelled paratype.

DUBIOUS SPECIMENS, RECORDS, NAMES

Dubious Bolbohamatum specimens

Material not identified to species. — 29 females (probably nearly all calanus or

phallosum); 4 males. These males had no phallus, or they were damaged or

misshapen. Localities as follows:

India: Bangalore; Bellahunsi; Belgaum; Bombay; Dehra Dun; Getara; Igatpuri;

Kasawa (Bombay); Madras; Mahe; Namakal*; Nilgiri Hills; Surat; Sylhet;

Travancore: S. Bombay; E. India; N. W. India. — Singapore, Java (mislabelled!?).

Months vi—xi (9 specimens). Collections BM, P, SMT; Institut Royal des

Sciences naturelles, Brussels; Zoologiske Museum, Copenhagen; Senckenberg-

Museum, Frankfurt.

Before me are also 2 females seen by Westwood, one originating from the “Boys

Sale”, with a label in what seems to be Westwood’s hand, reading “calanus 97;

this specimen is in Oxford. The other, from the BM, is labelled “E. Ind//48/22”

(round label), and I wouldn’t be surprised if this were the specimen on which

Westwood based his nigriceps. There is a third old female specimen in the Paris

museum, from the collection of ‘“Laferté./5894”, “Ex-Musaeo/D. Sharp 1890”.

The female from Sylhet mentioned above (SMT, length 15.5 mm) has unusually

developed fossorial protrusions on the hind tibiae; the formula of the right tibia is

1c—2—3ps—4po—Spo, of the left Ic—2ps—3ps—4ps. The female from the

Nilgiris mentioned above (Brussels museum), is very large (19 mm long) and seems

different from other calanus-like females, i.a. by the frontal carina (crest in frontal

view bisinuate).

Dubious Bolbohamatum records

Material not recovered. — Under Bolboceras calanus Westwood, Paulian (1945:

41): Vietnam: Chapa; Pays Moi; Taiwan. The last record may be based on the

unreliable work of Miwa (1931).

38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFI. 1, 1980

Dubious names: possibly Bolbohamatum

Bolboceras bicarinatum Westwood, 1852: 24, pl. 4 fig. 15. — East India. — Typee

not recognized; female?

Bolboceras nigriceps Westwood, 1852: 25, pl. 4 fig. 17; 17 a—b (where are these?),),

— Distribution not mentioned. — Type not recognized (see also under dubiouss

specimens).

ACKNOWLEDGEMENTS

For the loan of specimens I am indebted to the following persons:

S. Adisoemarto (Bogor), M. E. Bacchus (London), A. Bons (Paris), You

Cambefort (Toulouse), P. N. Chatterjee (Dehra Dun), R. Damoisean (Brussels), J. .

C. Daniel (Bombay), A. Descarpentries (Paris), H. Freude (Munich), J. L. Gressitt

(Honolulu), W. Th. T. P. Gunawardana (Colombo), F. Hieke (Berlin), H. F.

Howden (Ottawa), J. C. Ismay (Oxford), R. Krause (Dresden), N. Mgller

Andersen (Copenhagen), R. Poggi (Genoa), R. D. Pope (London), R. zur Strassen

(Frankfurt), E. Taylor (Oxford).

Our staff artist A. Bos produced the habitus plate.

Financial support came from the Uyttenboogaart-Eliasen-Stichting (Amster-

dam).

REFERENCES

Boucomont, A., 1912. Scarabaeidae: Taurocerastinae, Geotrupinae. — Col. Catalogus 46: 47 pp.

Boucomont, A. & J. Gillet, 1921. Faune entomologique de l’Indochine française. Fasc. 4. Fam. Scara-

baeidae: 76 pp. — Saigon: Portail.

Fabricius, J. C., 1792. Entomologia Systematica: xx + (2) + 538 pp. — Hafniae: Proft.

Krikken, J., 1977a. The genus Bolbelasmus Boucomont in Asia, with notes on species occurring in other

regions (Coleoptera: Geotrupidae). — Zool. Meded. Leiden 51: 277—292, 19 figs, 1 pl.

———, 1977b. Asian bolboceratine scarabs of the genus Bolbogonium Boucomont (Coleoptera: Geo-

trupidae). — Tijdschr. Ent. 120: 77—108, 30 figs., 3 pls.

——., 1978. The Afro-Asian Bolboceroides validus group (Coleoptera: Geotrupidae). — Zool. Meded.

Leiden 52: 301—311, 13 figs., 2 pls.

Mayr, E., 1963. Animal species and evolution: vii + 797 PP., figs. — Cambridge, Mass.: Belknap Press.

Miwa, Y., 1931. A systematic catalogue of Formosan Coleoptera. — Report Dep. Agr. Govt Res. Inst.,

Taihoku, 55: xi + A-B + 359 pp.

Olivier, G. A., 1789. Entomologie, ou histoire naturelle des insectes. Coléoptéres. Tome 1: xx + 190 +

84 + 88 + 4 + 19 + ix + (2) pp., frontis-piece, 62 pls. — Paris: Baudoin.

Paulian, R., 1945. Coléoptéres Scarabéides de Indochine. 1. — Faune de l'Empire français 3: 228 pp.,

105 figs., | map.

Westwood, J. O., 1848a (Oct. 2). Descriptions of some new or imperfectly known species of Bolboceras.

— Proc. Linn. Soc. London 1: 384—387.

———, 1848b (Nov.). [Same title.] — Ann. Mag. nat. Hist. (2) 2: 353356. [Same text as 1848a.].

——., 1852. Descriptions of some new or imperfectly known species of Bolboceras, Kirby. — Trans.

Linn. Soc. London 21: 19—30, 2 pls.

wen.) Pab + us di in iar

rr menen girato Asa - cali

alber Sane Nick DEN cA et. ate

LOIR

ern, ar Wh. MS St ee à

mn 7 oe ads ren

eee

pe ma =

le a kad dm De sa aa

8 nt bec à fi 1

D “SS

EN Ul ve

| à

à Ar; ENGE a ei tote

n si 8 \ è

2 = 1e 6 :

a En ti

> i è

4 123 AFLEVERING 2 1980

| TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

FER WILLEMSE. — Classification and distribution of the species of Eupholidoptera

Ramme of Greece (Orthoptera, Tettigonioidea, Decticinae), p. 39—69,

fig. 1—190, pl. 1—2.

Tijdschrift voor Entomologie, deel 123, afl. 2, 1980 Gepubliceerd 29-11-1980

sf £ + DE

CLASSIFICATION AND DISTRIBUTION OF THE SPECIES

OF EUPHOLIDOPTERA RAMME OF GREECE

(ORTHOPTERA, TETTIGONIOIDEA, DECTICINAE)

FER WILLEMSE

Laurastraat 67, Eygelshoven, Netherlands

With 190 text-figures and 2 plates

ABSTRACT

On the basis of variability, several species and subspecies of the genus Eupholidoptera Ramme, 1951,

are redefined. Consequently some modifications of nomenclature are proposed: i.e., to synonymise E.

chabrieri magnifica (Costa, 1863) and E. chabrieri kaltenbachi Adamovic, 1972, with E. chabrieri schmidti

(Fieber, 1861); to consider E. garganica La Greca, 1959, a subspecies of E. chabrieri (Charpentier, 1825)

and E. chabrieri epirotica (Ramme, 1927) a distinct species; to synonymise E. danconai La Greca, 1959,

with E. megastyla (Ramme, 1939) and E. smyrnensis festae (Giglio Tos, 1914) with E. smyrnensis

(Brunner von Wattenwyl, 1882). Two new species are described: E. leucasi from the Ionian island of

Levkás and £. icariensis from the Aegean island of Ikaria. A key to the 14 species and two subspecies

known from Greece, including Uvarovistia (Karabagia) uvarovi (Karabag, 1952) together with a

distribution map are given.

INTRODUCTION

During the course of preliminary investigations of the Greek Orthoptera, the

important point emerged that the classification of some species of Eupholidoptera

Ramme, 1951, was not satisfactory. Apparently the amount of variation was not

‚always sufficiently known as some features seem to have been weighted

excessively. The need became obvious to revise some taxa. Since the erection of

thè genus in 1951, the number of species known from Greece has about doubled,

including two species described in this paper as new. The distribution of the

species in the Greek area was badly known due to insufficient material from the

mainland of Greece and confusing nomenclature in the past. On account of rich

material now available and the reasons mentioned above, it seems useful to give a

survey of the systematics and faunistics of the species of the genus occurring in

Kareece.

MATERIAL AND METHODS

Some material has been borrowed from the Instituut voor Taxonomische

Zoölogie, Amsterdam (ITZ), the Naturhistorisches Museum, Wien (NMW) and

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

the Istituto Policattedra di Biologia Animale, Catania (IBA) (abbreviations used in

the subsequent text are in parentheses). Additional Greek material (CW) has been

collected by the author and his son Lucas (accompagnied by J. Tilmans in 1976) in

1971 and all successive years up to 1978. The dates of capture range from Ist July

to 11th August, the altitudes of the localities from sealevel up to 2000 m. The

Greek orthography is transliterated into Roman characters in agreement with the

system proposed by the Permanent Committee on Geographical Names for British

Official Use, London. References listed under the species concern important

taxonomical ones, those recording original material from Greece and synonyms.

The figures are original unless otherwise stated, those of the epiphallus drawn after

dissection, slight maceration in 10% KOH solution and soaking in distilled water.

ACKNOWLEDGEMENTS

For loan of material and valuable information I am much indebted to Dr. A.

Kaltenbach, Vienna; Dr. G. Kruseman, Amsterdam and Dr. M. La Greca,

Catania.

SYSTEMATIC PART

Eupholidoptera Ramme, 1951

Eupholidoptera Ramme, 1951: 195, 197—211.

Type-species: Locusta chabrieri Charpentier, 1825.

Diagnosis. — Reference is made to the original description. Mainly defined as

follows: last abdominal tergite completely black in male, partly black in female;

Ovipositor narrow, as seen from above, i.e. strongly compressed laterally, and

comparatively long.

Differential diagnosis. — Reference is made to the key to the genera of the

Pholidopterini in Ramme (1951: 195).

Distribution. — The range of the genus covers southern Europe, Anatolia,

Syria, Lebanon and Israel. The distribution of the species in Greece is shown on

map |.

Remarks. — Hitherto about 30 species and some subspecies of Eupholidoptera

have been recognized. Ramme (1951) recorded eight species and one subspecies

from Greece. This number had increased considerably since. In this study the

following taxa are recorded from Greece:

E. chabrieri schmidti (Fieber, 1861)

E. chabrieri garganica La Greca, 1959, stat. nov.

E. leucasi sp.n.

E. epirotica (Ramme, 1927)

E. smyrnensis (Brunner von Wattenwyl, 1882)

E. megastyla (Ramme, 1939)

WILLEMSE: Eupholidoptera of Greece 41

E. prasina (Brunner von Wattenwyl, 1882)

E. icariensis sp.n.

E. spinigera (Ramme, 1930)

E. astyla (Ramme, 1927)

E. cretica Ramme, 1951

E. forcipata Willemse & Kruseman, 1976

E. latens Willemse & Kruseman, 1976

E. pallipes Willemse & Kruseman, 1976

E. gemellata Willemse & Kruseman, 1976

Eupholidoptera seems to be an unnatural grouping of species, as too much

emphasis has been placed on morphological features. Presumably quite a number

of species of Pholidopterini are not yet known. At this stage not much can be said

on generic or specific groupings. Regarding the species of Eupholidoptera from

Greece the following general observations can be made. Using the morphology of

Eupholidoptera

Ochabrieri schmidti

@chabrieri garganica

Mieucasi

[] epirotica

@ megastyla

* smyrnensis

S latens

Z pallipes

gemellata © Uvarovistia uvarovi

Map |. Distribution of Eupholidoptera species in Greece

42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

the male and female subgenital plate, the male cercus and the coloration, E.

chabrieri, E. leucasi, E. epirotica, E. smyrnensis and E. megastyla form a distinct

group. E. pallipes and E. gemellata differ from this group in more rounded pronotal

dorsum, form of epiphallus and the coloration. Similarly E. prasina, E. icariensis

and E. spinigera may be grouped together on account of the form of the male and

female subgenital plate and the unarmed male cercus. This leaves a more

heterogeneous assemblage of species, i.e. E. astyla, E. cretica, E. forcipata and E.

latens, with unarmed male cercus and resembling coloration but with diverse forms

of male subgenital plate and epiphallus.

Uvarovistia uvarovi (Karabag, 1952), originally described under Eupholidoptera,

seems closer to some species of the latter genus rather than to the remainder of

Uvarovistia. Therefore this species is treated in this paper together with the species

of Eupholidoptera.

Specific characters are found mainly in the external abdominal terminalia and

the epiphallus of the male, the female subgenital plate and the coloration in both

sexes. Sometimes the shape of the pronotal dorsum and the proportions of the

hind femur are useful. Taxonomic distinction is usually apparent, sometimes,

however, difficult. The male provides the most reliable, specifically diagnostic

features. The female characters are often not useful to differentiate between

closely allied taxa.

Key to the species and subspecies of Eupholidoptera and Uvarovistia from Greece

(males). (It is emphasized here that identification should be based on a study of the

male and its epiphallus.)

1. Black coloration of distal abdominal tergites confined to last tergite. Cercus

slender, much longer than greatest width of base, slightly tapering apically

(igs: JAS "eS ee RN ee 2

— Black coloration of distal abdominal tergites not confined to last tergite, but

extending proximally over lateral parts of penultimate tergite. Cercus, if

viewed from above, almost triangular, greatest width of base almost equal to

length of cercus (fig. 70)(Karpathos) ....... Uvarovistia uvarovi (Karabag)

2. Cercus with hook-shaped basal or subbasal tooth (figs. 34— 65) ...... 3

— Cercusunarmed (figs? 66—69)". Ae A ee U ODE 10

3. Tooth of cercus situated at base (figs. 34—51,54—65) ............ -

— Tooth of cercus situated just distal to base (figs. 52—53) ........... 8

4. Emargination of hind margin of last abdominal tergite very wide, about as

wide as length of cercus (figs. 28—29) (Makedhonia, Thraki, Thasos, Limnos,

Lesbos, Samos, Nisiros, Ródhos) .. . . smyrnensis (Brunner von Wattenwyl)

— This emargination much narrower (figs. 1—27) ................ 5

5. Extending apical parts of epiphallus very close together, slender and straight

(figs. 173—180, 184, 185) (Zákinthos, Pelopónnisos, Steréa, Ellas, Thessalia,

SE Ipiros, SW Makedhonia) - . BR... oe ya oe megastyla (Ramme)

— These parts widely separated from each other (figs. 119, 164, 169—170) .. 6

6. Pronotal dorsum of general colour but middle of prozona black (pl. | figs.

3—4). Fused part of epiphallus slender, convex laterally, without plate-like

WILLEMSE: Eupholidoptera of Greece 43

lateral expansions, as in figs. 169—170 (Ipiros, Kérkira) . . epirotica(Ramme)

Pronotal dorsum entirely of general colour. Fused part of epiphallus wider,

plate-like, lateral expansions more or less developed (figs. 119—164) ... 7

. Epiphallus with fused part strongly inflated and extending apical parts strongly

recurved, ranging as in figs. 153—162 (Kerkira, W. Ípiros)

een Else ER SF ODE chabrieri garganica La Greca

These characters less apparent, as in figs. 125—148 (E. Ipiros, W. & N.

KELDER EN chabrieri schmidti (Fieber)

. Pronotal dorsum slightly flattened. Pronotal lateral lobe black, lower margin

broadly bordered yellowish (pl. 1 fig. 2). Epiphallus with fused part very short,

wide and extending apical parts well separated from each other, very long and

strongly recurved (figs. 165—168) (Levkás) ............. leucasi sp.n.

Pronotal dorsum smoothly rounded. Pronotal lateral lobe of general colour

with black dot dorso-posteriorly (as in pl. 1 fig. 6, pl. 2 fig. 8). Epiphallus with

fused part long, without well separated extending apical parts (Willemse &

U an OO 1195.35 38), sr an sa BORA ns ee ah fs à 9

. Tip of epiphallus with a lateral spine on each side (Willemse & Kruseman,

PWG pl. 1185, 35—36) (Kriti). tane pallipes Willemse & Kruseman

Tip of epiphallus laterally rounded and without spines (Willemse & Kruseman,

[9N6#plS/fies.37—38)(Kriti), en gemellata Willemse & Kruseman

. Lobes of subgenital plate with needle-like apex (figs. 112—115) ..... Il

These lobes with obtuse or rounded apex (figs. 117—118; Willemse &

Kruseman, 1976: figs. 2, 7, pls. 4—5 figs. 14—15,21—22) ......... 13

. Pronotal lateral lobe black, lower margin widely bordered yellowish. Styli set

more proximad, their tips not reaching apex of subgenital plate (Ramme, 1930:

CLORO TSO N Mean spinigera (Ramme)

Pronotal lateral lobe of general colour with black dot dorso-posteriorly (pl. 1

fig. 6, pl. 2 fig. 8). Styli set more distad, their tips reaching apex of subgenital

zie bns Le o ee EE EE 12

. Epiphallus with extending apical parts distinctly divergent, as in fig. 186

TT ER oe ere eee en er ON PP icariensis sp.n.

Epiphallus with apical part expanding laterally with median apical incision, as

in figs. 189— 190 (Samos, Khios) ....... prasina (Brunner von Wattenwyl)

. Lobes of subgenital plate rounded apically, styli set pre-apically on ventral

surface and pointing ventrad (figs. 117—118; Willemse & Kruseman, 1976: pls.

ei EDEN RO ERE PAT OT O 14

Lobes of subgenital plate narrowly obtuse apically, styli set on apex and

pointing distad (Willemse & Kruseman, 1976: fig. 7, pl. 5 fig. 21) ..... 15

. Epiphallus asymmetrical, extending apical parts close together and pointing

Bien (ele) NAXOS AK EH) A. sont IL) mart Jota! ’y (a) oe astyla(Ramme)

Epiphallus symmetrical, extending apical parts divergent and pointing dorsad

(Willemse & Kruseman, 1976: pl. 7 figs. 32—34) (Kriti)

Sy ON latens Willemse & Kruseman

. Apical parts of epiphallus fused over almost whole length, tip only slightly

divergent (Willemse & Kruseman, 1976: fig.9)(Kriti) ..... cretica Ramme

Apical parts of epiphallus fused in proximal half, strongly divergent, hook-like

44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

and recurved in distal half (Willemse & Kruseman, 1976: pl. 6 figs. 29—31)

ARTE À a a AOT forcipata Willemse & Kruseman

13 21 25

OAN

da x Pu NG Vai

oe A iba

Figs. 1—33. Emargination of the hind margin of the last abdominal tergite in the male of Eupholidoptera

species: 1—15, E. chabrieri (s.1.) (1—2, E. chabrieri chabrieri (Charpentier): (1) Col de l’Ablé, Luceram;

(2) Mt. Puzzillo, Velino Mts.; 3—10, E. chabrieri schmidti (Fieber): (3) Sperlonga, Latina; (4) Kanesta,

Istra; (5) Uéka, Istra; (6) Alona, Florina; (7) Evzonoi, Kilkis; (8) Klisoúra, Kastoria; (9— 10) Kalpakion,

Ioannina; 11—14, E. chabrieri garganica La Greca: (11—12) Panayia, Preveza; (13) lake Antinioti, Ker-

kira; (14) Benitzes, Kerkira; 15, E. chabrieri bimucronata (Ramme), Colle S. Rizzo, Messina; 16—18,

E. leucasi sp. n., paratypes; 19—20, E. epirotica (Ramme): (19) holotype; (20) paratype; 21—27, E. me-

gastyla (Ramme): (21—22) Kámbos, Messinia; (23—24) Lekhaina, Ilia; (25) Mt. Pilion, Magnisia;

(26—27) Rámia-Livádhion, Arta; 28—29, E. smyrnensis (Brunner von Wattenwyl): (28) Souflion; (29)

Amörion, both Evros; 30—31, E. prasina (Brunner von Wattenwyl): (30) holotype; (31) Samos; 32,

E. icariensis sp. n., holotype; 33, E. astyla (Ramme), Skonef.

WILLEMSE: Eupholidoptera of Greece 45

Eupholidoptera chabrieri (Charpentier, 1825)

(figs. 1—15, 34—51, 71—90, 119—164)

Locusta chabrieri Charpentier, 1825: 119.

Thamnotrizon chabrieri; Brunner von Wattenwyl, 1861: 293, pl. 10 figs. 6a—d.

Pholidoptera schmidti; Ramme, 1927: 130, figs. 10a, 11a; 1930: 809, figs. 5—7, 9.

Pholidoptera chabrieri; Chopard, 1951: 128, figs. 210, 211, 223.

Eupholidoptera chabrieri; Ramme, 1951: 198, figs. 44, 45, 48, 50; Harz, 1969: 370, figs. 1116, 1121—1124,

1131—1142 (partim).

Diagnosis. — Reference is made to general descriptions by previous authors.

The species is well defined in the male, i.e. by the moderate emargination of the

hind margin of the last abdominal tergite (figs. 1—5), cercus with basal tooth (figs.

34—51), form, spines (figs. 71—90) and styli of subgenital plate, epiphallus (figs.

119—164) and coloration.

Differential diagnosis. — The male is, with few exceptions, readily

distinguishable from other species. Judging from the descriptions and figures, the

distinction between E. chabrieri and E. marani Peshev, 1960, and E. beybienkoi

Peshev, 1962, is not clear. The female of E. chabrieri is indistinguishable from that

of its closest relatives, e.g. in Greece E. leucasi and E. megastyla.

Remarks. — Hitherto the following subspecies have been recognized by La

Greca (1959), Harz (1969) and Adamovié (1972): E. chabrieri chabrieri

(Charpentier, 1825), S. France, S. Switzerland, N.W. & N. & (?) Central Italy; E.

chabrieri brunneri (Targioni-Tozzetti, 1881), considered of doubtful validity, (?) N.

& Central Italy; E. chabrieri magnifica (Costa, 1863), (?) Central Italy, S.W. Italy,

(?) Sardegna; E. chabrieri bimucronata (Ramme, 1927), Sicilia, (?) Sardegna; E.

chabrieri schmidti (Fieber, 1861), N.E. Italy, Istra; E. chabrieri usi Adamovié, 1972,

Adriatic Islands of Krk, Cres, Loëinj; E. chabrieri galvagnii Adamovié, 1972,

Hercegovina, Montenegro; E. chabrieri kaltenbachi Adamovié, 1972, Serbia,

Yugoslav Makedonija; E. chabrieri epirotica (Ramme, 1927), N.W. Greece.

These subspecies were recognized mainly on the basis of differences of the

epiphallus. In some cases also very slight distinction of external male abdominal

terminalia, ovipositor, measurements and coloration were noted or figured.

Analogously, an attempt was made to arrange the Greek material at hand. This,

however, failed largely, due to considerable variation. Exception is made for E.

epirotica, which is regarded here a distinct species. But the remainder could be

arranged only roughly, i.e., populations agreeing with several known or

representing unknown forms of E. chabrieri (s.1.), those agreeing with E. garganica,

and others linking both. Thus the need became obvious for some review of the E.

chabrieri-complex.

A careful study of the literature and rich material before me (ITZ, CW, IBA),

covering most of the range of the species, reveals that, with few exceptions, the

characters overlap to such an extent that they cannot be considered reliable (figs.

I—15, 34—51, 71—90). Particularly it seems that, at least in some cases, too great

a diagnostic emphasis was placed on the form of the epiphallus (figs. 119—164).

Distinction of very few geographical races or subspecies seems justified, e.g. the

nominate form (figs. 119—124) and E. chabrieri bimucronata (figs. 163—164).

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

However, the distinction between E. chabrieri magnifica (figs. 125—126) and E.

kaltenbachi (figs. 131—136) is not clear and the distinction between these taxa and

42 43 | | |

6 54

34 35 | 36 37 38 39 40 | 41

Figs. 34—54. Left male cercus of Eupholidoptera species, dorsal views: 34—S1, E. chabrieri (s.l.)

(34—35, E. chabrieri chabrieri (Charpentier): (34) le Plan du Peiron, Grasse; (35) Col de Brouis, Castel-

lane; 36—48, E. chabrieri schmidti (Fieber): (36—37) Ucka, Istra; (38—39) Alona, Florina; (40) Klisou-

ra, Kastorfa; (41) Botzarás, (42) Kaléntzion, (43—44) Kalpakion, (45—46) Asfaka, all Ioannina; (47)

Katarräktis, Arta; (48) Sperlonga, Latina; 49—50, E. chabrieri garganica La Greca, Gazatika, Kérkira;

51, E. bimucronata (Ramme), Colle S. Rizzo, Messina; 52—S3, E. leucasi sp. n., paratypes; 54, E. epiroti-

ca (Ramme), paratype.

WILLEMSE: Eupholidoptera of Greece 47

E. chabrieri schmidti (from Istra) (figs. 127—130) refers merely to different

measurements. On account of these observations it appears necessary to propose

some modifications of the nomenclature (see below).

Kaltenbach (1967) gave a survey of the confusing way in which the name

chabrieri has been used previously. Records under this name from the period 1927

to 1951 usually refer the name to E. smyrnensis, while for chabrieri as now

understood usually the name schmidti has been used. Previous records of Greek

material referred to E. chabrieri are now assigned to E. chabrieri garganica, E.

chabrieri schmidti, E. epirotica and E. megastyla.

Distribution. — The range of chabrieri (s.1.) covers S. France, S. Switzerland, all

of Italy except for its extreme South East, Sicilia, Sardegna and some neighbouring

islands, the Adriatic islands, all of Yugoslavia, Albania, extending into N.W.

Greece, Bulgaria (?) and Romania. Kaltenbach (1967) pointed out that the species

does not occur in Anatolia.

Eupholidoptera chabrieri schmidti (Fieber, 1861)

(figs. 3—10, 36—48, 73—83, 125—148)

Thamnotrizon schmidti Fieber, 1861: 197 (nec Corfu).

Thamnotrizon magnificum Costa, 1863: 28. Syn. nov.

Thamnotrizon chabrieri; Brunner von Wattenwyl, 1882: 334 (partim).

Pholidoptera schmidti; Ramme, 1927: 130, figs. 10a, lla.

Eupholidoptera chabrieri; Ramme, 1951: 198, 204, 206, 207, 209, figs. 44, 45, 48, 50.

Eupholidoptera chabrieri schmidti; La Greca, 1959: 62, figs. 68, 69; Harz, 1969: 370, 371, fig. 1116; Ada-

movic, 1972: 195, 198, figs. 1—3.

Eupholidoptera chabrieri kaltenbachi Adamovic, 1972: 198, figs. 1—3. Syn. nov.

Material studied. — Makedhonia: Kilkis: Evzonoi, 34; Plataniá, 19; Florina:

Alona, 2 km E., 24 39; Kastoria: Klisoura, 1g (all CW). Ípiros: (=Epirus), Erber,

Coll. Br. v. W., 6885, Pholidoptera schmidti Fieb. det. Ramme, 14 19 (NMW);

Ioannina: Pápingon, 4¢ 19; Kalpákion, 44 49; Asfäka, 35 59; Kónitsa -

Eléftheron, 13; A. Paraskevi, Mt. Smölikas, 24; Koutseliön, 13; Kaléntzion, 2¢

39 ; Botzarás, 2¢; Vrosina, 34 19; Arta: Katarraktis, 1g (all CW).

Diagnosis. — Epiphallus with extending apical parts usually divergent and, in

lateral view, slightly to moderately recurved; fused part inflated only

longitudinally in the middle and provided with well developed plate-like lateral

expansions, as in figs. 125—148.

Variation. — It seems appropriate to give series of figures covering the

considerable amount of variation: epiphallus in figs. 125—148, male cercus in figs.

36—48, spines of male subgenital plate in figs. 73—83, emargination of hind

margin of male last abdominal tergite in figs. 3—10.

Differential diagnosis. — The distinction between E. chabrieri schmidti and the

nominate subspecies appears arbitrary, being difficult to define. It is supposed that

the extending apical parts of the epiphallus are parallel or almost so in the

nominate form (figs. 119—124) against slightly divergent in E. chabrieri schmidti

(figs. 127—130). The characters of some material at hand, however, disagree with

this assumption (figs. 137—138, 147—148). The distinction from E. chabrieri

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

aude

pi È FUGA 93

ae 103

wh 80 LE E N ae a Ton

= Ar sue oo A

AL a

83 ARTI SY!

4 Ne

90 95

100

/ 106

/ ae

/ Ai =

Figs. 55—70. Left male cercus of Eupholidoptera species, dorsal views: 55—63, E. megastyla (Ramme):

(55—56) Kallithéa, Ilia; (57—59) Ramia — Livädhion, Arta; (60) Akhladhokambos, Argolis; (61) Me-

lidhoni, Fthiótis; (62) Mt. Olimbos, Larissa; (63) Lekhaina, Ilia; 64—65, E. smyrnensis (Brunner von

Wattenwyl): (64) Souflion, Evros; (65) Yerakini, Khalkidhiki; 66—67, E. prasina (Brunner von Watten-

wyl): (66) Samos; (67) holotype; 68, E. icariensis sp. n., holotype; 69, E. astyla (Ramme), Skonef; 70,

Uvarovistia (Karabagia) uvarovi (Karabag), Mt. Lastros, Karpathos. Figs. 71—106. Spines on apical mar-

gin of lobes of male subgenital plate in Eupholidoptera species: 71—90, E. chabrieri (s.1.) (71—72),

E. chabrieri chabrieri (Charpentier): (71) Col de Brouis, Castellane; (72) le Plan du Peiron, Grasse;

73—83, E. chabrieri schmidti (Fieber): (73) Uéka, Istra; (74) Evzonoi, Kilkis; (75) Klisoúra, Kastoria;

(76) Epirus, Erber leg.; (77—78) Botzaräs, (79) Kaléntzion, (80) Kalpákion, (81—82) Asfaka, all loanni-

(

\ \ \ | | \ |

Zord Ae 7

we / >

WILLEMSE: Eupholidoptera of Greece 49

garganica is pointed out below, that from E. chabrieri bimucronata is given in figs.

51, 90, 163— 164.

Remarks. — As outlined above, part of the Greek material agrees with several

subspecies as hitherto understood. This may be demonstrated by comparison of

figs. 125—126 and 131—132 with figs. 133—136 showing the epiphalli of E.

chabrieri magnifica, E. chabrieri kaltenbachi and Greek specimens, respectively; or

of figs. 127—130, representing E. chabrieri schmidti from Istra, and figs. 139— 140

and 145—146 made after Greek specimens. The remainder of the Greek material

does not fit precisely into any known form (figs. 137—138, 141—144, 147—148).

Apparently the degree of variation, at least among Greek populations, is larger

than has been thought. I propose to allocate all this material to E. chabrieri schmidti

and consider E. chabrieri magnifica (Costa, 1863) and E. chabrieri kaltenbachi

Adamovic, 1972, synonyms of E. chabrieri schmidti (Fieber, 1861). Whether the

distinction between this subspecies and the nominate form is justifiable, is quite

doubtful. This question, however, is not settled here.

Distribution. — The range of the subspecies as here understood is not well

defined. I studied material from Istra and Slovenija, Hrvatska, Serbia, Yugoslav

Makedonija, N.W. Greek Makedhonia, E. fpiros (map), Central and S.W. Italy.

Eupholidoptera chabrieri garganica La Greca, 1959, stat. nov.

(figs. 11—14, 49— 50, 84—89, 153— 162)

Thamnotrizon schmidti Fieber, 1861: 197 (only Corfu).

Eupholidoptera garganica La Greca, 1959: 66, figs. 46, 50, 64, 65, 70, 71; Kaltenbach, 1967: 190; Harz,

1969: 363, 370, 372, figs. 1118, 1119, 1145.

Eupholidoptera chabrieri schmidti; Cejchan, 1963: 771 (partim?).

Material studied. — Kerkira: (=Corfu), Erber, coll. Br. v. W., 5860, Pholidoptera

schmidti Fieb. det. Ramme, 1g; Benitses, 27.vii.1965, A. Kaltenbach, 1g (both

NMW); Yimärion, 7¢ 79; Gazatika, 7Z 69; Pélekas, 14 19; Tsaki, 1¢ 19;

Lavkion, 1g; Lake Antinioti, 34 49; Lake Korission, 4g 39; Petália, 7g 49;

Perithia, 73 39; Thesprotia: Igoumenitsa, 1g; Préveza: Panayia, 34 20:

S. & N. of Koroni, 1g; Nikópolis, 343 229; Ioannina: Mt. Tómaros 94 29 (all

CW). Italy: Foresta Umbra (Gargano), 24.vii.1957, Eupholidoptera garganica n.sp.

La Greca det., 13 paratype, 19 allotype (IBA).

Diagnosis. — Epiphallus with extending apical parts, in lateral view, strongly

recurved; fused part strongly inflated in combination with strongly reduced size of

plate-like lateral expansions, as in figs. 153—162.

Variation. — The Greek material varies considerably. The shape of the

epiphallus varies from the type of E. garganica all to that met with in E. chabrieri

schmidti (figs. 153—162). Variation of the cercus, spines of the subgenital plate and

na; (83) Sperlonga, Latina; 84—89, E. chabrieri garganica La Greca: (84—86) Gazätika, (87) lake Anti-

nioti, (88) Benitses, all Kérkira; (89) Panayia, Préveza; 90, E. chabrieri bimucronata (Ramme), Colle S.

Rizzo, Messina; 91—94, E. leucasi sp. n., paratypes; 95, E. epirotica (Ramme), holotype; 96—102,

E. megastyla (Ramme): (96—98) Mt. Olimbos, Larissa; (99—100) Bassae, Ilia; (101—102) Mt. Eriman-

thos, Akhaia; 103—105, E. smyrnensis (Brunner von Wattenwyl): (103) Thäsos; (104) Tikherón, Evros;

(105) Yerakini, Khalkidhiki; 106, Uvarovistia (Karabagia) uvarovi (Karabag), Mt. Lastros, Karpathos

50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

emargination of the hind margin of the last abdominal tergite of the male as shown

in figs. 11—14, 49—50, and 153—162.

Differential diagnosis. — The subspecies may be recognized only by the form of

the epiphallus. As a result of the large amount of variation the decision whether a

given specimen has to be arranged under E. chabrieri garganica or not is sometimes

arbitrary. The series of figures of the epiphallus of E. chabrieri garganica and other

subspecies of E. chabrieri suggest their distinction proposed here (figs. 119— 164).

Remarks. — The occurrence of transitional forms, in Greece, between typical

E. garganica (figs. 157—158) and E. chabrieri schmidti indicates that both taxa are

closely related. Further indication of their close relationship is found in the forms

described by Adamovié (1972) under the names E. chabrieri usi and E. chabrieri

galvagnii, the former from some northern Adriatic islands and the latter from

Hercegovina and Montenegro. These forms, defined by the form of the epiphallus

(figs. 149—152), also link E. chabrieri schmidti and E. garganica. In a general way

one can see that the epiphallus is subject to a geographical variation in which the

recurvation of the extending apical parts, the inflation of the fused part and the

reduction in size of the plate-like lateral expansions increases from Istra in the

North along the Adriatic coastal area of Yugoslavia into the Gargano area of Italy

and the coastal area of N.W. Greece in the South. Distinction at species level

between E. garganica and E. chabrieri appears no longer justifiable and I propose

to merge E. garganica in the E. chabrieri complex as subspecies. The question

whether or not E. chabrieri usi and E. chabrieri galvagnii should be arranged under

either the subspecies schmidti or garganica is not settled here.

Distribution. — The range of the subspecies as now understood covers the

Gargano-Foggia area of E. Italy, the Ionian island of Kérkira and the opposite part

of the mainland of Greece (map). As already noted by Kaltenbach (1967) it is quite

possible that the subspecies also occurs in Albania (Cejchan, 1963).

Eupholidoptera leucasi sp.n.

(pl. 1 figs. 1—2; figs. 16—18, 52—53, 91—94, 165— 168)

Material studied. — ¢ holotype, © allotype, 9 4, 16 © paratypes, labelled:

Hellas, N. Levkádos, Levkas, Ano Exánthia, 600 m, 15.vii.1976; additional

paratypes: Levkás, Nikiäna, N. of Nidhrion, 5 m, 14.vii.1976, 4 3,3 © (all F. & L.

Willemse & J. Tilmans, CW).

Diagnosis. — Differs from type-species as follows: &, pl. 1 figs. 1—2. Median

emargination of hind margin of last abdominal tergite slightly larger (figs. 16—18).

Cercus with strong inner tooth located about rectangularly and shortly distad from

base of cercus (figs. 52—53). Hind margin of lobes of subgenital plate above styli

stretched and terminating in two more or less closely set spines of about equal

length (figs. 91—94). Epiphallus of remarkable form (figs. 165—168): fused part

very short, very wide and conspicuously inflated without lateral plate-like

expansions, anterior surface with median longitudinal groove and crossed by some

transverse grooves; extending apical parts arising from dorso-lateral edges of fused

WILLEMSE: Eupholidoptera of Greece SI

| 2 ii

| \ 4 | ì NS NS

= ZE LC

Figs. 107—111. Stylus of male subgenital plate in E. megastyla (Ramme), ventral views: 107—108,

Dhimitra-Anatoli, Larissa; 109, Elasson, Larissa; 110—111, Kallithéa, Ilia. Figs. 112—118. Subgenital

plate in Eupholidoptera species, ventral and/or lateral views: 112—113, E. prasina (Brunner von Watten-

wyl): (112) 4, Khios; (113) g, Samos; 114—116, E. icariensis sp. n.: (114—115) g, holotype; (116) 9,

allotype; 117—118, E. astyla(Ramme), g, Skonef

part, parallel or slightly converging or diverging, comparatively very long, their

tips reaching in situ the tooth of the cercus and in lateral view always strongly

recurved in basal fourth.

9. Much resembling type-species. Subgenital plate with moderately deep

median V-like incision, lobes angulately rounded, the very hind margin sometimes

curled inward. Last abdominal sternite slightly projecting in the middle.

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

Measurements (length in mm): body & 26—32, 9 25—32; pronotum

& 11.3—13.2, © 11.4—13.9; hind femur 8 24.5—28.8, 9 25.0—29.5; ovipositor

20.1—23.5.

Variation. — Shape and size of median emargination of hind margin of the male

last abdominal tergite vary slightly, as in figs. 16—18. Inner tooth of male cercus

varies somewhat in size, the location, however, is invariable (figs. 52—53). Spines

of male subgenital plate range from divergent to very close to each other, as in figs.

91—94. The inflation of the fused part of the epiphallus may be less strong, the

grooves less apparent, but the short, wide, transverse appearance is always obvious

(figs. 165— 168).

Differential diagnosis. — The species is well defined in the male by its

resemblance with the type-species, E. chabrieri, in combination with the location

of the cercal inner tooth and the unique form of the epiphallus. The main feature

of the latter refers to the combination of strongly recurved and basally widely

separated extending apical parts with very short and widely inflated fused part. In

E. hesperica La Greca, 1959, the fused part is also very short but apparently less

wide and the extending apical parts by far less strongly recurved. Somewhat

similar extending apical parts are found in E. chabrieri garganica La Greca,

E. palaestinensis (Ramme), EF. anatolica (Ramme), E. forcipata Willemse &

Kruseman and E. ledereri (Fieber). In these species, however, the fused part of the

epiphallus is quite different. Opportunity is taken here to figure more fully the

epiphallus of E. palaestinensis (figs. 171—172) (Ramme, 1939: fig. 26p, dorsal view

only). The female of the new species is indistinguishable or almost so from that of

the type-species and some others, e.g. E. megastyla.

Remarks. — The distinction of E. leucasi at species level is justifiable on account

of obvious and stable differences in at least two important characters and absence

of intermediate forms, especially in regard to E. chabrieri garganica, whose range

adjoins that of the new species. The tendency of the geographical variation of the

epiphallus among E. chabrieri in the Adriatic area, as pointed out above, reaches a

climax in E. leucasi. The new species was found along the road in Rubus, Cirsium

and other prickly shrubs.

Distribution. Known only from the Ionian island of Levkás (map).

Eupholidoptera epirotica (Ramme, 1927)

(pl. 1 figs 3—4; figs. 19—20, 54, 95, 169—170)

Thamnotrizon chabrieri; Brunner von Wattenwyl, 1882: 335 (Varietät aus dem Epirus).

Pholidoptera epirotica Ramme, 1927: 132, figs. 11b, 12.

Pholidoptera schmidti epirotica; Ramme, 1930: 812.

Eupholidoptera epirotica; Ramme, 1951: 198, 203, 206, 207, 209, figs. 44e, 48e, 50; La Greca, 1959: 67;

Kaltenbach, 1967: 197.

Eupholidoptera chabrieri epirotica; Harz, 1969: 371, figs. 1122, 1140, 1142.

Material studied. — & holotype, ¢ paratype, labelled: Epirus, Erber Coll. Br. v.

W., Pholidoptera epirotica Ramme det. Ramme, Type & Paratypus; Kerkira:

(= Corfu), coll. Werner, 1 © (all NMW).

WILLEMSE: Eupholidoptera of Greece 53

124

133 134

130

Figs. 119—130. Epiphallus in Eupholidoptera species, anterior and left lateral views: 119—130, E. cha-

brieri (s.1.): 119—124, E. chabrieri chabrieri (Charpentier): (119—120) Col de Brouis, Castellane;

(121—122) Col de l’Able, Luceram; (123—124) Mt. Puzzillo, Velino Mts.; 125—130, E. chabrieri

schmidti (Fieber): (125—126) Sperlonga, Latina; (127—130) Ucka, Istra

Diagnosis. — Differs from type-species as follows: &, pl. 1 fig. 3. Emargination

of hind margin of last abdominal tergite slightly wider (figs. 19—20). Cercus

slightly longer and more slender (fig. 54). Epiphallus with extending apical parts, in

lateral view, very slightly recurved; fused part without plate-like lateral

expansions, much longer than wide, tapering basally, regularly convex laterally,

anterior surface with median longitudinal groove (figs. 169—170).

Occiput and prozona of pronotal dorsum broadly black in the middle.

Q, pl. 1 fig. 4. Lobes of subgenital plate slightly more broadly rounded.

Coloration as male.

Variation. — Morphological features of both males are similar. The coloration

of the type and the female are similar, that of the paratypic male, however, differs

in the less conspicuously black coloration of the occiput and the pronotum.

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

Differential diagnosis. — The species is defined by the resemblance to the type-

species, E. chabrieri, in combination with the characteristic form of the epiphallus.

The reliability of other features, e.g. the black coloration of occiput and

pronotum, cannot be judged because of lack of material.

Remarks. — In the collection of Brunner von Wattenwyl there is another pair

bearing locality labels similar to those of the types of E. epirotica. This pair is quite

distinct and agrees fully with E. chabrieri schmidti. Similarly, no specimens were

found among other material linking E. epirotica with E. chabrieri.

On account of the apparently distinct form of the epiphallus, lack of

intermediate forms and the sympatric occurrence of E. chabrieri (s.l.), E. epirotica

is provisionally regarded a distinct species. Additional material, however, is

needed to establish this assumed status.

Distribution. — Known only after the types from Ipiros and the female from

Kérkira (Kaltenbach, 1967), unfortunately both without precise localities (map).

Eupholidoptera megastyla (Ramme, 1939)

(figs. 21—27, 56—63, 96— 102, 107—111, 173—180, 184, 185)

Thamnotrizon chabrieri; Brunner von Wattenwyl, 1882: 334 (only Parnass).

Pholidoptera chabrieri; Werner, 1933a: 407 (?); 1938: 169 (2).

Pholidoptera megastyla Ramme, 1939: 101, fig. 28.

Eupholidoptera megastyla; Ramme, 1951: 198, 203, 205—207, 211, figs. 48, 50; La Greca, 1959: 65; Kal-

tenbach, 1967: 197; Harz, 1969: 362, 367, 373, figs. 1110, 1148—1150; Willemse, 1971: 17; 1974: 352.

Eupholidoptera danconai La Greca, 1959: 65, figs. 47, 49, 63. Syn. nov.

Material studied. — Makedhonia: Pélla: Edhessa, 23.viii.1965, Blommers e.a.,

1g 49 (ITZ); Kozani: Velvendós, 9g 6 9; Neápolis, Sg 19; Grevena:

Paraskevi, near Dheskáti, 2 2; Piéria: Ftéri, 1 3; Litokhoron, 15 g 17 9 (all CW);

Mt. Olimbos, W. of Litökhoron, 1450 & 1750 m, 13.vii. & 2.viii.1965, 3 9;

Kolindrós, 10 km W., 24—26.vii.1965, 1 3 (both Blommers e.a., ITZ); Leptokaria

- Kariá, 2 &. Thessalia: Larissa: Mt. Olimbos, refuge B, 8 ¢ 19 9; Elassón, 20 km

NW., 1 4, 4kmW.,2 g 2 9 (all CW); Tirnavos, 17.viii.1965, Blommers e.a., | g

4 9 (ITZ); Dhimitra - Anatoli, 5 24 9; Magnisia: Mt. Pilion, above Portaria,

1 3 (CW); Lekhónia, 7—11.viii.1965, Blommers e.a, 1 & (ITZ). Ipiros: Arta:

Ramia-Livadhion, 6 4 8 ©. Stereá Ellas: Aitolia - Akarnania: Kokkinokhorion,

28 39 (both CW). Fthiótis: Pelasyia, 6.vii.1976, J. P. Duffels, 1 & (ITZ);

Timfristós village, 1 & 19; Mt. Oiti, above Pavliani, 1 9; Melidhoni, 6 3 109;

Fokis: Mt. Vardhousia, above Mousounitsa, 1¢; Evvoia: Steni-Kathenoi, 23

29 (all CW); Kimi, 30.vi.1971, A. Kaltenbach, 1 & (NMW); Voiotia:

Mt. Parnassós above Arákhova, 24 (CW), Krüper coll. Br. v. W., 1704,

Pholidoptera schmidti Fieb. det. Ramme, 1 © (NMW). Pelopónnisos: (= Morea),

Staudinger, coll. Br. v. W., 19.524, Eupholidoptera megastyla Ramme det.

Kaltenbach, 1 ¢ (NMW); Korinthia: A. Eléni, 1 g 4 9; Argolis: Navplion, 3 9;

Ligourion, 2 3; Mt. Kteniäs, Akhladhókambos, 11 ¢ 7 9; Lakonia: Mistrás, 2 3;

Skála, | & 1 9; Mt. Taiyetos, above Anóyia, 4 ¢ 10 9, E. of Spárti, 38 3 9;

Messinia: Kambos, 8 3 9 9; Mikrá Mandhinia, 1 g; Arkadhia: Karitaina, | &;

Ipsous 5 & 5 9; Khrissovitsi, 8 & 2 9; Kapsia, 1 g 1 9; Mt. Párnon 2 4 1 9 (all

WILLEMSE: Eupholidoptera of Greece

Figs. 131—148. Epiphallus in Eupholidoptera species, anterior and left lateral views: 131—148, E. cha-

brieri schmidti (Fieber): (131—132) Yugoslav Makedonija (E. chabrieri kaltenbachi Adamovic, after

Adamovié, 1972); (133—134) Alona, Florina; (135—136) Evzonoi, Kilkis; (137—138) Klisoura, Kas-

toria; (139—142) Kalpákion, (143— 144) Kaléntzion, (145—146) Botzaras, all Ioannina; (147—148) Ka-

tarraktis, Arta

CW); Dhimitsana, 14.vii.1977, M. & J. Duffels, 1g 19 (ITZ); Akhaia:

Mt. Panakhaikon, above Zástova, 5 & 2 9; Mt. Frimanthos, above Káléntzion,

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

150

Figs. 149—160. Epiphallus in Eupholidoptera species, anterior and left lateral views: 149—150, E. cha-

brieri galvagnii Adamovié, 151—152, E. chabrieri usi Adamovié (both after Adamovié, 1972); 153— 160,

E. chabrieri garganica la Greca: (153—154) Perithia, (155—156) lake Antinioti, both Kérkira;

(157— 158) Foresta Umbra, Gargano (paratype of E. garganica); (159—160) Gazätika, Kérkira

WILLEMSE: Eupholidoptera of Greece 57

3 &5 9; Aroania village, 6 ¢ 3 9; Mt. Aroänia, above Kalávrita, 3 ¢; Páos, 1 9;

Vlasia, 1 g 2 9; Ilia: Lekhaina, 3 g 1 9; Kallithéa, 6 Z 3 9; Bassae, ruins, 3 ¢

49; Rovia, 19 (all CW). Italy: Irsima (Matera), 21.vii.1957, Eupholidoptera

danconai n.sp. La Greca det., 1 3 paratype, 1 9 allotype (IBA).

Diagnosis. — Differs from type-species as follows. ¢: Styli of subgenital plate of

variable length (figs. 107—111). Epiphallus very slender, extending apical parts

close together, in lateral view not or scarcely recurved, almost as long as length of

fused part, which is slender with plate-like lateral expansions but slightly

developed, as in figs. 173—180, 184, 185.

Variation. — Width of emargination of the hind margin of the last abdominal

tergite in the male varies, as in figs. 21—27. Length, incurvation and shape of the

basal tooth of the male cercus as in figs. 55—63. One or two spines on the apical

margin of the lobes of the male subgenital plate, as in figs. 96— 102. Length of the

styli quite variable, sometimes very long, reaching two thirds length of cercus, as in

figs. 107—111. Also the epiphallus varies considerably in length, the shortest one

about half as long as the longest (figs. 173—180, 184, 185). The coloration usually

resembles that of the type-species. Several specimens from some localities

(Kambos, Melidhoni, Neápolis), however, differ in the degree of reduction of the

black pattern of the pronotum and hind femur. The populations from Mt. Ktenias

and Mt. Parnon differ even more conspicuously from the remainder: general

colour yellow to orange brown and bright malachite green instead of dirty brown

and dark green; pronotum without any black, lower margin bordered castaneous

brown; elytra brown; hind femur with few apical streaks and tip black.

Differential diagnosis. — The male of this species may be distinguished from

others by its combination of resemblance with the type-species and the

characteristic epiphallus. The length of the styli seems to be no reliable character.

The female is indistinguishable from that of the type-species.

Remarks. — The large amount of variation has been given special attention in

regard to the existence of subspecies. However, these could not be established.

Instead it came out that E. danconai La Greca, 1959 (fig. 176), known from the

extreme southeastern part of Italy, agrees fully with some samples before me from

diverse localities assigned to E. megastyla. Therefore I propose to consider E.

danconai a synonym of E. megastyla.

Hitherto E. megastyla was known only from the type-species from the island of

Zakinthos and scanty material recorded from the Peloponnisos (Kaltenbach, 1967;

Harz, 1969; Willemse, 1971) and Mt. Olimbos (Willemse, 1974). The Parnass

record (Brunner von Wattenwyl, 1882, under E. chabrieri) refers to a single female,

which I have now at hand and is indistinguishable from the female of E. chabrieri.

Whether this locality refers to the presently named Mt. Parnassos (= Parnassus) or

to Mt. Parnis (=Parness), NE of Athinai is an open question. On account of the

ranges of E. chabrieri and E. megastyla the specimen is now assigned to the latter.

Werner’s records of E. smyrnensis (= chabrieri auct.) from the islands of Skopelos

(1933a) and Skiathos (1938) refer to E. megastyla rather than to E. smyrnensis (these

localities are omitted on the map).

Distribution. — The range of E. megastyla as now understood (map) covers the

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

lonian island of Zákinthos, the Pelopónnisos, southern continental Greece

including Evvoia, S.W. Makedhonia, and extreme S.E. Ipiros, extending into S.E.

Italy. In Arta E. megastyla occurs very close to E. chabrieri, but sympatric

occurrence could not be established.

Figs. 161—172. Epiphallus in Eupholidoptera species, anterior and left lateral views, of: 161—162, E.

chabrieri garganica La Greca, Gazätika, Kérkira; 163—164, E. chabrieri bimucronata (Ramme), Manda-

nica, Messina; 165—168, E. leucasi sp. n., paratypes; 169—170, E. epirotica (Ramme), holotype:

171—172, E. palaestinensis (Ramme), Tiberias, Israel

WILLEMSE: Eupholidoptera of Greece 59

Eupholidoptera smyrnensis (Brunner von Wattenwyl, 1882)

(figs. 28—29, 64—65, 103—105, 181—183)

Thamnotrizon smyrnensis Brunner von Wattenwyl, 1882: 336; Werner, 1901: 293.

Pholidoptera smyrnensis; Berland & Chopard, 1922: 168; Burr, Campbell & Uvarov, 1923: 124, 140, 152.

Pholidoptera chabrieri; Ramme, 1927: 128; Werner, 1933a: 407 (?); 1933b: 194; 1937b: 109; 1938: 169 (?).

Eupholidoptera smyrnensis; Ramme, 1951: 198, 204, figs. 44—46, 48, 50; Kaltenbach, 1965; 473; 1967:

195; Harz, 1969: 373, figs. 1129, 1151—1154 (? Skopelos; not Karpathos & Ikaria).

Pholidoptera festae Giglio Tos, 1914: 5. Syn. nov.

Pachytrachelurus festae Giglio Tos, 1914: 6; Ramme, 1930: 809; Harz, 1969: 373.

Pholidoptera chabrieri festae; Ramme, 1930: 809; Jannone, 1936: 135; Salfi, 1937: 3.

Eupholidoptera smyrnensis festae; Ramme, 1951: figs. 44, 50; Kaltenbach, 1967: 192.

Material studied. — Makedhonia: Khalkidhiki: Yerakini, 1 ¢; Drama: Adhriani,

19 (both CW); Kavälla: Thasos, Panayia, 17.vii.1963, S. Daan & V. v. Laar, 13 19

(ITZ). Dhitiki Thráki: Évros: Tikheròn, 18 39; Amórion, 1g; Souflion, 14 (all

CW). Samos: Koútsi, 25—27.vi. & Iraion, 13.vi. & Pedhiás Valmaris, 29.vi.1977,

M.C. & G. Kruseman, 54 29 (all ITZ).

Diagnosis. — Differs from type-species as follows. 3: Median emargination of

hind margin of last abdominal tergite much wider, lobes acute (figs. 28—29).

Cercus shorter (figs. 64—65). Pair of short spines of equal length on apical margin

of lobes of subgenital plate (figs. 103105). Epiphallus with extending apical parts

from slightly convergent to distinctly divergent, in lateral view moderately

recurved; fused part of variable width, plate-like lateral expansions well developed

(figs. 181—183). Pronotal lateral lobe usually solid black with well delimited

yellowish fascia bordering lower margin. Elytron less solid black. Black markings

of hind femur usually confined to upper half.

Q: Lobes of subgenital plate triangular, apex more acute. Coloration as in male.

Variation. — Lobes of the last abdominal tergite of the male range from pointing

ventrad to ventro-laterad (figs. 23—29). Variation of epiphallus is considerable,

already noted by Ramme (1951). Some extreme examples are figured here (figs.

181—183).

Differential diagnosis. — The species comes near the type-species but may

readily be distinguished by the very wide median emargination of the hind margin

of the last abdominal tergite in the male. The distinction of the female, however, is

more difficult. The form of the lobes of the subgenital plate and the coloration

may be helpful.

Remarks. — The subspecific distinction of the Ródhos populations under the

name E. smyrnensis festae (Giglio Tos, 1914) seems not justified. The characters

noted by Ramme (1930; 1951: figs. 44, 50) are widely overlapped by the variation

of the material before me. I propose to merge the taxon in E. smyrnensis.

The species has been recorded from numerous localities in Greece. Most of

these records seem correct, some however are doubtful and a few incorrect.

Reliable records are indicated on the map and are summarized here: Makedhonia

— Vassilika (Berland & Chopard, 1922); Lembet, Ak Bunar, Aivatli, Naresh and

Hortiack plateau (Burr, Campbell & Uvarov, 1923); Kalamika (Kaltenbach, 1967);

Aegean Is — Limnos (=Lemnos), Kastron (Werner, 1937b; Kaltenbach, 1967);

Lésvos (=Mytilene) (Werner, 1901; Berland & Chopard, 1922; Ramme, 1927;

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

Werner, 1933a?); Samos, Marathókambos (Werner, 1933b, 1937b; Kaltenbach,

1967); Nisiros (Ramme, 1927); Ródhos (Giglio Tos, 1914; Ramme, 1927, 1930;

Salfi, 1937), Villanova (Jannone, 1936), Rodhini (Kaltenbach, 1967). Other records

of E. smyrnensis (or E. chabrieri auct.), however, are omitted or referred to other

species, viz., Kithira (Werner, 1937a) referred now to E. spinigera; Kumani, the

Peloponnisos (Ramme, 1927) omitted, being far out of the range of E. smyrnensis

and presumably based on a misidentified female of E. megastyla; Skopelos

(Werner, 1933a) omitted, referring to a juvenile female of which identification is

considered unreliable; Skiathos (Werner, 1938) omitted as E. megastyla rather

than E. smyrnensis occurs in that island; Yiannitsà (= Yenidjé-Vardar), Pella distr.,

(Berland & Chopard, 1922) omitted, as the record refers to a female and its

identification needs confirmation, because all E. smyrnensis, E. megastyla and

E. chabrieri may occur in this area; Ikaria (Werner, 1933b) discussed now under

E. ikariensis; Karpathos (Werner, 1936) assigned now to Uvarovistia (Karabagia)

uvarovi.

Distribution. — The range extends from central Greek Makedhonia, via Dhitiki

Thraki, S. Bulgaria into Anatolia and some eastern Aegean islands (map).

Eupholidoptera prasina (Brunner von Wattenwyl, 1882)

(pl. 1 fig. 6, pl. 2 fig. 5; figs. 30—31, 66—67, 112—113, 189— 190)

Thamnotrizon prasinus Brunner von Wattenwyl, 1882: 337; Werner, 1901: 293.

Pholidoptera prasina; Ramme, 1930: 816, figs. 5, 6, 8, 9; Werner, 1933b: 194; 1934: 326; 1937b: 109.

Eupholidoptera prasina; Ramme, 1951: 198, 203, 206, 211, figs. 49, 51; Harz, 1969: 376, figs. 1113, 1115,

1157, 1158.

Material studied. — ¢ holotype, labelled: Smyrna, Coll. Br. v. W. Br. v. W. leg.,

prasinus m. det. Br. v. W., 5942, Type; Samos: Marathókambos, v. Oertzen,

Pholidoptera prasina Br. Ramme det., 13; Khios: 30.vi.1936, Werner, Pholidoptera

prasina Br. coll. Werner, 24 (all NMW).

Diagnosis. — 4, pl. 1 fig. 6, pl. 2 fig. 5. Pronotal dorsum smoothly rounded

laterally. Elytron reaching hind margin of first abdominal tergite. Hind femur

comparatively short. Last abdominal tergite strongly downcurved, hind margin

with moderately deep and wide median emargination, lobes acute (figs. 30—31).

Cercus without tooth, slender, proximal half very slightly incurved, in the middle

of length abruptly tapering into distal half which is straight, apex obtusely pointed

(figs. 66—67). Subgenital plate divided into small basal triangular part and pair of

elongate lobes; lobes strongly extending and narrowing distad, terminating in

needle-like spine, ventral surface with longitudinal ridge with stylus on its distal

end, located on base of terminal spine; length of stylus and spine about equal (figs.

112—113). Epiphallus small, fused part narrow, short and cylindrical, extending

apical parts widening laterally, in antero-posterior view roughly circular to heart-

shaped, fused basally, incised apically, as in figs. 189— 190.

General colour yellowish brown to pale green. Face with some symmetrically

arranged black points. Hind edge of occiput black. Black stripes from upper

margin of eye distad. Pronotum with black spot in dorso-posterior part of lateral

WILLEMSE: Eupholidoptera of Greece 61

lobe, not reaching hind margin. Middle of elytron and first abdominal tergite

black. Last abdominal tergite completely black. Fore and middle legs with few

black markings, hind femur near the knee and often dorsally in the middle of

length black.

Q. Not at hand. Reference is made to the description by Ramme (1930).

Variation. — Among the scanty material before me, the variation of the

epiphallus is worth mentioning (figs. 189—190).

Differential diagnosis. — The species is well defined. It comes near E. icariensis

(pl. 2 figs. 7, 8), which differs in the form of the epiphallus, and E. spinigera. The

latter has the styli located more proximad, the whole upper part of the pronotal

lateral lobe is black and the black pattern of the hind femur is different,

resembling that of the type-species.

Remarks. — Previous records of the species are confirmed by the material at

hand except for the specimen from Psérimos (=Kappari) (Ramme, 1930). This

material could not be traced and the locality is omitted on the map.

Distribution. — The range covers W. Anatolia, extending into the neighbouring

Aegean islands of Samos and Khios.

Eupholidoptera spinigera (Ramme, 1930)

Pholidoptera spinigera Ramme, 1930: 818, figs. 5, 6, 8, 9.

Pholidoptera chabrieri; Werner, 1937a: 147 (?).

Eupholidoptera spinigera; Ramme, 1951: 198, 204, 207, 209, 211, figs. 49; Harz, 1969: 376, figs. 1114,

1128, 1155, 1156.

Diagnosis. — Reference is made to the original description.

Differential diagnosis. — The rounded pronotal dorsum and the male subgenital

plate show the species to be closely related to E. prasina and E. icariensis. It is

distinguished from these two by the emargination of the last abdominal tergite and

the location of the styli in the male and the coloration of the pronotal lateral lobe

and hind femur.

Remarks. — The species is known only from the ¢ holo- and 9 allotype. The

epiphallus was not figured, being lost (Ramme, 1951: 209). Werner’s record

(1937a) of E. smyrnensis (= chabrieri auct.) refers to a juvenile pair from the island

of Kithira (= Cerigo), the type-locality of E. prasina. This material more probably

represents either E. spinigera or E. megastyla.

Distribution. — Known only from the island of Kithira.

Eupholidoptera icariensis sp.n.

(pl. 2 figs. 7—8; figs. 32, 68, 114—116, 186)

Pholidoptera chabrieri; Werner, 1933b: 194 (2).

Material studied. — ¢ holo-, 9 allotype, 19 paratype, labelled: Hellas Ikaria

Therma 14/21.vi.1977, M. C. & G. Kruseman; additional paratypes: Ikaria,

Thérma Loútra, 16.vi.1963 & Ayios Kirikós, 17.vi.1963, S. Daan & V. v. Laar, 49 |

juv. Q (all ITZ).

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

Figs. 173—185. Epiphallus in Eupholidoptera species, anterior and left lateral views, of: 173—180, E. me-

gastyla (Ramme): (173) Litokhoron, Piéria; (174) Akhladhókambos, Argolis; (175) Neápolis, Kozani;

(176) Irsina, Matera (paratype of E. dancomai La Greca); (177) Melidhoni, Fthiótis: (178—180) Ramia-

Livadhion, Arta; 181—183, E. smyrnensis (Brunner von Wattenwyl): (181—182) Samos; (183) Souflión,

Evros; 184—185, E. megastyla(Ramme), Mt. Panakhaikön, Akhaia

Diagnosis. — Differs from E. prasina as follows: &, pl. 2 fig. 7. Hind margin of

last abdominal tergite narrowly and not deeply incised medially (fig. 32). Cercus

slightly longer (fig. 68), subgenital plate as in figs. 114—115. Epiphallus small,

fused part cylindrical without plate-like lateral expansions, extending apical parts

robust and V-like diverging apically, as in fig. 186. Hind femur with few transverse

dorso-basal stripes, a dorsal spot halfway its length and black apex.

Q, pl. 2 fig. 8. Last abdominal sternite slightly projecting in the middle.

WILLEMSE: Eupholidoptera of Greece 63

Subgenital plate slightly wider than long, divided in a pair of lobes along median

sulcus in proximal half; lobes with hind margin widely rounded laterally,

concavely emarginate medially (fig. 116). Coloration as in male.

Measurements (length in mm): body ¢ 23.0, 9 22.0 — 23.0; pronotum Z 10.3,

Q 9.7 — 10.2; hind femur ¢ 21.6, 9 21.0 — 23.0; ovipositor 15.0 — 19.0.

Variation. — Among the scanty available material the coloration of one female

from A. Kirikós is worth mentioning. In this specimen the black spot of the

pronotal lateral lobe is larger, extends dorsad over the pronotal dorsum and an

additional pair of black spots is present on the prozona of the pronotal dorsum.

Differential diagnosis. — The species differs from E. prasina mainly in the form

of the epiphallus and from E. spinigera in the location of the styli and the

coloration of the pronotum and hind femur. Besides, the species is closely related

to E. krueperi (Ramme) and E. unimacula Karabag. The former species (Ramme,

1930: 819, figs.; 1951: fig.) differs clearly in the epiphallus, the female subgenital

plate and the coloration of the pronotal lateral lobe and furthermore in the male

cercus, subgenital plate and last abdominal tergite. The latter species (Karabag,

1956: 13, figs.) has vestigial styli and slightly different form of the epiphallus and

female subgenital plate.

Remarks. — The only records of a species of the genus from Ikaria is that in

Werner (1933b). It refers to a juvenile female, assigned to E. smyrnensis (=

chabrieri auct.). That identification however is considered unreliable and Werner’s

record is referred here to E. icariensis.

Distribution. — Known only from the type-series from the Aegean island of

Ikaria.

Eupholidoptera astyla (Ramme, 1927)

(figs. 33, 69, 117—118, 187)

Pholidoptera astyla Ramme, 1927: 133, 196, 198, figs. 11d, 13, 14; 1930: 799, 821, figs. S—7, 9; 1939:

94—96, fig. 27.

Eupholidoptera astyla; Ramme, 1951: 198, 203, 206, 209, 211, fig. 51; Harz, 1969: 362, 377, figs. 1109,

1130, 1159—1161; Willemse & Kruseman, 1976: 130, 131, figs. 1—5.

Material studied. — Kriti (?): Skonef, 23.vi, 14 (CW).

Diagnosis. — Reference is made to the descriptions in Ramme (1927, 1930,

1939).

Differential diagnosis. — The species is well defined by quite a number of

characters, cf. figs. 33, 69, 117—118, 187. An asymmetrical epiphallus is found also

in Uvarovistia (Karabagia) uvarovi.

Remarks. — Hitherto the species was known only from the original material.

The male at hand agrees fully with the original descriptions and figures. The styli

of the subgenital plate are not completely reduced, but minute, scale-like,

seemingly immovable and located pre-apically on the ventral surface of the plate

(figs. 117—118).

Distribution. — The species is known from the island of Naxos, the Kikladhes,

and Kriti. The locality of the specimen before me could not be traced.

64 TIDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

Figs. 186—190. Epiphallus, anterior and left lateral views, of: 186, Eupholidoptera icariensis sp. n., holo-

type: 187, E. astyla (Ramme), Skonef; 188, Uvarovistia (Karabagia) uvarovi (Karabag), Mt. Lästros, Kär-

pathos; 189— 190, E. prasina (Brunner von Wattenwvl), (189) holotype, (190) Khios

Eupholidoptera cretica Ramme, 1951

Eupholidoptera cretica Ramme, 1951: 198, 202, 203, 211. figs. 47, Sl; Harz, 1969: 362, 377, figs.

11111112; Willemse & Kruseman, 1976: 130, 131, figs. 6—9.

Diagnosis. — Reference is made to the original description.

Differential diagnosis. — The species is readily distinguished by a number of

characters (see key).

Remarks. — Hitherto the species is known only from the male holotype.

Distribution. — The type is from Samariä, western Kriti.

Eupholidoptera forcipata Willemse & Kruseman, 1976

Eupholidoptera latens Willemse & Kruseman, 1976

Eupholidoptera pallipes Willemse & Kruseman. 1976

Eupholidoptera gemellata Willemse & Kruseman. 1976

WILLEMSE: Eupholidoptera of Greece 65

Reference is made to the original descriptions and discussions of these species

(Willemse & Kruseman, 1976: 130—137, pls. 2—8 figs. 7—41). They are well

defined and all from Kriti.

Uvarovistia Ramme, 1951

Uvarovistia Ramme, 1951: 195, 197, 212.

Type-species: Olynthoscelis zebra Uvarov, 1916.

Diagnosis. — Reference is made to the original description.

Differential diagnosis. — The genus is close to Eupholidoptera and may be

distinguished from the latter by the coloration of the distal abdominal tergites and

the ovipositor. In Uvarovistia the last abdominal tergite is partly or completely

black and the colour extends over the lateral parts of the penultimate tergite; the

ovipositor is comparatively more robust, shorter and wider in dorsal view, i.e. less

compressed laterally.

Remarks. — Ramme erected the genus on the basis of a well defined group of

four species. He studied the epiphalli of three species and found their forms to

resemble each other remarkably.

Kaltenbach (1967) arranged Eupholidoptera uvarovi Karabag under Uvarovistia on

account of the black coloration of the distal abdominal tergites. Other characters

of this species, however, agree with some species of Eupholidoptera rather than

with members of Uvarovistia. Presumably for these reasons Harz (1969) erected for

U. uvarovi the subgenus Karabagia, but he omitted to point out the distinction

between this new subgenus and the nominate taxon. As remarked under

Eupholidoptera, however, a review of supra-specific groupings in the tribe Pholi-

dopterini can best be postponed for the time being.

Distribution. — The range of the nominate subgenus covers Iran, Iraq and

Armenia, that of monotypic Uvarovistia (Karabagia) the Aegean Island of

Karpathos.

Uvarovistia (Karabagia) uvarovi (Karabag, 1952)

(pl. 2 figs. 9—10; figs. 70, 106, 188)

Pholidoptera chabrieri; Werner, 1936: 12.

Pholidoptera (Eupholidoptera) uvarovi Karabag, 1952: 135, figs. 1—5.

Uvarovistia uvarovi; Kaltenbach, 1967: 197.

Uvarovistia (Karabagia) uvarovi; Harz, 1969: 384, figs. 653, 654, 1182—1186.

Material studied. — Karpathos: Chamili-Sattel bei Volada, 15—18.vi.1935 &

Lastros Gebirge, 15.vi.1935, O. Wettstein, Pholidoptera chabrieri Charp. det.

Werner, 13 29 (NMW).

Diagnosis. — Reference is made to the original description.

Variation. — Insufficiently known.

Differential diagnosis. — From the species of nominate Uvarovistia this species

is quite distinct in the asymmetrical epiphallus (fig. 188), the very short male

cercus (fig. 70), the slender ovipositor and the well defined yellow fascia bordering

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980

throughout the lower margin of the pronotal lateral lobe (pl. 2. fig. 9). From

Eupholidoptera it differs in the black coloration of the lateral parts of the

penultimate abdominal tergite.

Remarks. — The species seems to link nominate Uvarovistia with Eupholidoptera.

An asymmetrical epiphallus is found also in E. astyla, the male subgenital plate

(fig. 106) is much the same as in E. smyrnensis, the emargination of the hind margin

of the last abdominal tergite in the male and the coloration of the pronotum and

hind femur are found in several species of Eupholidoptera, and the slender

ovipositor even in all species of that genus,

The material at hand refers to that recorded by Kaltenbach (1967) and by

Werner (1936).

Distribution. — Known only from the island of Karpathos, the Dhodhekanisos

(map).

REFERENCES

Adamovié, 2., 1972. Drei neue Unterarten von Eupholidoptera chabrieri aus Jugoslavia (Tettigoniidae,

Orthoptera). — Glasn. Muz. Beogr. (B) 27: 193—200, figs.

Berland, L., & L. Chopard, 1922. Travaux scientifiques de l'Armée d'Orient (1916-1918). Orthoptères.

— Bull. Mus. natn. Hist. nat. Paris 1922: 166—170, 230—235, figs.

Brunner von Wattenwyl, C., 1861. Disquisitiones orthopterologicae. Dissertatio II. Nonnulla Orthopte-

ra europaea nova vel minus cognita. — Verh. zool.-bot. Ver. Wien 11: 285—310, pls.

———., 1882. Prodromus der Europäischen Orthopteren: i—xxxii, 1—466, pls., map. — Engelmann,

Leipzig.

Burr, M., B. Campbell & B. Uvarov, 1923. A contribution to our knowledge of the Orthoptera of Ma-

cedonia. — Trans. ent. Soc. Lond. 1923: 110— 169, figs., map.

Cejchan, A., 1963. Ergebnisse der Albanien-Expedition 1961 des Deutschen Entomologischen Institu-

tes. 10, Beitrag. Saltatoria. — Beitr. Ent. 13: 761— 796.

Charpentier, T. de, 1825. Horae entomologicae, adjectis tabulis novem coloratis: i—xvi, 1—255 +5, pls.

— Gosohorsky, Wratislava.

Chopard, L., 1951, Orthopteroides. Faune de France, 56: 1—359, figs. — Lechevalier, Paris.

Costa, A., 1863. Nuovi studii sulla entomologia della Calabria Ulteriore. — Atti Accad. Sci. fis. mat.

Napoli 1: 1—80, pls.

Fieber, X., 1861. Beiträge zur Orthopteren-Kenntniss. I. — Wien. ent. Monatschr, 5: 193—198.

Giglio-Tos E., 1914. Escursioni zoologiche del Dr. Enrico Festa nell’Isola di Rodi. IX. Dermaptera et

Orthoptera. — Boll. Musei Zool. Anat. comp. R. Univ. Torino 29 (680): 1—7.

Harz, K., 1969. Die Orthopteren Europas. Vol. 1. Series entomologica 5: i—xx, 1— 749, figs. — Junk,

The Hague.

Jannone, G., 1936. Nuovi contributi alla conoscenza della fauna dell isole italiane dell’Egeo. V. — Boll.

Lab. Zool. gen. agr. R. Scuola Agric, Portici 29: 47—248, figs.

Karabag, T., 1952. A new species of Pholidoptera (Orthoptera, Tettigoniidae) from the Dodecanese Is-

lands. — Istanb, Univ. Fen. Fak. Mecm. (B) 17: 135—137, figs.

———., 1956. Some new and less known Tettigoniidae (Orthoptera) from Turkey. — Communs Fac.

Sci. Univ. Ankara (C) 5:1—19, figs.

Kaltenbach, A., 1965. Dictyoptera und Orthopteroidea von Nordost-Griechenland und der Insel Tha-

sos, — Annin naturh. Mus. Wien 68: 465—484, figs.

——., 1967. Mantodea und Saltatoria aus Griechenland. — Annin naturh. Mus. Wien 70: 183—199,

he.

La Greca, M., 1959, L’Ortotterofauna Pugliese ed il suo significato biogeografico. — Memorie Bioge-

ogr. adriat. 4: 33— 170, figs.

Ramme, W., 1927. Die Dermapteren und Orthopteren Siziliens und Kreta. — Eos. Madr. 3: 111—200,

figs.

WILLEMSE: Eupholidoptera of Greece 67

———, 1930. Revisionen und Neubeschreibungen in der Gattung Pholidoptera Wesm. (Orth., Tetti-

gon.). — Mitt. zool. Mus. Berl. 16: 798—821, figs.

——, 1939. Beiträge zur Kenntnis der palaearktischen Orthopterenfauna (Tettig. et Acrid.). — Mitt.

zool. Mus. Berl. 24: 41—150, figs.

—— , 1951. Zur Systematik, Faunistik und Biologie der Orthopteren von Südost-Europa und Vorder-

asien. — Mitt. zool. Mus. Berl. 27 (1950): 1—431, figs.

Salfi, M., 1937. Ortotteri di Valona (Albania) e di Rodi (Egeo). — Annuar. Mus. zool. Univ. Napoli

(N.S.) 7: 1—6, figs.

Targioni-Tozzetti, A., 1881. Orthopterorum Italiae species novae in collectione R. Musei Florentini

digestae. — Boll. Soc. ent. ital. 13: 180—186.

Werner, F., 1901. Die Dermapteren- und Orthopterenfauna Kleinasiens. — Sber. Akad. Wiss. Wien (1)

110: 259—306, figs.

—_, 1933a. Ueber Orthopteren aus Ost-Griechenland und von den Inseln des Aegaeischen Meeres.

— Mitt. zool. Mus. Berl. 18: 395—415, figs.

——, 1933b. Ergebnisse einer zoologischen Studien- und Sammelreise nach den Inseln des Ägäi-

schen Meeres. II. Orthopteren. — Sber. Akad. Wiss. Wien (1) 142: 185— 204.

——, 1934. Dritter Beitrag zur Kenntnis der Tierwelt der Ägäischen Inseln. — Sber. Akad. Wiss.

Wien (1) 143: 313— 337, figs.

—_, 1936. Ergebnisse einer zoologischen Forschungsreise nach dem Dodekanes, unternommen

von Kustos Dr. Otto Wettstein. I. Orthopteren aus dem Dodekanes. — Sber. Akad. Wiss.

Wien (1) 145: 9—16.

——, 1937a. Beiträge zur Kenntnis der Tierwelt des Peloponnes, der Inseln Kythira und Euboea so-

wie der kleinen Inseln im Saronischen Golf. — Sber. Akad. Wiss. Wien (1) 146: 135—153, figs.

——, 1937b. Ergebnisse der vierten zoologischen Forschungsreise in die Ägäis (1936). — Sber.

Akad. Wiss. Wien (1) 146: 89—118, figs.

—, 1938. Ergebnisse der achten zoologischen Forschungsreise nach Griechenland (Euboea, Tinos,

Skiathos, Thasos usw.). — Sber. Akad. Wiss. Wien (1) 147: 157—173, figs.

Willemse, F., 1971. A new species of Conocephalus, subgen. Xiphidium, from Greece (Orthoptera, Ensi-

fera, Conocephalinae). — Publties natuurh. Genoot. Limburg 20 (1970): 15—17, figs.

———, 1974. A new subspecies of Paranocaracris bulgaricus (Ebn. & Dren.) from Greece (Orthoptera,

Acridoidea, Pamphaginae). — Biol. gallo-hellenica 5: 351—352, figs.

———, &G. Kruseman, 1976. Orthopteroidea of Crete. — Tijdschr. Ent. 119: 123— 164, figs.

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 2, 1980 PI.

Figs. 14, 6, Eupholidoptera species: 1—2, E. leucasi sp. n., holotype; 3—4, E. epirotica (Ramme): (3) &

holotype, (4) 9, Kerkira; 6, E. prasina (Brunner von Wattenwyl), £, Khios

Pie 2 WILLEMSE: Eupholidoptera of Greece 69

Figs. 5, 7, 8. Eupholidoptera species: 5, E. prasina (Brunner von Wattenwyl), ¢ holotype; 7—8, E. icarien-

sis sp. n.: (7) 3 holotype, (8) 9 allotype. Figs. 9—10. Uvarovistia (Karabagia) uvarovi (Karabag), 9, 3, 10,

Ik 1

DEEL 123 AFLEVERING 3 1980

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

C. A. W. JEEKEL. — A revision of the Burmese Paradoxosomatidae (Diplopoda,

Polydesmida) in the Museo Civico di Storia Naturale at Genova (Part III),

p. 71—88, fig. 1—21.

Tijdschrift voor Entomologie, deel 123, afl. 3 Gepubliceerd 29-11-1980

A REVISION OF THE BURMESE PARADOXOSOMATIDAE

(DIPLOPODA, POLYDESMIDA) IN THE MUSEO CIVICO

DI STORIA NATURALE AT GENOA (PART III)')

C. A. W. JEEKEL

Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Amsterdam

With 21 text-figures

ABSTRACT

The genus Antheromorpha Jeekel, 1968, is diagnosed and its relationship with other genera in the tribe

Orthomorphini is discussed. Redescriptions of a number of its species from Burma originally described

by Pocock, 1895, under the names of Orthomorpha miranda, O. melanopleuris, O. bistriata, O. bivittata,

O. comotti and O. pardalis are given and their relationship is discussed. The distributional pattern of the

species of the group of genera to which Antheromorpha belongs, and the relative abundance of them in

Burma, Borneo and Java, seems to indicate a dispersal route which bypassed the Malay Peninsula and

Sumatra.

INTRODUCTION

The third and last part of the revision of the Pocock types of Burmese

Paradoxosomatidae in the Genoa Museum deals with the genus Antheromorpha

Jeekel, 1968, and concerns the species which Pocock (1895) described under the

names of Orthomorpha miranda, O.melanopleuris, O. bistriata, O. bivittata,

O. comotti, and O. pardalis. Unfortunately these species, with the exception of

O. miranda and O. bistriata, were based on female specimens only, which renders

an evaluation of their systematic status almost impossible. In spite of this it

appeared worthwhile to extend the original descriptions of O. melanopleuris,

O. bivittata, O.comotti, and O.pardalis too, in order to facilitate future

identifications.

DESCRIPTIONS

Antheromorpha Jeekel

Brachytropis Silvestri, 1896: 812 (preoccupied).

Antheromorpha Jeekel, 1968: 149.

!) Based on data accumulated through the aid of grant I 954—36 from the Netherlands Organisation

for the Advancement of Pure Research (Z.W.O.) and the Italian National Council of Research

(C.N.R.).

Parts I and II of this series appeared in Tijdschr. Ent. 108: 95—144, and Tijdschr. Ent. 118:

289— 301.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

seneric diagnosis. — Medium-sized Orthomorphini with moderately or well-

eveloped paranota having distinct marginal calluses. Colour pattern usually

conspicuous, often with pale paramedian stripes in addition to the pale paranotal

margins. Pleural keels well developed in anterior somites, disappearing gradually

towards the middle or posterior somites. Sternite of Sth somite of male with a

process between the anterior legs. Legs of male without particular modifications

except the usual scopulae on tibiae and tarsi, which extend caudad to the legs of

the second half of the body before completely disappearing. Gonopods with coxa

straight, of medium size. Prefemur only slightly elongate. Femorite well-

developed, somewhat widening distad. Spermal channel running along the medial

side of the femorite. No distinct laminate crest at the medial side of the base of the

femorite, and in general no sharp demarcation between the femoral and

postfemoral sections. Tibiotarsus and solenomerite relatively short. The

tibiotarsus curving rather strongly caudad, consisting of a well-developed lamina

medialis and lamina lateralis. The lamina medialis about halfway bearing a well

developed prong. Apex of tibiotarsus bilobate.

Type-species. — Orthomorpha miranda Pocock, 1895.

Antheromorpha miranda (Pocock)

(figs. 1—6)

Orthomorpha miranda Pocock, 1895: 812, fig. 13—13a.

Antheromorpha miranda; Jeekel, 1968: 57.

Material. — This species was described after an unrecorded, but apparently

rather large number of specimens from four localities. The material taken by

Oates at Rangoon and Tharrawaddy is in the British Museum (Natural History),

London, and has not been incorporated in the present study. Fea collected

material at Palon in Pegu and at Thigian, upper Irrawaddy. From these two

localities the Genoa museum has three males and one female, and one female,

respectively. Weidner (1960) has mentioned four syntypes from Palon in the

Hamburg museum. In the loan register of the Genoa museum I found evidence

that Silvestri at one time borrowed some material from Palon, which apparently

was never returned and probably is still in his collection at Portici.

Pocock originally designated Rangoon as the type locality; hence I have

labelled the material in the Genoa museum as paratypes (= paralectotypes),

pending the designation of a lectotype in the British Museum.

Palon in Pegu, coll. L. Fea, 3 3, 1 9 paratypes. Thigian, upper Irrawaddy, coll.

L. Fea, 1 9 paratype.

Description. — Colour: see Pocock.

Width: &: 3.2, 3.6, 3.7 mm; 9: 4.6, 3.6 mm.

Head and antennae: Labrum deeply and rather widely emarginate, tridentate.

Clypeus moderately convex, rather weakly impressed towards the labrum; the

lateral border weakly rounded, with a slight notch near the labrum. Headplate

rather densely to sparsely punctulate and setiferous from the clypeus up to the

JEEKEL: Burmese Paradoxosomatidae 73

Figs. I—6. Antheromorpha miranda (Pocock). 1: head and four anterior somites of ¢ paralectotype, la-

teral aspect. 2: left side of 10th and 11th somites of same, dorsal aspect. 3: the same, lateral aspect. 4:

hypoproct of same, ventral aspect. 5: right gonopod of same, medial aspect. 6: left side of 10th somite of

9 paralectotype from Palon, dorsal aspect.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

stal region, otherwise smooth; vertex hairless. Antennal sockets separated by

ae and two fifths times the diameter of a socket, or by just over two thirds of the

length of the 2nd antennomere. Postantennal groove deep and rather wide; the

wall in front moderately prominent. Vertex moderately convex, the vertigial sulcus

rather weakly impressed, running downward to about the upper level of the

sockets. Antennae of moderate length, moderately stout, distinctly clavate.

Pubescence moderate in the proximal antennomeres becoming dense in the distal

ones. Relative length of antennomeres: 2nd = 3rd > 4th > Sth = 6th; the 6th

antennomere nine tenths of the length of the 2nd.

Collum: a little wider than the head, subtrapezoidal in dorsal outline (fig. 1).

Anterior border faintly convex in the middle, a little more strongly rounded

towards the sides, and laterally straight. Posterior border widely emarginate in the

middle, becoming faintly convex more laterally; no notch or a very faint one above

the lateral rounding. Lateral border rather widely and a little asymmetrically

rounded. Surface of collum transversely alnost evenly convex, only slightly

flattened in the middle. Marginal rin narrow along the lateral border, disappearing

almost completely towards the middle of the anterior border. Surface smooth,

shiny, hairless or with a few hairs.

Somites: Constriction rather weak. Prosomites dull, somewhat silky. Stricture

narrow, distinctly demarcated from the prosomite, dorsally distinctly beaded

down to the level of the paranota, smooth below that level. Metatergites smooth,

shiny. Transverse furrow present from the Sth to the 18th somite, weakly indicated

also on the 4th and 19th somites. The furrow rather deep, rather wide, distinctly

longitudinally striate. Anterior and posterior metatergites with some hairs; the

others hairless, but in front of the furrow a transverse row of four minute, now

hairless but probably originally setiferous granules. Sides rather coarsely granulate

in the anterior somites, to finely rugulose-granulose in the posterior somites.

Pleural keels of the 2nd to the 4th somites represented by well-developed curved

ridges, which are caudally produced into an acute-angled triangular lappet which

projects behind the caudal margin. On the Sth and subsequent somites there is

only a triangular lappet at the posterior margin, which up to the 7th somite is

pointed and projects slightly behind the caudal margin of the somites. From the

9th somite onwards the lappet is obtuse and does not project, it becomes gradually

more rounded caudally in subsequent somites to gradually disappear towards the

17th somite.

Paranota: (figs. 1—3) 2nd somite wider than the collum, but narrower than the

3rd; 4th somite wider than 3rd. Paranota of 2nd somite with its anterior border

widely rounded, a little thrust forward, the latero-anterior edge somewhat obtuse,

with a weak lateral tooth. The lateral border very weakly convex, with the

indication of a tooth halfway. Posterior border practically straight, incurved at

base. Latero-posterior edge acutely angular, acuminate. Marginal rim narrow, but

sharply demarcated by a furrow. Paranota slightly declined in the lateral direction,

the caudal edges projecting well caudad of the posterior margin of the somite.

Paranota of 3rd and 4th somites with the anterior border rather widely rounded,

not shouldered at base, passing into the practically straight lateral border with an

indication of a small marginal tooth. Posterior border straight, not incurved at

JEEKEL: Burmese Paradoxosomatidae 75

base. Caudal edges acutely angular, acuminate, projecting well beyond the

margins of the somites. Paranota horizontal, their marginal rims somewhat wider

than in the 2nd somite. Paranota of 5th and subsequent somites with the latero-

anterior border weakly rounded anteriorly, becoming practically straight and

distinctly divergent laterally. Posterior border straight or faintly concave. The

caudal edges acutely angular, acuminate, becoming subspiniform in the 15th to

19th somites, projecting well behind the margin in all somites. Level of paranota

high, raised a little above the horizontal level, the caudal edges generally pointing

slightly upwards. Marginal rims rather narrow dorsoventrally, especially in the

poreless paranota. The rims also ventrally demarcated by a furrow. Pores lateral,

in an elongate oval excavation.

Sternites and legs: Sternites of middle somites as long as wide. Cross impressions

with the longitudinal furrow rather weak, the transverse furrow deeper.

Pubescence moderate. Sternal cones absent. Sternite of 5th somite with a simply

rounded conical knob between the anterior legs, occupying about two fifths of the

width between the coxae. The process is a little broader than long. Transverse

furrow distinct. Posterior part of sternite rather widely excavated. Sternite of 6th

somite deeply excavated, the transverse furrow almost obsolete, the posterior part

of the caudal half of the sternite not elevated above the ventral level of the

metasomal ring. Sternite of 7th and 8th somites not modified. Legs rather long,

moderately slender, not incrassate. Pubescence moderate becoming rather dense

in distal podomeres, equally distributed on all sides. Tarsal and tibial brushes

moderately thick in the anterior legs, thinning out gradually to become almost

obsolete in the legs of the 13th or 14th somite and totally absent in the legs of the

17th somite. Ultimate pairs of legs not modified. Relative length of

podomeres: 3rd > 6th > Sth = 2nd > 4th, the 6th podomere just over three

quarters of the length of the 3rd.

Anal somite: (fig. 4) Epiproct thick, rather long, the ventral side slightly

concave. The base rather broad, the sides moderately converging, practically

straight, the lateral setiferous tubercles rather small. The end rather narrowly and

straightly truncate, with a pair of rather widely separated slender cones directed

backward and very slightly downward. Paraprocts slightly rugulose, the setae on

weak tubercles, the marginal rims rather narrow but moderately high. Hypoproct

triangular, broad, the sides widely concave, the end obtusely rounded. Setiferous

tubercles projecting slightly outside the margin, but not equalling the caudal edge.

Gonopods: (fig. 5) Coxa of moderate size, straight, widening slightly distad, with

a latero-anterior setiferous area. Prefemur only slightly elongate, laterally

distinctly demarcated from the femorite, the demarcation transverse on the

longitudinal axis of the femorite. Femorite long, widely curved, broadening

slightly distally. Postfemoral region not demarcated. Spermal channel running

straight along the medial side of the femorite towards the base of the solenomerite.

Tibiotarsus arising from the caudal side of the distal end of the femorite, short,

rather strongly curved in caudal direction. Lamina medialis with an elongate

serrulate prong, pointing meso-distad. Apex of tibiotarsus with a sub-uncate lobe.

Solenomerite short, tapering distad, largely sheathed by the lamina medialis and

lamina lateralis.

6 PH DSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL, 3, 1980

Female: differing from the male in the following characters, Antennal sockets

separated by five sixths of the length of the 2nd antennomere. Antennae relatively

a little shorter, Collum about as wide as head, Somites more robust with the

stricture relatively a little less constricted, and the dorsum of the metatergites a

little more convex. Pleural keels of 2nd to 4th somites as in the male, those of the

Sth somite represented by a rounded lappet near the posterior margin, scarcely

projecting, From the 7th somite onwards there is only a slight ridge near the

posterior margin, which is rounded and gradually disappears towards the 16th

somite, Paranota (fig. 6) extending less laterad, with the lateral borders somewhat

more convex, Sternites as long as wide, pubescence as in male, Legs comparatively

a little shorter, rather slender, Relative length of podomeres: 3rd > 6th > 2nd > Sth

> 4th; the 6th podomere just over three quarters of the length of the 3rd. Two

ultimate pairs of legs slightly shorter than the preceding, but not modified,

Antheromorpha miranda (Pocock), var.

(figs. 7, 8)

Orthomorpha melanopleuris Pocock, 1895: 813 (pro parte),

Material, The male specimen from Minhla which Pocock referred to

Orthomorpha melanopleuris is certainly not conspecific with the female type of that

species, It may represent a geographical form of Antheromorpha miranda,

Minhla, coll, G. B. Comotto, 1 &.

Description, — Differing from A, miranda in the following particulars,

Colour: The lower side of the pleurae of the metasomites is yellowish instead of

dark,

Width: 3.3 mm.

Head and antennae: Antennal sockets separated by three fifths of the length of

the Ind antennomere.

Collum: The posterior margin has a distinct notch above the lateral rounding.

The lateral margin is somewhat more pronouncedly asymmetrically rounded,

Somites: The granulation of the pleurae is distinct only in the anterior half of the

body, and becomes more dispersed in the middle part and restricted to the lower

half of the pleurae. In the posterior part of the body the surface becomes

subgranulose. The pleural keels gradually disappear from the 9th somite onwards

to become only indicated on the 13th and very faintly indicated on the 14th somite.

Paranota: (fig. 7) The lateral borders of the paranota are less diverging caudad,

being more parallel-sided. Paranota not thicker dorso-ventrally than in 4. miranda.

Sternites and legs: Sternites one and one eighth times longer than broad.

Sternite of Sth somite with process slightly thicker than in A. miranda; caudal

portion of this sternite not modified. Sternite of 6th somite rather deeply

excavated, in the caudal half only the median part is scarcely raised above the

ventral level of the metasomal ring. The tibial and tarsal brushes of the legs are still

visible in the distal part of the tarsi of the legs of the 16th somite, are almost

entirely absent in the legs of the 17th somite and completely lacking in the legs of

the 18th somite

JEEKEL: Burmese Paradoxosomatidae 77

Figs. 7—8. Antheromorpha miranda (Pocock), var. 4 specimen from Minhla, identified by Pocock as

Orthomorpha melanopleuris Pocock. 7: left side of 10th and 11th somites, dorsal aspect. 8: right gono-

pod, medial aspect.

Anal somite: Hypoproct more elongate triangular, less trapezoidal. The

setiferous tubercles project distinctly outside the border.

Gonopods: (fig. 8) The telopodite on the whole slightly more elongate and

slightly less curved. Otherwise as in A. miranda.

Antheromorpha melanopleuris (Pocock)

(figs. 9, 10)

Orthomorpha melanopleuris Pocock, 1895: 813 (pro parte).

Antheromorpha melanopleuris; Jeekel, 1968: 57.

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

Figs. 9—10. Antheromorpha melanopleuris (Pocock), © lectotype. 9: left side of 10th and 11th somites,

dorsal aspect. 10: the same, lateral aspect.

Material. — The number of specimens upon which this species was based has

not been recorded. The collection of the Genoa museum has two female

specimens, one of which now has been labelled as lectotype. A syntype (now

paralectotype), probably female, is according to Weidner, 1960, in the Hamburg

museum; an additional female is in the British Museum, London.

The male specimen from Minhla, tentatively considered by Pocock to belong to

Orthomorpha melanopleuris, appears to be a variety of A. miranda and is described

above.

Teinzo, coll. L. Fea, 9 lectotype, 9 paralectotype.

Description. — Differing from A. miranda 9 in the following particulars.

Colour: see Pocock. The yellowish paramedian bands of the metatergites are

slightly more parallel than in 4. miranda, where they are distinctly diverging in a

caudal direction.

Width: 3.9 mm in both specimens.

Head and antennae: Clypeus moderately impressed towards the labrum; its

lateral border widely emarginate above labrum. Headplate punctulate-rugulose in

clypeal area, otherwise smooth and shiny. Vertex with two plus two hairs.

Antennal sockets separated by three quarters of the length of the 2nd

antennomere. Vertigial sulcus well impressed. 2nd to 6th antennomeres of

subequal length.

Collum: a little narrower than the head. Posterior border with a slight

emargination above the lateral rounding. Surface almost evenly convex, with some

dispersed hairs.

Somites: Stricture with a faint lateral striation down to about the level of the

stigmata, Metatergites with a faint median furrow in some somites: the transverse

furrow weakly striate. Sides in most somites smooth or with some coarse wrinkles,

finely dispersedly granulate. In 2nd to about Sth somites a more dense granulation,

in posterior half of body granulation in particular in lower half of sides. Pleural

keels of 2nd somite caudally rounded, in somites 3 and 4 acutely pointed, all

JEEKEL: Burmese Paradoxosomatidae 79

projecting caudad of posterior margin. In Sth and 6th somites the pleural keels are

represented by obtusely angular lappets near the posterior margins, which become

more rounded in the 7th somite and gradually disappear towards the 12th somite.

Paranota: (figs. 9—10) of 2nd somite with obtusely angular posterior edges.

Paranota of 3rd, 4th and Sth somites wider dorsoventrally, the posterior edges less

acute. Paranota of subsequent somites with latero-anterior border more strongly

curved, the caudal edges less acuminate. Pore area marked by a distinct

emargination. Only in the 16th and subsequent somites the caudal edges become

spiniform.

Sternites and legs: Middle sternites one and one eighth times broader than long.

Anal somite: Terminal knobs of epiproct shorter. Hypoproct broader, shorter,

trapezoidal with sides concave and end rounded. Setiferous tubercles moderately

developed, not projecting outside the margin.

Antheromorpha bistriata (Pocock)

(figs. 11—14)

Orthomorpha bistriata Pocock, 1895: 814.

Antheromorpha bistriata; Jeekel, 1968: 57.

Material. — This species was described after a single male specimen, which is

preserved in the Genoa museum and which I have labelled as holotype.

Bhamo, 1.x.1886, coll. L. Fea, ¢ holotype.

Description. — Differing from A. miranda in the following particulars.

Colour: The dark middorsal stripe is narrower than in A. miranda.

Width: 3.4 mm.

Head and antennae: Lateral border of clypeus faintly rounded, and distinctly

emarginate near the labrum. Clypeal and frontal regions punctulate, the clypeus

somewhat rugulose; vertex with two pairs of setae. Postantennal groove

moderately deep and rather wide; the wall in front rather weakly prominent.

Antennomeres decreasing in length from the 2nd to the 6th.

Collum: a little narrower than the head, subtrapezoidal or oblong reniform in

dorsal outline. Posterior border with a very weak notch above the lateral rounding.

Lateral border almost symmetrically rounded. Lateral sides of collum slightly

flaring. Surface of collum with some hairs along the anterior margin and some in

the middle.

Somites: Constriction moderate. Metatergites mostly hairless, sometimes four

hairs in front of the transverse furrow and some along the posterior margin.

Transverse furrow without distinct sculpture, sometimes faintly striate. Sides

smooth to weakly granulose, not distinctly granulate in the anterior somites.

Pleural keels of 2nd to 4th somites with the caudal lappets rectangular, pointed,

scarcely projecting caudad. In the Sth and 6th somites the caudal lappets are

obtusely angular, blunt, and do not project caudad. In the 7th somite the caudal

end is rounded, and the keels disappear gradually towards the 10th somite.

Paranota: (figs. 11—12) of 2nd somite caudally less produced, the caudal angle

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

[o |

14 \\ |

\ |

cy ee

fi]

{ef sf

Nae aH sx

È VA

a, )

Figs. 1114. Antheromorpha bistriata (Pocock), £ holotype. 11: left side of 10th and 11th somites, lat-

eral aspect. 12: the same, dorsal aspect. 13: hypoproct, ventral aspect. 14: right gonopod, medial aspect.

less acute than in A. miranda. Paranota of subsequent somites dorsoventrally

relatively a little thicker, and projecting less caudad. The lateral borders of the

paranota are more parallel, less diverging than in A. miranda.

Sternites and legs: Sternites of middle somites one and one eighth times longer

than broad. Sternite of 5th somite swollen between anterior legs and bearing a low

knob. Sternite of 6th somite rather deeply excavated, but not down to the level of

the ventral side of the metasomal ring. Tibial and tarsal brushes are dense in the

anterior legs up to the 6th somite, thinning out gradually to become very thin from

the 10th somite onwards and totally absent from the 15th somite onwards.

Anal somite: Epiproct with terminal knobs a little shorter. Hypoproct (fig. 13)

subtriangular, narrower at base than in A. miranda and with the lateral sides more

distinctly convex.

Gonopods: (fig. 14) very similar to those of A. miranda but with the femorite

notably shorter in relation to size of coxa, prefemur and tibiotarsus.

JEEKEL: Burmese Paradoxosomatidae 81

Antheromorpha bivittata (Pocock)

(figs. 15, 16)

Orthomorpha bivittata Pocock, 1895: 814.

Antheromorpha bivittata; Jeekel, 1968: 57.

Material. — The two specimens, both female, upon which this species was based

are in the Genoa museum. One has now been designated as lectotype.

Shenmaga, coll. L. Fea, 9 lectotype, 9 paralectotype.

Description. — Differing from A. miranda ® in the following particulars.

Colour: see Pocock. The dark median band is rather broad, slightly constricted

in the area of the stricture. Lower side of pleurae yellowish.

Width: lectotype 3.9 mm, paralectotype 3.5 mm.

Collum: a little wider than the head. Posterior border distinctly notched above

the lateral rounding.

Somites: Whole surface of lateral sides distinctly granulate. Pleural keels of 5th

and 6th somites represented by obtuse triangular lappets, projecting scarcely

caudad in the 5th, not projecting in the 6th somite. In the 7th somite the caudal

lappet is rounded, from the 8th somite onwards it is very weakly developed to

become obsolete in the I Ith or 12th somite.

Paranota: (figs. 15—16) Posterior edge of paranota of 2nd somite about

rectangular. Paranota laterally slightly more convex, the caudal edges slightly less

produced.

Sternites and legs: Sternites of middle somites one and one quarter times

broader than long. Cross impressions with the longitudinal furrow also distinct.

Pubescence moderate to rather dense. The two ultimate pairs of legs a little

shorter than the preceding ones. Relative length of podomeres: 3 rd > 6th > 2nd =

Sth > 4th.

Anal somite: Hypoproct broadly triangular.

Figs. 15—16. Antheromorpha bivittata (Pocock), 9 lectotype. 15: left side of 10th and 11th somites, dor-

sal aspect. 16: the same, lateral aspect.

82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

Antheromorpha pardalis (Pocock)

(figs. 17, 18)

Orthomorpha pardalis Pocock, 1895: 815.

Antheromorpha pardalis; Jeekel, 1968: 57.

Material. — According to Pocock this species was based on a unique female

specimen. This specimen is preserved in the Genoa museum (it lacks its collum),

and has been labelled as holotype.

Palon in Pegu, coll. L. Fea, 9 holotype.

Description. — Differing from A. miranda Q in the following particulars.

Colour: see Pocock.

Width: 4.1 mm.

Head and antennae: Lateral borders of clypeus widely emarginate near the

labrum. Headplate rugulose in clypeal part. Antennal sockets separated by one

and three quarters times the diameter of a socket. Vertex more convex, the sulcus

well impressed. Relative length of antennomeres: 2nd > 3rd > 4th = Sth > 6th; the

6th antennomere four fifths of the length of the 2nd.

Collum: lacking.

Somites: Metatergites with weakly leathery integument. Transverse furrow well

impressed, almost reaching the dorsal delimitation of the paranota, present from

the 2nd to the 18th somite; without distinct sculpture. A fine middorsal furrow is

visible in most metatergites. Sides moderately densely granular. Pleural keels of

2nd to 4th somites with pointed caudal lappets projecting caudad of posterior

margin. In the 5th somite a weakly developed caudal ridge which gradually

disappears in subsequent somites and is absent from the 8th somite onwards.

Paranota: (figs. 17—18) of 2nd somite with anterior border straight and lateral

border widely rounded. Latero-posterior edge obtuse. Marginal rim moderately

wide, distinctly demarcated, on all sides the furrow along the caudal margin

continued upwards to the metasomal margin. Paranota of 3rd somite rather

Figs. 17—18. Antheromorpha pardalis (Pocock), 9 holotype. 17: left side of 10th and 11th somites, dorsal

aspect. 18: left side of 11th and 12th somites, lateral aspect.

JEEKEL: Burmese Paradoxosomatidae 83

weakly and evenly convex, the posterior edge acute, the posterior border weakly

concave. Paranota of 4th somite somewhat less convex than those of 3rd. The

posterior edge slightly acutely angular. Posterior edges of 2nd to 4th somites

projecting beyond posterior margin of the somites. Caudal edge of paranota of

5th somite about rectangular, not projecting beyond posterior margin. Paranota of

6th and subsequent somites with acute edges, which become spiniform and

curving a little inward in the 14th to 19th somites, projecting caudad of posterior

margin in all these somites.

Sternites and legs: Sternites of middle somites one and one quarter times

broader than long. Cross impressions rather weak. Pubescence of legs ventrally

moderate. Relative length of podomeres: 3rd > 6th > 2nd > Sth > 4th.

Anal somite: Epiproct of moderate width and thickness. Sides slightly concave.

Near the end a pair of well-developed setiferous tubercles; the end itself with a

pair of small cones pointing backwards. Paraprocts rugulose, the setae not arising

from tubercles. Marginal rims moderately high and wide. Hypoproct

subtriangular, with the sides concave at base, widely rounded more distally, the

end obtusely angular. Setiferous tubercles weakly developed, not projecting.

Antheromorpha comotti (Pocock)

(figs. 19—21)

Orthomorpha comotti Pocock, 1895: 814.

Antheromorpha comotti; Jeekel, 1968: 57.

Material. — The type material consists of a single female specimen, which is

preserved in the Genoa museum and is now labelled as holotype.

Minhla, coll. G. B. Comotto, 1883, 9 holotype.

Description. — Differing from A. miranda Q in the following particulars.

Colour: see Pocock.

Width: 4.5 mm.

Head and antennae: Lateral border of clypeus faintly convex, widely emarginate

above the labrum. Clypeus rugulose-punctulate, remainder of headplate smooth

and shiny. Antennal sockets separated by one and five eighths times the diameter

of a socket or by about three fifths of the length of the 2nd antennomere. Vertex

with median sulcus moderately impressed and running downward to halfway

between the antennal sockets. Antennae not clavate, pubescence rather dense in

proximal antennomeres to dense in the distal ones. Relative length of

antennomeres: 2nd = 3rd > 4th = Sth > 6th; the 6th antennomere almost three

quarters of the length of the 2nd.

Collum: (fig. 19) about as wide as head. Its posterior border faintly emarginate,

convex towards the lateral sides and emarginate above the lateral rounding.

Lateral margin widely rounded, becoming narrowly rounded caudally. Surface of

collum hairless, dull, rugulose in the middle becoming sudgranulose towards the

lateral sides. Marginal rim narrow and very thin, laterally weakly defined, finely

continued along the anterior border. Lateral sides of collum slightly flaring,

caudally slightly inflated.

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

Somites: weakly constricted. The stricture rather narrow, distinctly demarcated.

Only the caudal part of the stricture finely but distinctly longitudinally ribbed

down to upper level of the paranota, below thafevel finely striate, but no striation

below the lower level of the paranota. Metatergites very dull, silky, finely rugulose

becoming subgranulose near base of iaranota. In front of the transverse furrow a

row of four minute granules. Transverse furrow moderately impressed, linear,

finely longitudinally striolate; a fine median furrow on metatergites in front and

behind the transverse furrow. Sides densely covered with minute granulae,

especially in the 2nd to 4th somites. Pleural keels represented by distinct ridges in

the 2nd to 4th somites, caudally produced into slightly acutely angular pointed

lappets, which project slightly caudad of posterior margin. In Sth to 7th somites

the pleural keels are present only as rounded ridges near the posterior margin of

the somites, in the 8th, 9th and 10th somites these ridges gradually disappear.

Paranota: (figs. 19—21) Lateral border of paranota of 2nd somite widely

rounded; latero-posterior edge obtusely angular and projecting slightly caudad of

margin. Posterior border widely rounded, incurved at base. Marginal rim thin,

laterally rather weakly defined. Paranota of 3rd somite with the caudal edge

obtusely angular and slightly projecting. The marginal rim rather thick. Posterior

border somewhat convex, slightly incurved at base. Paranota of 4th somite with

lateral border slightly more widely rounded than in the 3rd somite, the marginal

Figs. 19—21. Antheromorpha comotti (Pocock), 2 holotype. 19: left side of head and three anterior so-

mites, lateral aspect. 20: left side of 10th and I Ith somites, lateral aspect. 21: the same, dorsal aspect.

JEEKEL: Burmese Paradoxosomatidae 85

rim thicker than in the 3rd somite. Posterior border straight or faintly concave, the

latero-posterior edge about rectanrular, slightly projecting caudad of posterior

margin. Paranota of 5th and subsequent somites notably different from those of

the other species treated in this paper. Their lateral border faintly convex; the

posterior edges acutely angular, pointed, becoming spiniform in the 14th to 19th

somites.

Sternites and legs: Sternites with cross impressions well-developed, the

transverse furrow a little deeper than the longitudinal one. Pubescence of all

sternites rather dense. Legs of moderate length, rather slender. Ventral

pubescence of podomeres and dorsal pubescence of distal end of tibiae and the

entire tarsi rather dense; dorsal pubescence of proximal podomeres moderate.

Last pairs of legs lacking in type specimen. Relative length of podomeres: 3rd >

6th > 2nd > Sth > 4th; the 6th podomere almost four fifths of the length of the 3rd.

Anal somite: Epiproct of moderate length, width and thickness. The lateral sides

converging concavely, becoming almost parallel-sided near the apex. Lateral

setiferous tubercles very distinct. The end with two rather small, rather widely

separated rounded knobs. Paraprocts rugulose, subgranulose. The marginal rims

moderately wide, rather low. Setiferous tubercles small and weakly prominent.

Hypoproct triangular to subtrapezoidal. The sides concave, the end rounded. The

setiferous tubercles rather large but weakly prominent and not projecting outside

the margin.

CLASSIFICATION

The name Antheromorpha, proposed in 1968 as a substitute for the almost totally

forgotten but preoccupied name Brachytropis Silvestri, was introduced as a generic

name for a rather large group of species occurring in Burma, Siam and Indochina

formerly referred to Orthomorp a Bollman, 1893, in an attempt to eliminate the

heterogeneity in the last genus. To Antheromorpha the following species were

assigned tentatively (Jeekel, 1968: 57):

. bistriata (Pocock, 1895) — Burma

. bivittata (Pocock, 1895) — Burma

. comotti (Pocock, 1895) — Burma

. festiva (Brölemann, 1896) — Siam

. harpaga (Attems, 1937) — Indochina

. mediovirgata (Carl, 1941) — Burma

melanopleuris (Pocock, 1895) — Burma

. minlana (Pocock, 1895) — Burma

. miranda (Pocock, 1895) — Burma

. orophila (Carl, 1941) — Burma

. pardalis (Pocock, 1895) — Burma

. uncinata (Attems, 1931) — Siam

In this concept the genus is a fairly large one, and it seems appropriate,

therefore, to discuss the interrelationship of the species, a number of which have

been redescribed in the previous pages of this paper.

Closely related to the type-species of the genus, A. miranda, are A. bistriata and

> à RRR RRR à À d Da

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

A. mediovirgata. The gonopods of these species are very similar, except that in

A. bistriata and A. mediovirgata, the femorite is relatively shorter. A. mediovirgata

can be separated from A. bistriata by its much smaller size (width of male 2.0 mm

is against 3.4 mm), the development of the pleural keels (present up to the Sth

somite as against up to the 10th somite), and the development of the tarsal

scopulae (disappearing more rapidly in the anterior legs).

The typical colour pattern and general morphology of these three species is

shared by two other Burmese species, viz., A. bivittata and A. melanopleuris, and

although these species are known only in the female sex and their gonopod

characters are unknown, secondary evidence leaves no doubt as to their close

relationship to the type-siecies of the genus.

Less certain is the status of A. pardalis. In this species the colour pattern of the

metatergites is different, the yellowish paramedian stripes being reduced to a pair

of spots in front of the transverse furrow. Still more important is the fact that in

this species the transverse furrow of the metatergites is present from the 2nd

somite onwards instead of from the 5th onwards. The reference to Antheromorpha

of this species must be considered more or less arbitrary, and the discovery of the

male has to be awaited before a more definite conclusion can be reached.

The position of A. comotti, also based on a female, is also not yet settled.

Although it has the same colour pattern as A. miranda, it differs notably in the

structure of the paranota and in the leathery sculpture of collum and metatergites.

In this case, however, more can be said with regard to the relationship to

A. miranda, because A. comotti is evidently closely related tot A. orophila (Carl),

and in fact the latter name may prove to refer to the same species. The male of

A. orophila has gonopods basically similar to those of A. miranda, except that the

femorite apically is more strongly curved caudad, whereas the tibiotarsus is a little

more elongate and less curved than in A. miranda.

The only remaining Burmese species, A. minlana (Pocock), was not represented

in the collection of the Genoa museum, and has not yet been re-examined. It is a

relatively smaller species (width of male 2.6 mm, of female 3.0 mm), with the

paramedian light bands vestigial. According to the drawing given by Pocock the

gonopods are very slender, with the tibiotarsus more elongate than in A. miranda,

but otherwise they are essentially similar to those of the latter species.

Of the three non-Burmese species, A. festiva (Brölemann) from Siam is evidently

quite closely related to A. miranda. Apparently, Attems (1937: 69) knew only the

brief original description of this species and overlooked the lengthy and lavishly

illustrated account by Brölemann of 1904. The species has about the same size as

A. miranda (width of male 3.2 mm, of female 3.7 mm), the same colour pattern, and

the same characteristic habit with the diverging lateral borders of the paranota.

The gonopods are of the same general type and differ mainly in the somewhat

different course of the spermal channel in the basal part of the femorite, and the

relatively larger process of the lamina medialis of the tibiotarsus.

A. uncinata (Attems) from Siam also appears to be related to A. miranda, but

differs in the stronger curve of the femorite of the gonopods, the relatively larger

tibiotarsus, and in colour. Instead of two paramedian yellowish bands, this species

has a single median stripe. It is a large species (width of male 5.0 mm) which is also

JEEKEL: Burmese Paradoxosomatidae 87

characterized by the presence of two strongly developed cones at the apex of the

epiproct pointing ventrad.

A. harpaga (Attems) from Indochina, is a relatively small species (width of male

2.5 mm) which apparently has a colour pattern largely similar to that of

A. uncinata. In the gonopods it is characterized by the sigmoid curve of the spermal

channel in the basal part of the femorite, the presence of a rather distinct

denarcation between femur and postfemur, the relatively large tibiotarsus and the

large size of the process of the lamina medialis. Like A. uncinata it has the epiproct

provided with a pair of terminal cones pointing ventrad. Apparently this is the only

species in which the lateral sides of the collum are angular.

In conclusion we may distinguish the following groups of species in the genus

Antheromorpha:

|. A. bistriata, A. bivittata, A. festiva, A. mediovirgata, A. melanopleuris,

A. miranda;

2. A. comotti, A. orophila;

3. A. uncinata;

4. A. harpaga;

5. A. minlana;

6. A. pardalis.

Future studies may reveal the necessity of giving one or more of these groups

separate generic status.

In 1968 Antheromorpha was referred to the tribe Orthomorphini together with

Dajakina Jeekel, 1963, Eudasypeltis Pocock, 1895, Gigantomorpha Jeekel, 1963,

Orthomorpha Bollman, 1893, and Piccola Attems, 1953 (Jeekel, 1968: 128). Within

this tribe two groups of genera can be distinguished: one, including Orthomorpha,

Eudasypeltis and Piccola, in which the tibiotarsus of the gonopods is simple and

undivided, bearing only one or several tiny lobes at the extreme apex, and one,

including Antheromorpha, Dajakina and Gigantomorpha, in which the tibiotarsus

has a secondary branch arising from the lamina medialis. To the latter group will

be added two new genera to be proposed elsewhere, viz. Nesorthomorpha Jeekel (in

press) and Diglossosternum Jeekel (in press), based respectively on Orthomorpha

coriacea Carl, 1902, and Orthomorpha bipulvillata Carl, 1902, both from Java.

The interrelationship of these genera is evident on account of the gonopod

characters, but their position amongst each other is hard to define.

Antheromorpha may come closest to Dajakina from Borneo and Sumatra, but the

latter genus differs in the absence of a sternal process on the fifth sternite of the

male, the very weak development of the paranota, and the presence of pleural

keels in a few anterior somites only.

Gigantomorpha is particularly characterized by the presence on the medial side

of the gonopod femorite of two longitudinal chitinous crests. The paranota in this

genus, which occurs in Borneo as well as in Celebes, are also different in having

the latero-anterior border much more strongly rounded.

For the differential characters of the two new genera the reader is referred to

the forthcoming paper (Jeekel, in press).

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 3, 1980

ZOOGEOGRAPHY

The geographical distribution of the group of orthomorphine genera to which

Antheromorpha belongs, shows a remarkable aspect which deserves some

attention. The area in which these genera occur includes Burma, Siam, Indochina,

Sumatra, Java, Borneo and Celebes, but according to our present knowledge the

representation of the group in the various countries and islands is quite unequal.

The following survey may illustrate this.

Of a total of 27 species, Burma has 9, Siam 2, Indochina 1, the Malay Peninsula

0, Sumatra 1, Java 4, Borneo 9 and Celebes |. There is no doubt that these

numbers are biased by insufficient exploration of certain areas, but this affects the

absolute numbers and in a much less degree the relative part of the species as

against the total number of paradoxosomatid species in the countries. In this

respect it is evident that the species of the group are relatively numerous in the

faunas of Burma and Borneo, and that they form only a minor part of the faunas of

Indochina and Sumatra.

In particular the aspect of the fauna of Sumatra seems to indicate that in the

period of dispersal of the group faunal exchanges were possible only between Java,

Borneo and Farther India, whereas Sumatra and the Malay Peninsula were

apparently isolated and largely inaccessible.

A rather similar distributional pattern of the genus Orthomorpha, occurring in

Farther India, the Malay Peninsula and Java, was mentioned earlier (Jeekel, 1968:

133). Although this pattern differs from that of the group of genera to which

Antheromorpha belongs, it agrees with regard to the remarkable dissimilarity in the

faunas of Java and Sumatra.

REFERENCES

Attems, C., 1937. Polydesmoidea I. Fam. Strongylosomidae. Tierreich 68: I-xxii, I—-300.

Brölemann, H. W., 1904. Myriapodes recueillis par M. A. Pavie en Indo-Chine. Mission Pavie Indochi-

ne 1879-1895, Etudes diverses 3: 296— 309, pis. 13—15.

Jeekel, C. A. W., 1968. On the classification and geographical distribution of the family Paradoxosoma-

tidae (Diplopoda, Polydesmida): i-vii, I— 162. — Amsterdam, privately printed.

——, (in press). On the classification of some poorly known Paradoxosomatidae from South-East

Asia (Diplopoda, Polydesmida). — Rev. suisse zool.

Pocock, R. I., 1895. The Myriapoda of Burma, Pt. IV. Report upon the Polydesmoidea collected by

Sig. L. Fea, Mr. E. W. Oates and others. — Annali Mus. civ. Stor. nat. Genova 34: 787—834.

Silvestri, F., 1896. I Diplopodi. — Annali Mus. civ. Stor. nat. Genova 36: 121—254, figs. 1—26.

Weidner, H., 1960. Die entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologi-

schen Museums Hamburg. III. Teil. Chilopoda und Progoneata. — Mitt. Hamburg. Zool.

Mus. Inst. 58: 57— 104.

DI ‘gets 5 NM

ue, v |

= es Pr

I =» © 5

St it

I a

Br.

= es

DEL

sees

Hine peter n

sch ders

Hip Sia 1 i

CI Know na

quite

= td

_ DEEL 123 AFLEVERING 4 1980

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

B. L. BRADOO. — Feeding behaviour and recruitment display in the social spider

Stegodyphus sarasinorum Karsch (Araneae, Eresidae).

Tijdschrift voor Entomologie, deel 123, afl. 4 Gepubliceerd 9-V-1980

FEEDING BEHAVIOUR AND RECRUITMENT DISPLAY IN

THE SOCIAL SPIDER STEGODYPHUS SARASINORUM

KARSCH (ARANEAE, ERESIDAE)

B.L. BRADOO

Department of Zoology, D.A.V. College, Chandigarh, India

With one plate

ABSTRACT

The author describes the sequence of feeding behaviour and recruitment display in the social spider

Stegodyphus sarasinorum Karsch (Araneae, Eresidae) from India. The display informs the fellow

members of the colony about the presence of a prey in the web. The prey is subdued, transported and

consumed by these social spiders on a co-operative and social basis.

INTRODUCTION

Some general observations on the feeding habits and social behaviour of

Stegodyphus sarasinorum Karsch have been reported by Jambunathan (1905),

Gravely (1915), Kullmann (1968, 1969, 1972), Bradoo (1972) and Jacson & Joseph

(1973); on African species, particularly S. mimosarum Pavesi and S. simoni Giltay

has been reported by Cambridge (1889), Marshall (1898), Ghesquiere (1926) and

Giltay (1927a, b). But detailed studies on the feeding behaviour and recruitment

display of these spiders have not been made so far. Hence the present study was

undertaken to investigate the sequence of feeding behaviour in S. sarasinorum. For

the present study material has been used originating from Kerala State in India.

MATERIAL AND METHODS

Nests of S. sarasinorum, collected from Ferok and Kadalundi, District Calicut,

Kerala State, were fixed near the laboratory for making detailed observations on

these social spiders. They were fed on cockroaches, house crickets and other live

insects collected from the field. Some nests were collected at an interval of few

weeks and dissected in the laboratory for determining the types of prey captured

by these spiders. Such preys were identified from their chitinous remains and other

hard parts of their exoskeleton present in the nest materials. They were also

identified during the actual captures made by spiders of natural colonies in the

field.

S. sarasinorum is not strictly a nocturnal species. During the day, most spiders

Stay inside their silken nest but they at once emerge from the nest when diurnal

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

insects like bees, wasps, flies or odonates strike their webs and get entangled in

them. However, the maximum general activity begins only late in the evening after

sunset, when most spiders come out on the web sheets or on the nest surface, or

both.

The preys accepted by the laboratory colonies include different types of

mantids, grasshoppers, tettigonids, Gryllotalpa, house crickets, termite

reproductives, Forficula sp., dragon flies, house flies, robberflies, cockroaches,

adult antlions, Eumenes sp., paussid beetles and lady bird beetles. The preys

identified from the nest analysis include, membracids, honey bees, wasps,

dipterans, odonates, grasshoppers, mantids and a variety of Coleoptera; the

beetles include Carabidae, Curculionidae, Cicindelidae, Coccinellidae,

Elateridae, Buprestidae, Hydrophilidae, Staphylinidae, Scarabaeidae and

Lampyridae.

STRUCTURE OF THE NEST

S. sarasinorum is a social spider (Jambunathan, 1905). It makes large complex

silken nests on thorny bushes, shrubs and trees like Acacia, Ficus and Zizyphus sp.

as reported by Jambunathan (1905), Gravely (1915), Subrahmanyan (1953) and

Phanuel (1960). Bradoo (1967, 1972) found the nests of this species on bamboo

fencing around paddy fields in Kerala State and described its nest architecture and

Figs. 1, 2. Nests of the social spider Stegodyphus sarasinorum Karsch. 1, nest among the leaves of a

plant, showing small web-sheets and large nest exits; 2, a large nest with small exits on the surface

covered with adhesive cribellar silk.

BRADOO: Stegodyphus sarasinorum 9]

ecology. The nests are irregular silky constructions of variable shape and size

which incorporate leaves of plants (fig. 1), chitinous remains of preys eaten by the

spiders, their own exuviae and rolled bits of dry silk removed from their webs.

The nest is constructed by the joint and co-operative effort of the individuals of

the colony. Its size depends upon the number of spiders in the colony and the

number of generations that occupy the same nest, year after year. Normally each

generation lives in the nest for about a year. Some large nests (fig. 2) may be a few

years old and contain about 500 to 900 individuals. But this number decreases with

the increasing age and the longer life history of the spiders. At maturity, each nest

contains only few adult spiders. The number of males is less than the females and

polygamy is practised (Bradoo, 1975a, b).

The nests of S. sarasinorum are traversed by many branched, irregular and

interconnected tunnels of variable size which open on the nest surface or near its

margin by many exits. The size of the tunnels and the exits varies with the age and

the life history of the spiders, being small when the spiders are young, and larger

when they are full-grown. In Calicut, S. sarasinorum makes rain-proof nests during

the monsoons (Bradoo, 1972).

THE WEB

The nests of S. sarasinorum are surrounded by large, sticky sheets of cribellar

silk made by the joint and co-operative effort of the members of the colony. Some

adjacent nests may be interconnected by the common web sheets present between

them, as commonly observed among the younger colonies present on the same

bush, and among those nests that are formed by sociotomy (Jacson & Joseph,

1973). The structure and construction of the web has been described by Gravely

(1915) and Bradoo (1972), and the silk glands by Bradoo & Majupuria (1973).

Late in the evening, most spiders become active and after they come out of the

exits, begin to defecate and groom their bodies. Some of them also explore the

web and the surroundings, either by leaving fine silk threads floating in the air or

by suspending themselves from the nest surface or the web by fine drag lines.

Some individuals also begin to repair the damaged areas in the web by spinning

characteristic cribellar silk threads in a zigzag manner. The completion of the web

normally takes a long time, often a few days, and it attains its full size of 1—3

meters or more in a few days time. The size of the web depends on the life history

and the number of spiders present in the nest. Those colonies that contain several

hundred spiders usually have a much larger web than those that contain only few

individuals. The adult males take no part in web spinning or prey capture, but

perform these functions actively when still very young. Even the gravid females

and those that are looking after their cocoons take very little interest in spinning

and web repair, so that their web remains much smaller and considerably damaged

(fig. 3). Such gravid females use their silk for making special brood chambers in

the nest for storing their cocoons, as reported by the present author (1975b). They

obtain their prey from the surface of their nest, which is covered with sticky

cribellar silk (fig. 1).

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

Figs. 3, 4. Nests of the social spider Stegodyphus sarasinorum Karsch. 3, nest with damaged web during

the breeding season; 4, nest on Ficus plant covered with the drag-lines of spiderlings.

Some gravid females leave the home nest and construct new brood nests for

looking after their cocoons and the brood. Such new nests give rise to future

colonies of these social spiders (Jambunathan, 1905; Bradoo, 1972; Jacson &

Joseph, 1973).

The web sheets of these social spiders do not retain their sticky nature for a long

time. Various environmental factors, like excessive rain, heat, humidity and wind,

make the web dry and non-sticky. Moreover, the struggles of the ensnared preys

and the peculiar method of transportation also damage the web. Such dry and non-

sticky parts of the web are examined by the spiders each evening or even in the

early mornings at the time of sunrise. These dry webs are cut and rolled into small

balls, that are incorporated into the nest structure. But the complete web is never

removed or renewed. The damaged areas are jointly repaired by the spiders every

day. Krafft (1969) reported a similar co-operative web repair in the social spider

Agelena consociata Denis.

The structure, size and siting of S. sarasinorum webs is consistent with the view

that these spiders specialize on preys that are in flight above the ground surface.

That web plays a very prominent role and determines the manner of prey capture

among spiders has been discussed earlier by the present author (1973, 1977).

BRADOO: Stegodyphus sarasinorum 93

FEEDING BEHAVIOUR

The feeding behaviour of S. sarasinorum, as observed in the field and in the

experimental colonies, consists of the following sequence: (1) perception of

vibrations, (2) orientation and approach behaviour, (3) seizure and bite, involving

a recruitment display, (4) transportation of the prey, (5) feeding, and (6) treatment

of dry remains of the prey.

(1) Perception of vibrations

When some insect gets ensnared in the web sheets of S. sarasinorum, its struggles

set vibrations in the web, which are perceived by the spiders waiting on the web or

near the nest exits. The waiting spiders, hitherto called “pilot spiders”, are the first

individuals to reach and attack the prey in the web.

That the web spinning spiders respond to vibrations of the web has been

demonstrated by many workers (Boys, 1880; Peckham & Peckham, 1887;

Barrows, 1915; Rabaud, 1921; Grunbaum, 1927; Wells, 1936; Liesenfeld, 1956;

Walcott & Van der Kloot, 1959; Walcott, 1963). Tretzel (1961) found that the

female Coelotes terrestris (Wider) not only responds to the vibrations of the web

but can also distinguish her young from the prey by the differences in the web

vibrations. Walcott (1963) and Frings & Frings (1966) showed that the spiderweb

also transmits the sound vibrations. While the web vibrations are perceived by the

lyriform organs, the sound vibrations are perceived by the sensory hairs located on

the legs and the body. The detailed structure of spider lyriform organs has been

given by MacIndoo (1911) and Kaston (1935) and the electron microscopic study

by Salpeter & Walcott (1960).

(2) Orientation and approach behaviour

The sustained vibrations caused in the web by the ensnared preys elicit an

orientation and approach behaviour in the spiders. It involves a rapid locomotion

that brings the pilot spiders near the prey. The approach behaviour is released

even in the absence of the prey by touching a tunning-fork to the web. Such a

response is similar to that shown for a struggling prey. The intensity of web

vibrations is very important for the maintenance of orientation, otherwise the

spiders move in a random search on the web, keeping their first pair of legs wide

apart and finally they stop on the web.

The speed of approach behaviour is dependent on the intensity of the web

vibrations. The pilot spiders run straight and direct towards the source of

vibration. The “run” of an individual releases a similar behaviour in the other

neighbouring spiders present on the web. But certain preys that get ensnared far

away from the nest set only mild vibrations. In response, the pilot spiders start

rushing towards the prey. But when the prey stops its struggles (as observed in the

case of small acriids), the pilot spiders also stop and stay motionless on the web.

This is followed by the “plucking behaviour” which induces the prey to resume its

struggles and helps the spider to locate the prey. During plucking, the spiders

make characteristic web pulls with the help of their first pair of legs. This plucking

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

behaviour is similar to those of the orb-weavers as reported by Robinson (1969),

Robinson & Robinson (1971), Robinson & Mirick (1971) and Robinson & Graney

(1971).

The orientation behaviour helps the spiders to precisely locate the prey

ensnared in the web. As the pilot spiders are the first individuals to reach the prey,

they inform the other members of the colony by a special display through the web.

The locomotion of pilot spiders and few others that follow them to the prey, is

different from the normal locomotory behaviour in that the spiders are very much

excited and they run with a higher speed. The vibration receptors on the legs thus

play a significant role not only in perceiving the vibrations but also in the

orientation and approach behaviour towards the prey.

(3) Seizure and bite (prey-immobility)

The contact with the prey initiates seizure and biting of its appendages. It

involves a joint activity of the pilot spiders and many others that receive the signals

for subduing the prey. The pilot spiders recognize the prey by leg and palpal

contacts. They seize the movable appendages like mouth-parts, legs, wings and

antennae with the help of their chelicerae. The prey held taught in this manner

cannot escape from the firm hold of the spiders. To make its escape impossible,

the spiders exert a maximum pull on the appendages and begin to inject poison to

kill the prey. During these activities, the pilot spiders make repeated drumming

movements and special webpulls with the help of their hind legs on the web. This

display is very prominent when the prey continues to struggle in the web and it

induces many spiders (recruits or helpers) to rush out from the nest to assist the

pilot spiders in subduing the prey. Hence various preys that cannot be easily

captured are successfully subdued by the joint effort of the pilot spiders and the

recruits. Unlike in most other spiders, all types of preys receive the same

treatment.

The display that induces recruitment and communicates information to the

fellow members of the colony is called the recruitment display. It is connected

with the vibrations caused over the web. The pattern of recruitment display is

almost similar under different conditions. Under optimal conditions (attractive

ensnared prey, active and hungry colony) the pilot spiders, while holding and

attempting to drag the prey, perform a repeated display by drumming their hind

legs on the web and by making special web pulls accompanied by high excitation

of their pedipalps. During the display, the pilot spiders do not leave the prey unless

they themselves are attacked by the prey. Such a display was never seen “in

vacue”, meaning without the stimulation caused by a prey. This display is shown

only for the live prey and results in a mass-recruitment of the spiders from one or

many nests that may be interconnected by the common web sheets. The onset of

departure from the nest is induced by the display and the excited behaviour of the

pilot spiders, holding the prey. So new individuals (‘‘recruits’” or ‘thelpers’’)

emerge out from the nest and run towards the prey. They leave fine drag lines as

they run over the web.

BRADOO: Stegodyphus sarasinorum 95

The strength of the recruitment is, as a rule, determined by the strength of the

display but it also depends upon the struggles of the prey which induces web

vibrations and an active display in the pilot spiders. However, once the prey has

been subdued, the display alone is responsible for the recruitment. The

recruitment display not only stimulates the fellow members of the colony but also

directs them to the food source.

(4) Intensity of the recruitment display

The recruitment display is carried out only by the pilot spiders. High intensity

display is performed when the pilot spiders are few and its intensity goes down,

when many more spiders reach the prey. Finally it disappears completely.

Sometimes the display is interrupted when an appendage held by any pilot spider

gets detached from the rest of the prey. The spider carries it for a short distance on

the web or near the nest to feed on it all alone, an observation that Jambunathan

(1905) termed as a “selfish act”. The display of the pilot spiders together with the

excited running of the recruited spiders results in a greater activity in the nest, and

more spiders stream out towards the prey. It may cause a similar behaviour in the

other neighbouring nests, having a common snare between them. The display is

sometimes superimposed and very brief, when the prey gets ensnared close to the

nest where large number of spiders come out to cover the prey, as usually

observed late in the evening. The distance of the prey from the nest is very

important for the period of excitation.

The intensity of recruitment display gradually diminishes. The display pattern

becomes weaker, the drumming movements and the web pulls slower and shorter

lasting and finally disappear entirely. Meanwhile the prey is surrounded by large

number of spiders that begin to drag it towards the nest. The leg and palpal

excitation also slows down, and from the entire display, only hold of the prey

remains. The subsequent recruitment is very slow and finally nothing remains of it

and it disappears. The mass recruitment finally breaks off as everything grows

quiet around the prey.

The intensity of recruitment display, its dropping down and breaking off and the

length of the recruitment period, depends on the size and the attractive power of

the prey in the web and also on the time of the day. The recruitment display is best

observed when preys like flies (Diptera), bees (Hymenoptera), termite

reproductives (Isoptera) and other similar and active insects get ensnared in the

web, particularly during the day time, when only few pilot spiders are attracted to

the prey in the web. During the night, this display is observed when there are only

few spiders on the web and larger preys like grasshoppers (Orthoptera), beetles

(Coleoptera) or termites (Isoptera) get ensnared in the web sheets of these social

spiders. The strength of the recruitment is dependent on the conditions of the

colony, prey distance, the type of the prey and also the condition of the web.

Soon after the recruitment ends, all the individuals (pilot spiders, recruits and

others) get engaged in a joint and communal feeding on the prey. The dead preys

and the members of their own species did not evoke any display, as the pilot

spiders do not run and pull the prey. Individuals of S. sarasinorum, when thrown on

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

the webs of other colonies of this species, were not treated as prey by the resident

spiders. They would simply exchange mutual tactile contacts and would either run

to the nest or move in a normal fashion on the web. This indicates that these social

spiders do recognize the members of their own species. When they run towards the

nest, it induces the other spiders also to rush back to the nest and it indicates that

they understand the nature of the web vibrations caused by the fleeing spiders on

the web. The mutual recognition, though based on tactile sense, is possibly aided

by some chemicals and pheromones as suggested by Kullmann & Zimmermann

(1971). Krafft (1975a, b) reported a similar mutual tolerance behaviour among the

members of Agelena consociata colonies.

(5) Intensity of recruitment

The intensity of recruitment may be determined by the number of spiders

leaving the nest after a display of the pilot spiders and also by the number of

spiders arriving at the prey. This results in the formation of an aggregation and

coherence of many individuals around the prey. The number of recruited

individuals varies considerably. Many recruited spiders follow the drag lines left

by the pilot spiders and by those that leave the nest first and thus reach the prey.

Allthe recruited spiders do not emerge from the same exit but they come out from

several adjacent nest exits, but not from those exits that are far away or those that

lie on the different side of the nest.

Sometimes, all the recruited spiders may not arrive at the prey. They may

retreat back into the nest, particularly if it is day-time. This retreat behaviour may

be due to some mechanical disturbance in the web or may be a result of loss of

necessary signals through the web, that may be given by the pilot spiders from a

longer distance on the web. The retreat behaviour can also be induced by simply

touching the spider and preventing its course towards the prey. It can also be

induced by a strong and sudden flash light shown by the observer.

(6) The build up of recruitment

When some suitable prey gets ensnared in the web, the pilot spiders rush

towards the source of vibrations and begin to display the presence ofthe prey. This

display induces the recruits to rush out from the nest to help the pilot spiders. The

number of recruits depends on the prey size, the intensity of the display and the

stimulation received by them according to the distance of the prey from the nest.

The strength of the recruitment also depends upon the number of pilot spiders, the

struggles of the ensnared prey, colony needs and other possible factors. Thus in a

starved colony (which is given no food for two weeks or more) more pilot spiders

begin to move about and explore the web for any possible prey that may strike the

web. In such colonies, many spiders may even leave the. nest and try to start new

nests, as observed in the experimental colonies. In a starved colony the period of

recruitment is much shorter than usual. The strength of the recruitment markedly

diminishes after several preys are available in the snare. In such cases several

separate groups of spiders collect around the different preys on the web.

BRADOO: Stegodyphus sarasinorum 97

It is not surprising that these social spiders on account of their well developed

sense of touch have developed tactile releasers. Tretzel (1961) pointed out that the

Coelotes mother after prey capture shows remarkable movements and vibrations.

The manner of these movements and their repetition has a role as a releaser. Thus

the ‘‘web-shake”’ used for longer prey attracts the young from a closer range. The

““body-quivering” is given when the mother struggles hard with a beetle.

“cheliceral-shaking” is given when a small prey is held in the chelicerae. It may

also give a special palpal tap on the web. These movements associated with the

entanglement of the prey, act not only as a luring signal for the spiderlings to come

to the meal but also helps them to understand the details regarding the magnitude

and the situation of the prey. The Coelotes mother treats different preys in a

different manner.

In the same manner, the recruitment display in Stegodyphus spiders acquires a

signal value which not only stimulates the fellow members of the colony but also

directs them to the food source.

(7) Transportation of the prey

The method of transportion of the prey is characteristic, as it involves a joint

and co-operative effort of a large number of pilot spiders and numerous recruits.

Unlike in orb-weavers, the prey is never wrapped in silk, before or after the

transportation. During transportation, the prey irrespective of its size and kind is

practically dragged on the web in the direction of the nest. This damages the web

considerably but this joint action of many individuals is necessary for subduing and

transporting the prey which normally is much larger than the spiders.

Before the actual transportation begins, the spiders surround the prey from all

sides, hold its appendages and some of them even change their positions to get a

better hold of the prey. The prey is never separated from the web unless it is to be

refused and thrown down. While jointly dragging the prey on the web, the spiders

move backwards in the direction of the nest, guided by numerous drag-lines that

are laid down on the web by the recruits when they rush out to reach the prey.

Thus the recruits not only help in subduing the prey but also in transporting the

prey in the proper direction of the nest. The direction of orientation is therefore

predictable and determined by the number of recruits coming from a particular

nest.

The transportation of the prey finally stops when it reaches the nest. A few more

spiders emerge from the nest to assist them and the number of spiders soon

increases around the prey. The prey is never carried inside the nest unless it is too

small and the spiders are forced to do so. Normally the prey is so large that it

cannot be carried into the nest and if the spiders are disturbed, they rather leave

the prey and rush back to the nest.

The transportation of the prey is considerably affected by the presence or

absence of light. During the day, all types of prey are transported up to the nest,

but late in the evening and during the night the prey is consumed on the web itself

without transportation. The prey is covered by a large number of spiders. If a torch

light is shown on such feeding aggregations, many spiders leave the prey and

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

retreat into the nest. Though the dim light induces the spiders to pull the prey,

feeding is resumed soon.

A Mantis thrown on the web of an experimental colony was located by the pilot

spiders. They held it by the wings and began to perform the display. A few recruits

arrived to hold the legs of the mantis but were furiously bitten by the Mantis and

they ran back to the nest with injured legs. Meanwhile, some more recruits rushed

out to assist the pilot spiders and within half an hour, the mantis was surrounded

by 31 spiders and was completely subdued. When a dim torch light was shown, the

spiders began to move their hind legs and many of them moved away on the web,

but returned again to drag it towards the nest.

That the prey is invariably carried to the nest during the day has a survival value.

The large aggregations of spiders around the prey in the web would attract many

insectivorous birds. Such predators and other kinds of natural enemies of

S. sarasinorum have been reported by the author earlier (1972). Once the author

found a large number of nests of these spiders destroyed by house crows, at Ferok,

Kerala. As these avian predators are not active at night, the social spiders need not

transport the prey to the nest. But those spiders that continue to feed during the

night on the web itself, do carry the prey to the nest as the morning approaches, to

minimize the danger of predation and possible damage to the nest.

(8) Feeding

Large feeding aggregations of S. sarasinorum have been nicely photographed by

Jacson & Joseph (1973). Such feeding groups may have few to hundred or more

individuals, depending on the size of the prey, the size of the colony and the

physiological state of the spiders.

It should be recognized that in these feeding groups, some individuals may not

have actually participated in any way in the prey capture or transportation. But

they are permitted by the other active members to partake the food captured by

them. This is one of the interesting advantages of their social life, and almost every

member receives the required nourishment depending upon the availability of the

prey. In such aggregations, and in those colonies that have many individuals, not

all the members may receive an equal share of the prey due to insufficient number

of preys, in a particular period. Thus the various members of a colony mature

slowly in course of time due to differences in the growth rate and prey availability.

The prey sharing behaviour indicates a high degree of mutual tolerance and co-

operation in this species. A similar prey sharing behaviour has also been reported

in the social spider A. consociata by Pain (1964) and Darchen (1965), and in

A. republicana by Darchen (1967, 1973). Even in the solitary species, S. lineatus

(Latreille), the prey is jointly shared when many individuals were put together

under laboratory conditions (Millot & Bourgin, 1942).

In each feeding group, the spiders pour digestive secretions that dissolve the soft

interior of the prey. The semidigested liquid is then sucked jointly by the spiders.

The maintenance of feeding was found primarily to be dependent on the sensory

inputs from the chemoreceptors present around the mouth. The duration of

feeding depends upon the size of the prey and the number of feeding spiders. The

BRADOO: Stegodyphus sarasinorum 99

small acridiids were completely sucked within two hours, a dragonfly in one hour,

a large cockroach (Periplaneta americana) in four hours. Grasshoppers and larger

beetles took longer. These observations clearly show that social spiders feed and

consume larger preys in a shortest possible time and the spiders are soon ready for

new preys that may strike the web.

(9) Treatment of dry remains of the prey

After the cessation of feeding, the dry remains of the prey are separated and

thrown down from the web by the spiders that are the last individuals on the prey.

This behaviour is also shown by any individual that may come in contact with the

prey-remains (refuse) while exploring the web. Even the dry bits of grass, paper

bits or dry remains of the preys if placed on the web are removed and thrown

down, a behaviour also known among most other spiders, particularly the orb-

weavers. This behaviour of throwing the refuse away keeps the web clean and

possible predators away. But the remains of the prey that are transported to the

nest are not thrown out but are covered in silk and incorporated into the nest

structure. This evidently increases the size of the nest. Such dry remains of preys

are eaten by a number of insect commensals found in the nests of social spiders, as

reported by Bradoo (1967, 1971), Bradoo & Joseph (1970) and, among the African

species, by Marshall (1898).

(10) Drinking

The social spiders S. sarasinorum also accept drops of water during the summer

months when the colonies contain mature spiders. As soon as some water drops

are sprinkled on the nest surface, the spiders after initial contact with the drop of

water begin to drink it. This behaviour is particularly observed for the mature

males in the nest.

(11) Behaviour with unusual preys

The author studied the behaviour of these social spiders towards some unusual

preys that normally do not fall in their webs. The following observations are briefly

recorded:

(a) When a medium-sized female Gasteracanth spider was dropped on the web

of S. sarasinorum, a few pilot spiders rushed forward to hold its abdominal

appendages present in the form of small projections. But the pilot spiders ran back

to the nest as if in “distress”. The Gasteracanth was not accepted as a prey.

(b) A Scutellerid bug when placed on the web or the nest surface was

repeatedly thrown down and rejected by the spiders after an initial contact. When

placed inside a nest exit, it was again pushed out and rejected. In the same manner,

a blister beetle and a Phalangid were also rejected and thrown down from the web.

later, when placed again, was accepted as prey.

(d) When a large soldier ant was placed on the web, many pilot spiders

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

approached to hold its appendages but the ant struggled hard and injured some of :

them by biting their legs. Such injured individuals left the ant, which struggled

hard, and more spiders rushed out and successfully subdued it without

approaching its mouthparts.

(e) Large Rhinoceros beetles and Sphingid moths, when placed on the web,

induce a fleeing response in these spiders; the powerful wings of such a moth beat

so vigorously that the web gets damaged quickly and the moth finally escapes. The

social spiders were unable to hold the strong chitinous and powerful appendages

of the large rhinoceros beetle (5 cm long). The beetle damaged the web and fell

free to the ground. Such large insects are certainly difficult to subdue and the

spiders would face the danger of falling to the ground, along with the prey. The

size and suitability of the prey is therefore an essential factor for determining the

response of the spiders.

(f) Very minute insects like winged aphids, fig-insects and other chalcids,

minute diptera and micro-coleoptera also get ensnared in the sticky web sheets of

these spiders. But because of their small size and extremely weak web vibrations,

they are not perceived by the social spiders or they are at least not reacted upon.

Such minute insects are insignificant for the nutritional requirements of a large

colony of spiders. On several occasions, the author observed that the winged

aphids were actually separated and removed from the web and dropped down to

the ground by the spiders. However, these minute insects are actively captured by

a small cribellate spider Uloborus ferokus Bradoo (1979) that lives as a commensal

on the web sheets of this social spider.

(g) The young of S. sarasinorum do not spin the typical cribellar silk threads

until they complete the third or fourth moult. They cover the outer surface of the

nest with fine drag lines (fig. 4) and leave only minute nest exits for their own use.

Due to this close covering on all sides of the nest, the mother spiders are unable to

come out and hence they die inside the nest or beneath the covering of drag lines.

Their dead bodies are sucked by the spiderlings. They also feed jointly on those

small preys that fall on the nest surface.

DISCUSSION

From the results of this study, a complete description may be made of the

feeding behaviour of S. sarasinorum. The spiders that have not fed recently,

remain waiting on the web or near the nest exits, for any possible prey that may get

ensnared in the web. These waiting spiders, called “pilot spiders’, perceive the

vibrations of the web caused by the struggles of the ensnared prey. The intensity

and the continuity of the web vibrations releases an approach behaviour in the

pilot spiders. They perceive the vibrations with the help of the lyriform organs

located on their legs and these organs also help them to reach the prey. Contact

with the prey initiates its capture and seizure of its appendages and wings.

The sustained struggles of the prey make the pilot spiders to make repeated leg

taps and special web-pulls with their hind legs. This is particularly shown when

their number is low and they are far away from the nest. These movements release

an approach behaviour in other spiders, called ‘‘recruits’’ or “helpers”, to reach

BRADOO: Stegodyphus sarasinorum 101

the prey. They assist the pilot spiders in subduing the live prey which is normally

much bigger than the spiders. While holding the prey, the spiders inject poison and

begin to transport it towards the nest, if it is day-time. But late in the evening, the

prey is not transported and instead feeding commences on the web itself. The

digestive secretions of large number of spiders dissolves the soft parts of the prey

in a short time and this semidigested material is sucked jointly by the spiders.

The duration of feeding depends upon the size of the prey and the number of

spiders and their physiological state at the time of feeding. On larger preys like

grasshoppers and cockroaches, the spiders may continue to feed for several hours,

but very large beetles require a much longer time so that the spiders may continue

to feed even the second day. Unlike in most other spiders, particularly the orb-

weavers, the prey is never wrapped in silk, either before or after the

transportation.

The cessation of feeding is based mainly on decrease in the supply of

nourishment. The remains of preys consumed on the web are normally rejected

and thrown down, but those that are consumed near or on the nest surface are

usually incorporated into the nest structure.

The unsuitable preys and those with irritating chemical secretions are rejected

after the initial bite or after the first contact with the prey. This indicates the

importance of gustatory stimuli in food discrimination apart from those repellant

odours and chemicals that are encountered during or after the initial contact and

before the actual bite.

According to Buskirk (1975), those spiders that live in groups or colonies are

benefitted in several ways. The social life in S. sarasinorum also serves many

advantages, particularly in feeding, due to following reasons:

(1) the joint and co-operative activity helps these spiders to subdue and kill

preys that are normally much larger than the spiders;

(2) several preys that may fall in the web at the same time or at different times

one after the other, are successfully captured by the different batches of spiders;

(3) injuries caused by the stronger prey while subduing it are minimized by the

joint and co-operative action of many spiders;

(4) the group activity prevents the escape of active preys like flies, bees and

wasps from the web;

(5) unlike in most other spiders, all kinds of preys receive the same general

treatment by these social spiders;

(6) the joint action of many spiders helps in easy transportation of the prey on

the web and makes wrapping to prevent escape superfluous;

(7) the social life and mutual tolerance helps every individual of the colony to

get the necessary nourishment;

(8) the mutual prey-sharing behaviour helps even those individuals of the

colony that may not have actually participated in the prey capture and its

transportation;

(9) the group feeding on a single prey also brings about the quick digestion and

disposal of the prey, irrespective of its size; thus, unlike in solitary spiders, the

larger prey is consumed in a short time and the spiders are ready and free for the

next meal;

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

(10) the joint effort of these social spiders is very useful in the early part of their

life history, because the small spiderlings cannot yet make a typical sticky cribellar

silk sheet around their nest; they lack cribellum and calamistrum which develop

only after the third or fourth moult (Bradoo, 1972). Hence the juvenile web of

these spiderlings is mainly made of drag lines laid out on the outer surface of the

nest; this type of web is capable of trapping only smaller preys that are jointly

consumed by the spiderlings; they also eat the dead bodies of their mothers.

(11) the mother spiders are also observed catching smaller insects like bees,

flies, forficulids, etc., that are carried by them close to the nest and are shared

jointly with the numerous spiderlings; on such occasions, it is not unusual for the

spiderlings to move about over the body of the mother to reach the prey.

SUMMARY

The social spider S. sarasinorum mainly captures large flying insects that are

caught in the sticky web sheets around the nest. The ensnared prey sets the web

vibrations that are initially perceived by few “pilot spiders” waiting on the web or

near the nest exits. While holding the prey, the pilot spiders inform the fellow

members of the spider colony by repeated drumming movements of their hind legs

on the web, accompanied by the special web pulls. This type of display through the

web signals and induces the “recruits” or “helpers” to rush out of the nest and

reach the prey. The prey is then jointly subdued, transported and consumed by the

spiders on a co-operative and social basis. The prey is neither covered in silken-

cover nor is it separated from the web.

All kinds of preys, except those that are “‘distasteful” are treated in the same

manner, as described above. The remains of the preys that are consumed on the

web itself are separated and dropped down, but those preys that are transported

up to the nest are mostly incorporated into the nest structure.

The transportation of the prey is mainly dependent on the presence of light.

During the night, the prey is consumed on the web itself but during the day it is

carried up to the nest prior to feeding. The “distasteful” prey is either rejected by

a retreat behaviour or by separating and dropping it down from the web. Very

minute preys are not accepted because of their small size and extremely weak

vibrations caused in the web. Such minute preys are insignificant for the

nutritional requirements of a large spider colony.

ACKNOWLEDGEMENTS

I wish to express my deep sense of gratitude to Prof. Dr. K. J. Joseph, D. Sc.,

Head of the Department of Zoology, University of Calicut, presently at the

University of Basrah, Iraq, for his help, encouragement and facilities during the

course of this investigation at Calicut.

I also record my thanks to Dr. R. E. Buskirk, Dr. S. E. Reichert, Dr. J. Reiskind,

Dr. R. Darchen, Dr. B. Krafft and Dr. M. H. Robinson, for numerous reprints of

their papers on spider behaviour.

BRADOO: Stegodyphus sarasinorum 103

REFERENCES

Barrows, W. M., 1915. The reaction of an orb-weaving spider, Epeira sclopetaria Clerck, to rhythmic vi-

brations of its web. — Biol. Bull. 29: 316—332.

Boys, C. V., 1880. The influence of a tuning-fork on the garden spider. — Nature 23: 149—150.

Bradoo, B. L., 1967. Observations on the life history and bionomics of Oligotoma ceylonica ceylonica

(Embioptera), commensal in the nest of Stegodyphus sarasinorum Karsch. — J. Bombay nat.

Hist. Soc. 64 (3): 447—454.

—., 1971. Further records of Embiopteran and Microlepidopteran nest mates of S. sarasinorum

Karsch. — Indian J. Ent. 33 (1): 90.

—., 1972. Some observations on the ecology of social spider S. sarasinorum Karsch (Araneae: Ere-

sidae) from India. — Oriental. Ins. 6 (2): 193—204.

—., 1973. Spiders and webs. — Science Rep. C.S.I.R. Delhi 10 (8): 374—376.

———, 1975a. Notes on the sexual biology and morphology of the reproductive organs of S. sarasino-

rum Karsch. — Ent. month. Mag. 11: 239—247.

—., 1975b. The cocoon spinning behaviour and fecundity of S. sarasinorum Karsch from India. —

J. Bombay nat. Hist. Soc. 72 (2): 392—400.

—., 1977. How spiders catch their prey. — Science Rep. C.S.I.R. Delhi 14 (2): 89—94.

—.,, 1979. Uloborus ferokus sp. nov. (Araneae: Uloboridae) a commensal of S. sarasinorum Karsch.

— Bull. Brit. Arachnol. Soc. 4 (8): 353——355.

Bradoo, B. L., & K. J. Joseph, 1970. Life history and habits of Oligotoma greeniana End. (Embioptera)

commensal in the nest of S. sarasinorum Karsch. — Indian J. Ent. 32 (1): 16—21.

Bradoo, B. L. & T. C. Majupuria, 1973. Studies on the anatomy of social spider S. sarasinorum Karsch.

PartI: Spinnerets and silk glands. — Zool. Anz. 190 (1/2): 67— 73.

Buskirk, R. E., 1975. Coloniality, activity patterns and feeding in a tropical orb-weaving spider. — Eco-

logy 56 (6): 1314—1328.

Cambridge, O. P., 1889. On some new species and new genus of Araneida. — Proc. Zool. Soc. Lond.

4244,

Darchen, R., 1965. Ethologie d'une Araignée sociale, Agelena consociata Denis. — Biol. Gabon. | (2):

117— 146.

—, 1967. Une nouvelle araignée sociale du Gabon, A. republicana Darchen (Aranéide labidogna-

the). — Biol. Gabon. 3: 31—42.

—, 1973. L’Ecologie d'une araignée sociale A. consociata D. à la lumière de quelques expériences

laboratoire. — Insec. Soc. 20 (4): 379—384.

Frings, H. & M. Frings, 1966. Reactions of orb-weaving spiders (Argiopidae) to air borne sounds. —

Ecology 47 (4): 578—588.

Ghesquiere, M., 1926. Une nid d’Araignees sociales du Congo. — Rev. Zool. Afric. 14: 58—59.

Giltay, L., 1927a. Les Araignées sociales Congolaises. — Rev. Zool. bot. Afric. 15 suppl., Cercle Zool.

Cong. Bull. 4: 35.

——, 1927b. Une Araignée sociale du Kasai (Stegodyphus simoni n. sp.) — Rev. Zool. bot. Afric. 15

(1): 105—117.

Gravely, F. H., 1915. Notes on the habits of Indian Insects, Myriopods and Arachnids. — Rec. Ind.

Mus. 11: 483—539.

Grunbaum, A. A., 1927. Ueber das Verhalten der Spinne Epeira diademata besonders gegenüber vibra-

torischen Reizen. — Psychol. Forschung. 9: 275-299,

Jacson, C. C. & K. J. Joseph, 1973. Life, history, bionomics and behaviour of S. sarasinorum Karsch. —

Insec. Soc. 20 (2): 189—204.

Jambunathan, N.S., 1905. The habits and the life history of a social spider S. sarasinorum Karsch (Ere-

sidae). — Smiths. Misc. Coll. 47: 365—372.

Kaston, B. J., 1935. The slit sense organs of spiders. — J. Morphol. 58: 189—208.

Krafft, B., 1969. Various aspects of the biology of Agelena consociata Denis, when raised in the laborato-

ry. — Amer. Zool. 9: 201—210.

———, 1975a. Les interactions limitant le cannibalisme chez les Araignees solitaires et sociales. —

Bull. Soc. Zool. Fr. 100 (2): 203—221.

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 4, 1980

—, 1975b. La tolérance réciproque chez l’araignée sociale Agelena consociata Denis. — Proc. 6th

Int. Arachn. Congr. Amsterdam: 107—112, pl. 1.

Kullmann, E., 1968. Soziale Phenomene bei Spinnen. — Insec. Soc. 14 (3): 289—297.

, 1969. Beobachtungen zum Sozialverhalten von Stegodyphus sarasinorum Karsch. — Bull. Mus.

Nat. Hist. Nat. 41: 76—81.

, 1972. Evolution of social behaviour in spiders. — Amer. Zool. 12: 419—426.

Kullmann, E. & W. Zimmermann, 1971. Versuche zur Toleranz bei der permanent-sozialen Spinnenart

Stegodyphus sarasinorum Karsch. (Fam.: Eresidae). — Proc. Sth Int. Congress Arachn. Brno:

175— 182.

Liesenfeld, F. J., 1956. Untersuchungen am Netz und uber den Erschütterungssinn von Zygiella x-nota-

ta. — Zeit. vergl. Physiol. 38: 563—592.

MaclIndoo, N. E., 1911. The lyriform organs and tactile hairs of Araneads. — Proc. Acad. Nat. Sci. Phil.

63: 375—418.

Marshall, G. A. K., 1898. Notes on South African social spiders. — Zoologist 688 (2): 417—422.

Millot, J. & P. Bourgin, 1942. Sur la biologie des Stegodyphus solitaires. — Bull. Biol. Fr. et Belg. 76:

299—314.

Pain, G., 1964. Premières observations sur une espèce nouvelle d’Araignées sociales, Agelena consociata

Denis. — Biol. Gabon. 1: 47—58.

Peckham, G. W. & E. G. Peckham, 1887. Some observations on the mental powers of spiders. — J.

Morphol. 1: 383—419.

Phanuel, G. J., 1960. Unusual nest site of the social spider S. sarasinorum Karsch. — J. Bombay nat.

Hist. Soc. 57 (3): 686—688.

Rabaud, E., 1921. Influence des vibrations mécaniques d'une araignée Tetragnatha extensa Lin. — C.

R. Soc. Biol. 84: 763— 765.

Robinson, M. H., 1969. Predatory behaviour of Argiope argentata (Fabricius). — Am. Zool. 9:

161—173.

Robinson, M. H. & B. Robinson, 1971. The predatory behaviour of the ogre-faced spider Dinopis longi-

pes. — Amer. Midl. Nat. 85 (1): 85—96.

Robinson, M. H. & H. Mirick, 1971. The predatory behaviour of the golden-web spider Nephila clavi-

pes. — Psyche 78 (3): 123—139.

Robinson, M. H., B. Robinson & W. Graney, 1971. The predatory behaviour of the nocturnal orb web-

spider Eriophora fuliginea. — Rev. Per. Entom. 14 (2): 304—315.

Salpeter, M. M. & C. Walcott, 1960. An electron microscopical study of a vibration receptor in the spi-

der. — Exp. Neurol. 2: 232—250.

Subrahmanyan, T. V., 1953. On the habits of Indian Eresid spider S. sarasinorum Karsch (Eresidae). —

J. Bombay nat. Hist. Soc. 51: 521—522.

Tretzel, E., 1961. Biologie, Oekologie und Brutpflege von Coelotes terrestris (Wider). Teil Il. Brutpfle-

ge. — Z. Morph. Okol. Tiere 50: 375—542.

Walcott, C., 1963. The effect of the web on vibration sensitivity in the spider Achaeranea tepidariorum

(Koch). — J. Exp. Biol. 40: 595—611.

Walcott, C. & W. G. van der Kloot, 1959. The physiology of the spider vibration receptor. — J. Exp.

Zool. 141: 191—244.

Wells, F. L., 1936. Orb weavers differential responses to a tuning-fork. — Psyche 43: 10—13.

DEEL 123 AFLEVERING 5 1980

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

VAN DER VELDE & TH. C. M. Brock. — The life history and habits of

Notiphila brunnipes Robineau-Desvoidy (Diptera, Ephydridae), an aut-

ecological study on a fly associated with nymphaeid vegetations, p. 105—127,

text-figs. 1—7.

THE LIFE HISTORY AND HABITS OF NOTIPHILA

BRUNNIPES ROBINEAU-DESVOIDY (DIPTERA,

EPHYDRIDAE), AN AUTECOLOGICAL STUDY ON A FLY

ASSOCIATED WITH NYMPHAEID VEGETATIONS

G. VAN DER VELDE and Th. C. M. BROCK

Laboratory of Aquatic Ecology, Catholic University of Nijmegen, The Netherlands

With 7 figures

ABSTRACT

The life history and some ecological aspects of Notiphila brunnipes R.-D. are described. Special

attention is paid to the interrelations with the nymphaeid plant species Nuphar lutea (L.) Sm., Nymphaea

alba L., Nymphaea candida Presl and Nymphoides peltata (Gmel.) O. Kuntze. Observations have been

made on phenology, occurrence in space and time, habitat choice, food, general behaviour, sexual

behaviour, sex ratio, the deposition of eggs and mortality of the adults. Possible predators are

discussed. The eggs are described and the function of the chorion is discussed. Some experiments have

been carried out to study the development of the eggs in relation with humidity, oxygen and

temperature. The habitat choice, food, mortality of and predation on the larvae are described and

discussed. Some data on the puparia are given. Effects of the dry summer of 1976 are described. The

geographical distribution is discussed; the absence of N. brunnipes north of the limes norrlandicus can

be explained from the autecological data gathered during this study.

INTRODUCTION

Nymphaeid-dominated ecosystems, which are very common and show an

optimal development in the Netherlands, have been studied by us during several

years. Nymphaeids are aquatic plants with floating leaves, which root in the

bottom and have flowers above or floating on the water surface. By their structure

they characterize the ecosystem. They have an important function as habitat for

many organisms, among them semi-aquatic insects such as the Ephydridae.

The species Notiphila brunnipes Robineau-Desvoidy, 1830 (syn. N. stagnicola

Stenhammar, 1844, nec Robineau-Desvoidy, 1830, and N. chamaeleon Becker,

1896) occurred abundantly on the nymphaeids we studied and appeared to have

many interrelations with Nymphoides peltata (Gmel.) O. Kuntze, Nymphaea alba L.,

Nymphaea candida Presl and Nuphar lutea (L.) Sm. Because of our interest in these

interrelations, the life history and habits of Notiphila brunnipes were studied in

detail and a review was made of data from the literature.

105

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

DESCRIPTION OF THE STUDY AREAS

Most observations have been made in the Oude Waal near Nijmegen

(municipality of Ubbergen, Province of Gelderland) (fig. 1). The Oude Waal is an

old river branch cut off from the river Waal. Here extensive vegetations

dominated by Nuphar lutea, Nymphaea alba and Nymphoides peltata occur. The

100 150 200 250m

Fig. |. Map of the Oude Waal.

Oude Waal consists of a large shallow water, | km in length and about 250 m wide,

and three interconnected ponds (D, E and F). The depth of the large shallow water

varies from 0.5 to 1.5 m; in the dry summer of 1976 it became almost completely

dry. About once every two years this area is flooded in winter or spring by water

from the river Waal; summer high waters occur only occasionally. The depths of

the three interconnected ponds increase towards the centre to 2.5, 5.5 and 5.5 m,

respectively (D, E and F). The bottom of the Oude Waal consists of a pattern of

clay and sand, sometimes mixed, covered by a sapropelium layer of varying

thickness.

Some additional observations have been made in the Haarsteegse Wiel

(municipality of Vlijmen, Province of Noord-Brabant), which originated from two,

but not simultaneous, bursts in the dike of the river Meuse. This water has an area

of 17.9 ha and a maximum depth of 17 m. The lake has a vegetation dominated by

Nuphar lutea and Nymphaea candida, which are almost completely restricted to the

southern and western parts of the lake and which are sheltered against wave and

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 107

wind action. Nymphoides peltata is very scarce here. The bottom of the Haarsteegse

Wiel consists mainly of sand (‘“‘wash-over deposits”), but where nymphaeids grow

there is a layer of sapropelium, which becomes thicker towards the littoral border.

The hydrology of the lake is dependent on rain and ground water only and is more

stable than that of the Oude Waal; the water-level shows only small fluctuations.

OCCURRENCE OF ADULTS ON NYMPHAEID LEAVES AND FLOWERS IN SPACE AND TIME

Methods

N. brunnipes (fig. 7) is a small fly with a length of 2.8—4.5 mm, with a remarkable

silvery white frons and brownish fore tibiae (see for a description Robineau-

Desvoidy (1830), Stenhammar (1844), Grunberg (1910), Becker (1896, 1926),

Wahlgren (1927), Séguy (1934) and Dahl (1959)). It can be found in the flowers and

on the floating leaves of the nymphaeids mentioned; the fly is not adapted to walk

or stay on the water surface, so that they have to fly from leaf to leaf.

To study the changes in numbers during the season an insect-lime method was

used. Each week, from June 16th until October 1977, six undamaged floating

leaves (five of Nuphar lutea and one of Nymphaea alba) of the same size were

gathered in the Oude Waal, pond F, and covered with insect-lime (Tangletrap or

Stikem), which has no odour and consequently does not attract or repel the

insects.

These leaves, the petioles of which were removed, were stuck on six other

Fig. 2. Lime-trap for insects. For explanation, see text. The wire netting is kept floating by corks and is

attached with a string to the petiole.

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

floating leaves and then protected from predation by birds by wire netting, with

meshes of 3 cm, the netting being supported by cork floaters (fig. 2). The wire

netting with cork floaters was attached with a string to the petiole of the

supporting leaf. After 24 hours these leaves were taken to the laboratory in the

wire netting in plastic bags, so that the catch remained as undamaged as possible.

The fauna was washed from the leaves with refined petrol and then fixed in 70%

alcohol. The leaves were laid in transects from the littoral border vegetation to the

open water, so that the distribution of the animals within the nymphaeid zone

could also be studied.

Three Nuphar leaves were laid in the western part of pond F; leaf | near the

broad littoral border vegetation, consisting of Typha angustifolia L., Rumex

hydrolapathum Huds. and Scirpus lacustris L., here the floating leaves of Nuphar

lutea covered up to 60%; leaf 2 six metres from leaf 1 in the centre of the Nuphar

zone (maximum coverage 70%); leaf 3 twelve metres from leaf | at the margin of

the Nuphar zone with the open water, where Nuphar covered up to 60% of the

surface. Two leaves were laid in a narrow Nuphar zone at the eastern side of pond

F; leaf 4 near the littoral helophyte vegetation, which borders on grassland,

consisting of a narrow zone of the same plant species mentioned for the western

side, but also with Mentha aquatica L. (maximum coverage of Nuphar 50%) and leaf

5 at three metres distance from leaf 4 at the margin of the Nuphar zone with the

open water (maximum coverage 40%). Leaf 6, a Nymphaea alba leaf, was laid in an

isolated vegetation of N. alba, surrounded by open water and at six metres distance

from the littoral border vegetation at the southern side of the pond (maximum

coverage of Nymphaea 85%).

The development of floating leaves and flowers was followed in two plots of one

square metre, one of Nymphaea alba and one of Nuphar lutea, as described in Van

der Velde (1978). Temperatures of air and water were measured regularly.

OCCURRENCE IN TIME

The results with the insect-lime method are summarized in table 1. Among the

total catch of approximately 23,158 specimens (of which 99.4% insects, the rest

arachnids), 642 specimens were of N. brunnipes, i.e. 2.8% of the total catch (see for

some other results Van der Velde (1978)).

Another Notiphila species, N. dorsata Stenh., was caught in equally large

numbers (764 specimens = 3.3% of the total catch). This species has a goldish-

yellow frons, so that it cannot be confused with N. brunnipes. N. dorsata was not

observed in the flowers of the nymphaeids and according to Dahl (1959) this

species is more or less characteristic for the helophyte zone, although he also

observed N. dorsata in the nymphaeid zone.

In fig. 3 the water and air temperature, the number of floating leaves and flowers

of Nuphar and Nymphaea per square metre, and the catch of Notiphila per week are

plotted against time.

The highest number of N. brunnipes was caught on July 28th when flowering of

Nymphaea and Nuphar was at its peak.

Adults of N. brunnipes have been caught by us with a hand net from the floating

109

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes

LG Sd DIS 6 6 IRE ep 6 ctl sse azsqunu Te3olL

= SUL 0 = = = = = = XT "67

89°0 18S v = = “i = I € XT EZ

ELO GENE I = = > = I = XT GNT

87°0 Sen! 6 = = = = 8 9 XT°8

GTI Wain I Ja I G = S 9 G XT TI

TSO OS IE L T = I = G € TITA"SC

OBERE GSM WS, = = = = @ 6 È TITA*8I

81°? S/S Il 99 EG = = I 5, ce TITA*ZI

SIE O8ST OCT Sal Ji G = 8 v9 TITA"G

68" 6 vec (al 6 G DI = ONE 68 TTA’ 8¢

BOE [EGG T6 6 v DI T S DI TTA‘ TC

Dore oh Is C9 CE = 9 G WG OT TIA*ZI

BETZ DECC Lg CA C 9 = GG 61 TTA’ 8

WE gal SGI TE = = € = = gl TA" OE

Gm) Sy lols G @ = = = = = TA" EC

200 TOPT IL - - = = = I TA‘OT

Yoel Te302 FO yo eo 9 S ig € È i

sodruuniq *N 2 182303 IRÉTOE Ioqunu zeer

qubneo sedruunzq *N Jo szoqunu LL61 :23eq

‘(1X9] aas ‘uoneuejdxa 19Y]IN} 107) Jwıf-J998u1 YUM Sadiuunag DjIYdION JO Say9182 ‘| [QRL

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

off

2 e—e water temperature

22 =—- air temperature

20 je

è EN

OUDE WAAL gi N À N

(NA Nymphaea alba

eee

40 OUDE WAAL ee

36 Nuphar lutea

32 OUDE WAAL

I Notiphila brunnipes

[| Notiphila dorsata

M J J A

1977

S 0 N D

1977

Fig. 3. The course of water and air temperatures, the development of floating leaves and tlowers ol

Nymphaea alba and Nuphar lutea in two plots of | m?, and the numbers of Notiphila species caught per

week with insect-lime on the floating leaves on the Oude Waal in 1977.

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 111

leaves and flowers of nymphaeids between 5 June and 29 September 1975, from |

June until 29 August 1976 and from 31 May until 22 September 1977. Wahlgren

(1927) mentioned the months July and August as the time of occurrence.

Discussion. — From fig. 3 it is clear that the development of numbers of the

N. brunnipes population is highly correlated with the development of the floating

leaves and flowers of the nymphaeids mentioned; water and air temperature, the

number of floating leaves and flowers and the number of adult N. brunnipes have

their maximum in the same period. N. dorsata, however, has its maximum on the

floating leaves when decomposition dominates over production.

From the data obtained with the insect-lime method it can be suggested that

both Notiphila species have only one generation per year.

With respect to the insect-lime method one should keep in mind that the catch is

not only dependent on the size of the population but also on changes in the activity

pattern of N. brunnipes, for example caused by weather conditions. This also

applies to the other insects, so that the percentages of N. brunnipes with respect to

the total catch per date have been calculated. The absolute numbers and the

percentages show the same trend, so that it is very possible that the catches give a

real picture of the development of the numbers in time.

OCCURRENCE IN SPACE

The general distribution of the flies over the sampling leaves within the

nymphaeid zone can be read from table 1, by comparing the results from the

different leaves.

The flies appeared to have a distinctly higher abundance on the floating leaves

bordering the littoral helophyte vegetation (leaves | and 4); the lowest number of

flies has been caught towards the open water (leaves 3 and 5).

Adults of N. brunnipes also showed to have a higher abundance on the broad

Nuphar zone (leaves 1, 2 and 3) in comparison with the narrow one (leaves 4 and 5).

Relatively more specimens of N. brunnipes were caught on the Nymphaea leaf (leaf

6) which is comparable with the Nuphar leaves 3 and 5 because of their similar

location near the open water.

In the Oude Waal the flies were found abundantly on the floating leaves and in

the flowers of Nymphaea alba, Nuphar lutea and Nymphoides peltata and less so in

the littoral border vegetation on Typha angustifolia, Rumex hydrolapathum and

Scirpus lacustris. In the Haarsteegse Wiel they were found on the floating leaves

and in flowers of Nuphar lutea and Nymphaea candida and seldom on other plants

such as Acorus calamus L. and Iris pseudacorus L. Our data are in accordance with

those of Dahl (1959) who studied the ecological distribution of Ephydridae in

Scandinavia; he found that WN. brunnipes is characteristic for nymphaeid

vegetations and occurs in lesser amounts in the bordering reed vegetation.

Discussion. — The higher abundance of adult N. brunnipes on the nymphaeids

bordering the helophyte vegetation may be explained by the fact that the

environment is less dynamic (wind- and wave-action) close to the reed-belt than

112 T1IDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

ruil

Fig. 4. a, thorax of N. brunnipes with attached pollen of Nymphaea alba; b, enlarged part of a. Note the

many curved hydrophobous hairs on the body which prevent the insect from getting wet.

close to the open water. N. brunnipes is, as already mentioned, not able to walk

over the water surface, and after a longlasting contact with water it drowns in spite

of having numerous curved hydrophobous hairs all over the body (fig. 4).

GENERAL BEHAVIOUR

Generally the flies walk slowly over the floating leaves and fly only short

distances to reach other floating leaves or flowers in the neighbourhood. Under

warm, sunny weather conditions the flies are more active and fly away sooner than

under colder circumstances when they can be captured rather easily by putting a

glass tube over them. On a sunny afternoon in July the flies can be found in large

numbers on the floating leaves and flowers where they perform several activities

like feeding, walking, courting, copulating, polishing, sunning, and depositing

eggs.

Under bad weather conditions, for example, heavy rain, they seek shelter in the

various flowers, under the aerial leaves of Nymphaea alba (leaves of this species

often project entirely or partly above the water surface) and between the littoral

border vegetation, staying there more or less motionless.

De Meijere (1940b) reports that N. brunnipes often forms dense clusters on the

leaves of Nuphar and Nymphaea; the specimens are then situated in a circle with

their heads directed to the centre. This behaviour, however, has been noticed by

us only sporadically, perhaps because we made observations during day-time while

De Meijere (1940b) describes this behaviour for the evening (20.30 h.).

Dahl (1959) has expressed the frequency of a number of activities of the flies in

percentages, viz., feeding 8%, polishing 25%, walking 42%, copulating 8%, no

movement 17% (the rin circumstances such as temperature and

weather conditions and time of the observation were not mentioned, however).

VAN DER VELDE & Brock: Life history of Notiphila brunnipes 113

FEEDING BEHAVIOUR

According to Waitzbauer (1976) the species of the genus Notiphila generally feed

on fluids of decaying plant material. Adults of N. brunnipes were observed by us

obviously feeding on fluids of decaying floating leaves but also on exudates of

flowers of the nymphaeid species mentioned, and on the body fluids of dead

insects, such as Ephemeroptera, Trichoptera and aphids.

In the flowers of Nuphar, Nymphaea and Nymphoides the flies could be seen

licking the anthers (even unripe ones) and the stigma. The flowers of Nymphaea

alba and Nymphaea candida do not possess nectaria but on the first day of flowering

the stigma profusely produces a sweet-tasting exudate containing glucose and

fructose as could be proved by enzymatic tests. On the second day of flowering

this exudate has already disappeared. The flowers of Nuphar lutea possess nectaria

on the outer sides of the petals, which produce nectar, also containing glucose and

fructose. The production of nectar occurs especially on the first day of flowering;

the flower then has a strong brandy scent while the stigma is sticky or dry and

shiny; an extensive description of the development of the flowers of the

Nymphaeaceae mentioned is being prepared (Van der Velde, in prep.).

The flowers of Nuphar and Nymphaea have also many anthers which produce

abundant pollen. N. brunnipes can often be found with pollen on its body (fig. 4).

To check whether the flies also consume pollen the intestines of some of them

were studied under the microscope. In some intestines germinated and

ungerminated pollen of Nymphaea alba could be recognized together with diffuse

material and micro-organisms.

The flowers of Nymphoides peltata possess nectaria which are sheltered by hairy

staminodes. N. brunnipes can not reach these nectaria because of the staminodes,

although efforts of these flies to reach the nectar can be observed regularly.

Furthermore the flies can be observed licking the five anthers and the small stigma

of Nymphoides, which is papillate and wet according to Heslop Harrison &

Shivanna (1977).

This licking behaviour strongly suggests that food is taken up in the form of

nectar, stigmatic exudate and pollen in the flowers.

On the floating leaves, N. brunnipes can be seen licking on decaying parts, e.g.,

on decayed tissue around the tracks of the mining larvae of Hydromyza livens

(Fall.) (Scatomyzidae) in Nuphar leaves. Fungi play an important role in the

decomposition of the leaves of nymphaeids. By feeding on the fluids of leaf parts

infected by fungi the flies can be vectors of these fungi, e.g., via the passage of

spores in the digestion canals or by chance presence of spores on their body.

According to Fischer & Gäumann (1929) small flies such as Drosophila species

can be vectors of the spores of Glomerella cingulata (Ston.) Spauld. et Schrenk,

which is known from many plant species and also occurs on the floating leaves of

Nymphaea alba and Nymphaea candida; this may well apply to N. brunnipes and also

to N. dorsata.

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

SEXUAL BEHAVIOUR AND SEX RATIO

The flies can regularly be observed in copulation on the floating leaves and in

the flowers of the nymphaeids mentioned. Many of the flies which are found on

and in first-day flowers of Nuphar and Nymphaea are in copulation; sometimes

masses of the flies were present on these flowers.

N. brunnipes shows a distinct mating behaviour. This behaviour was described

by Dahl (1959) as follows: “During the posturing phase of courtship the male

approaches the female from behind and after performing encircling movements

around her with his head directed towards her, he stops in front of her head,

tapping her antennae with his own. After this phase the partners have been

observed slowly circling round face to face. If the female accepts the invitation,

she spreads her wings permitting the male to mount. During the insemination the

female continues feeding and walking”.

It appeared from our field and laboratory data that the mating behaviour as

described by Dahl (1959) is not always so elaborate. Sometimes the female spreads

her wings immediately when a male approaches her and so copulation is allowed

directly. It also has been observed that a male approached a female in a more

aggressive way leading to copulation.

—

Fig. 5. a, abdomen of N. brunnipes, male; b, abdomen of female; ventral aspects.

Males and females of N. brunnipes can easily be separated from each other

because they differ in size and in the position of the genital aperture. Males have a

length of 2.8—3.7 mm, so they are smaller than the females which measure

4.0—4.5 mm in length (10 specimens of each sex were measured). The genital

aperture is situated on the ventral side of the abdomen in the males and at the

caudal tip of the abdomen in the females (fig. 5).

Some samples from the insect-lime catch on the floating leaves were studied for

calculating the sex ratio. The results are summarized in table 2. More males than

females were caught on the floating leaves in July and more females than males in

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 115

Table 2. Sex ratio of Notiphila brunnipes in samples caught with insect-lime.

Date: 1977 number of males females sex ratio

specimens percentage

examined of males

30. vi 14 5 9 3527

2 Vd. 18 17 1 94,4

DVI 53 Sal 22 58.0

2B.vii 66 44 22 66.7

SI 12 33 39 45.8

MND ded 1 43 21 22 48.8

Yrs gar L 45 19 26 42% 2

Total STE 170 141 5459

June and August. Perhaps the females occur more often in flowers during July

when also flowering is at its peak; the deposition of eggs in flowers can be

mentioned in this respect. Another explanation can be that males are more active

in this month than the females because of their mating behaviour.

DEPOSITION OF EGGS

As described by Rousseau (1919) and De Meijere (1940a, b) the females of

N. brunnipes deposit eggs in the flowers of Nymphaea alba and Nuphar lutea; we

found the eggs also in the flowers of Nymphaea candida ancally observed them,

during the dry summer of 1976, in the flowers of Nymphoides peltata, further on

floating leaves of the various nymphaeids of which the margins were curled up and

desiccated. The eggs are especially deposited in first-day flowers. The flowers of

Nymphaea candida in the Haarsteegse Wiel contained relatively few egg clusters

because these flowers were very often half-filled with water (Van der Velde et al.,

1978).

When the numbers of N. brunnipes, and also flowering, are at their peak in the

Oude Waal, nearly all Nuphar (when intact) and Nymphaea flowers contain egg

clusters. The number of eggs per flower varies with the population size from year

to year; in July 1975 hundreds of eggs could be found per flower of Nymphaea alba

and Nuphar lutea. The maximum number of eggs found in an ovarium of a

N. brunnipes female was 24.

According to Miller (1921) Notiphila riparia Meigen also deposits its eggs on

Nymphaea (also cited by Hennig, 1943, and De Meijere, 1944), but this has not

been confirmed in the present study. With the insect-lime method only one

specimen of N. riparia was caught on a Nuphar leaf. N. riparia is more

characteristic for a reed vegetation according to Dahl (1959).

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

MORTALITY AND PREDATION

Especially after heavy rain the population of N. brunnipes seems to be

diminished. The flies drown easily when they become wet for a long time.

Fig. 3 shows that the numbers of N. brunnipes decrease at the end of the summer.

Then mortality of adults may have several reasons, e.g., competition with

N. dorsata, falling temperatures, and lack of food from flowers.

Although predation on the flies has not been observed by us, the following

species must be taken into consideration with respect to possible predation: the

spider Pirata piraticus (Cl.) (Lycosidae), some of the larger Dolichopodid species,

the fly Hydrophoria conica Wied. (Muscidae), some species of Gerridae, Odonata

(Coenagrion pulchellum Van der Linden, Erythromma najas Hansem., Ischnura

elegans Van der Linden) and the green frog Rana esculenta L. It has been observed

that specimens of Hydromyza livens approached N. brunnipes in an aggressive way,

but they were never consumed or killed by this predatory fly. In the flowers the

flies have a smaller chance to be caught by large predators such as Odonata.

DESCRIPTION OF THE EGGS

The eggs of N. brunnipes have a length of 1.1—1.3 mm and a width of 0.4—0.5

mm (fig. 7); they are white in colour and possess a short toadstool-like micropylar

protuberance on the rostral end. The other end of the egg is rounded (fig. 6).

The eggs are deposited in the flowers in rows of usually 2—7 eggs but larger

rows are not exceptional. Usually the eggs can be found on the bases of the petals,

sepals and even on the anthers of Nymphaea alba, Nymphaea candida and Nuphar

lutea. The eggs are attached to the substrate, but not very strongly so; they are,

however, very well attached to each other, in such a way that it is impossible to

separate the eggs without damaging the chorion.

Eggs have been studied by means of scanning electron microscopy; in this way

regular netlike structures and pores could be recognized on the chorion (fig. 6).

The eggs of N. brunnipes show a striking bipolarity, because they possess different

structures on the dorsal and ventral side. The structure on the dorsal side can be

described as ‘‘wall’’-structure, a structure in which concavities are flanked by

walls. The structure on the ventral side can be described as “‘island’’-structure;

here island-like structures are flanked by canal-like concavities. The “wall”-

structure was already described by De Meijere (1940a). On fig. 6 a possible glue

substance can be recognized at the margins of the “islands”; this side of the

chorion was fixed to a petal.

To study the internal structure of the chorion the eggs were sliced with a razor

blade and examined with the scanning electron microscope. The chorion appeared

to possess cavities which are in contact with the pores in the external walls of the

chorion (fig. 6). The surface structures of the chorion, described as ‘‘wall’’- and

‘“island’’-structure, coalesce on places where the eggs are attached to each other;

in this way the eggs support each other so that they maintain their position

between the floral leaves.

VAN DER VELDE & Brock: Life history of Notiphila brunnipes 117

25 pm 50 um

luz i #

Fig. 6. a, caudal part of N. brunnipes egg showing both structures; b, rostral part of the egg (note the mi-

cropylar protuberance and the beginning hatching split); c, wall-structure (dorsal); d, island-structure

(ventral): e and f, coupe of the chorion showing cavities and the attachment of two eggs by means of

both structures.

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

THE FUNCTION OF THE CHORION

The distinct structures of the chorion of the N. brunnipes egg were suspected to

hold an air layer which functions as a plastron. Hinton (1960, 1969, 1971) has

shown that structures such as a network of walls and pores on the egg surface have

the function to hold air, so that the egg can continue with respiration when

becoming wet, e.g. by a rain shower. According to this author a system of

hydrofuge structures can form the architecture for a permanent or unshrinkable,

physical gill which is called a plastron. An egg with a plastron can remain

immersed indefinitely and obtains the oxygen it requires from the ambient water,

provided that the water is well-aerated.

The flowers of Nymphaea close at night and when they close they sink somewhat,

to a third, under water, so that the eggs of N. brunnipes regularly become wet

(Rousseau, 1919; De Meijere, 1940b); the flowers of Nuphar do not close at night

and during flowering they are always above the water surface (Van der Velde, in

prep.).

When the eggs are held under water an air-film around them can be observed;

De Meijere (1940b) pointed out, however, that the eggs cannot tolerate a

longlasting contact with water, but in that case soon decay.

In the laboratory some experiments were carried out to study the development

of eggs with regard to the functioning of the air-layer in and around the chorion as

a plastron. For the experiments young first-day flowers of Nuphar and Nymphaea

were collected which contained eggs of N. brunnipes; the eggs in these flowers had

nearly the same age and thus could be regarded as being in the same

developmental stage.

For the first experiment four Nuphar flowers with eggs were placed in jars at

room temperature (25° C). Flower | was placed upright in a jar with water so that

the flower was some centimetres above the water surface; flower 2, was placed as

flower 1, but was sprayed two times per day with water; flower 3 was placed upside

down in aerated water, held in this position with wire netting; flower 4 was treated

as flower 3, but the water was not aerated. Furthermore some eggs were taken out

of a flower and placed in a dry petridish. Observations were made daily during 2

weeks.

After 4 days there were larvae in the water from the eggs in flower | and 2, after

6 days from those of flower 3. The eggs of flower 4 decayed instead of developing

into larvae. The eggs in the petridish also did not develop into larvae but

desiccated.

It can be concluded that eggs develop into larvae in a relatively dry environment

(flower 1), in a dry but temporarily wet environment (flower 2) and in well-aerated,

oxygen-rich water (flower 3). Eggs decay in water poor in oxygen (flower 4) and

desiccate in a dry environment outside the flower (petridish); probably the air

within a flower is humid enough to prevent desiccation. It can also be concluded

that the development of the eggs under dry but air-humid conditions goes faster

than in water and that the structures of the chorion hold an air-layer which

functions as a plastron as the eggs developed in well-aerated water, so that they

must have obtained oxygen from this medium.

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 119

Table 3. Development of eggs into larvae under three different conditions (for further explanation, see

text).

Time Number of larvae in jar

A B C

4 days 0 0 14

5 days 0 0 87

6 days 0 715 96

10 days 0 83 96

A second experiment has been carried out to study the development of the eggs

more quantitatively. Three-hundred eggs were collected from first-day flowers and

used for this experiment. In each of three flowers of Nymphaea alba 100 eggs were

deposited. In jar À a Nymphaea flower with eggs was held upside down in non-

aerated water by wire-netting, in jar B the same was done in well-aerated water

and in jar C the flower was placed upright (only its stem was in contact with water).

The experiment was carried out at room temperature (25° C) and the development

of the eggs into larvae was followed by daily observations until the remainder of

eggs was disintegrating. The results are summarized below (table 3).

This experiment confirms the results of the first experiment; furthermore the

number of eggs not developing into larvae is larger in water (B) than under air-

humid conditions (C).

DEVELOPMENT OF EGGS IN RELATION WITH TEMPERATURE

To measure the influence of temperature on the development rate of eggs an

experiment was carried out in the laboratory. In the field a number of first-day

flowers of Nuphar and Nymphaea were collected in which eggs had been deposited

by N. brunnipes. A cluster of eggs attached to the innerside of a sepal or petal was

placed in a petridish with a layer of water in order to maintain high humidity and

to prevent desiccation of eggs. The petridishes were placed in different climate

chambers which had constant temperatures of 5°, 10°, 14°, 18°, 20°, 24° and 30°

C, respectively. By daily observations the development of the eggs into larvae was

followed.

At temperatures of 5° and 10° C the eggs had not yet developed into larvae after

30 days. At 14° C larvae hatched after 13 days and at 18°, 20°, 24° and 30° C after

5,6, 4 and 6 days, respectively.

It can be concluded from the experiment that the eggs develop well and fast into

larvae at temperatures between 18—30° C.

Temperatures within this range indeed occurred in the flowers of Nymphaea alba

and Nuphar lutea, as measured in the field with the aid of a Wallac oy Universal

Thermometer (GST 32; probes Ni-101x and Ni-106). By day light, temperatures in

flowers can be several degrees higher than the surroundings, due to solar radiation

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

Table 4. Temperature measurements in °C in flowers and surroundings of Nymphaea alba and Nuphar

lutea.

Nymphaea alba Tw EL Aie Ts PET eels

Dates 23.01.1977 22 26.0 2320 24.5 4.5

Times 4 S00 hn. 22.30 | 207.12 ZITO Se

Sunny, unclouded 2155 26.0 28.4 25.0 3.4

ME) 26%. 1 29.4 De 4.9

22.5 26,1 29 ZAND Sl)

Dates sf Meth Sy, 2259 23.2 Aal 24155 1.0

Times 14.45 hy. 2350 23.6 228 DD Oren

Heavily clouded 23.0 2955 232" OEE) Chay,

2259 2502 2220 DAS 0.2

234 2250 2270 220 ied

Nuphar lutea

Date: 8,9711.1977 22.50 ZA DER: 2226 4.6

Time: „1.4.30 Rh. 2345 28,0 290 265 Sie es)

Sunny, unclouded 232 25,0 DE 244 Se

DS 26.0 DIES 25°00 0

234 26,5 280 2002 18

Date: VA 10977 1.9.0 79,4 1.8.8 73.8 0

mer Bla sa MOERS IK OG 1728 0.8

Heavily clouded OA IA Se di 18.4 Orn

IRON ee RITZ 1 18.0 One

109210 LOO 148%,0 ITE Os

Tw = water temperature, Tl = temperature upper surface floating leaf, Tf = temperature innerside

of the floral leaves, Ts = temperature outerside of the sepals of the flower.

(table 4). Table 4 also shows that under clouded weather conditions the differences

in temperature between the inner and outer side of flowers are not as spectacular

in comparison with those under sunny weather conditions.

A fast development of the eggs into larvae within 4—6 days also has to take

place under natural conditions. The flowers of Nymphaea alba and Nymphaea

candida flower 4—6 and 4 days, respectively, before they become definitively

closed (Van der Velde, in prep.); the larvae cannot leave the flower in case the

development should be longer. This is in accordance with the fact that adult

females of N. brunnipes deposit their eggs especially in young first-day flowers.

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 121

The flowers of Nuphar remain open after the flowering so that the larvae can

always leave.

MORTALITY AND PREDATION

The experiments prove, that the eggs cannot develop into larvae when they are

too long in water poor in oxygen; then the eggs soon decay.

In both study areas, the Coot (Fulica atra L.) consumes most of the petals and

sepals of the flowers of Nuphar lutea. In this way many eggs of N. brunnipes

attached to these floral leaves must be either consumed or lost into the water. The

oviposition substrate is thus strongly diminished in size. The floral leaves of

Nymphaea were seldom consumed by waterfowl. These facts might explain the

higher numbers of N. brunnipes adults and eggs on Nymphaea alba flowers in

comparison with Nuphar lutea.

A possible consumer of the eggs might be the fly Hydromyza livens, a regular

visitor of flowers of Nuphar lutea. In the laboratory this fly appeared to consume

the eggs but this could not be confirmed by field observations; it is difficult to see,

without disturbing them, what these flies are doing in the flowers. It is not known

whether there are other consumers of the eggs but in the flowers of Nymphaea alba

and Nuphar lutea some Braconid species have been collected, which are suspected

of this.

HATCHING OF THE LARVAE

De Meijere (1940b) described the hatching of larvae as follows (translated from

Dutch): “The eggs, on the time of hatching, were dark gray in colour and one saw

the larva moving in the egg-shell when using the pocket-lens. Then the egg-shell at

once tore cleft-like open on one end and the larva, which was situated with the

blackish headtop in front of the developed aperture, crept out. When that had

happened, the walls of the egg-shell came together again, so that a flat membrane

of white colour was left”. From the scanning photographs (fig. 6b) it is obvious

that the egg-shell splits open in the surroundings of the micropylar protuberance.

After the larvae have left the egg-shell through this opening, they also creep out

of the flower and drop themselves in the water to reach the bottom (see for a

description of the larvae, De Meijere (1940b), Hennig (1943), and further fig. 7).

HABITAT CHOICE OF THE LARVAE

When the larvae have reached thé bottom they make tracks in detritus and mud

as already described by De Meijere (1940b), which could be confirmed by us.

An experiment was carried out to test the response of the larvae to light.

Hundred larvae were placed in a petridish with water. The petridish was half

covered with aluminum foil and placed under a lamp. After an hour 87 larvae were

present in the covered half and only 13 in the uncovered half. Thus the larvae are

negatively phototactic and seek dark places.

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

The detritus, however, is poor in oxygen. It is known that larvae of certain

Notiphila species obtain oxygen from aquatic and littoral border plants; the larva

pierces its pointed caudal stigmata in the plant tissue and so oxygen is obtained

from the intercellular cavities (Grunberg, 1910; Varley, 1937; De Meijere, 1940b;

Berg, 1950; Oldroyd, 1964).

De Meijere (1940b) and Hennig (1943) recorded puparia of N. brunnipes from

the roots of Typha angustifolia and Typha spec.; we inspected various aquatic and

littoral border plants in May and July 1977 to find out if larvae or puparia occurred

on their roots or root stocks. The plant species Nymphaea alba, Nymphaea candida,

Nuphar lutea, Nymphoides peltata, Rumex hydrolapathum, Typha angustifolia,

Glyceria maxima (Hartm.) Holmb., fris pseudacorus, Acorus calamus, Phragmites

australis (Cav.) Trin. ex Steud. and Mentha aquatica were investigated.

In spite of investigations over two years we never found the larvae on roots or

other aquatic parts of Nymphaea alba, Nymphaea candida or Nuphar lutea, as also

recorded by De Meijere (1940b). Large numbers of puparia and some larvae,

however, have been found pierced with their stigmata in roots and short shoots of

Nymphoides peltata from the Oude Waal. A large number of puparia and full-grown

larvae (7—10 mm long) were found on the roots and root-stocks of Acorus calamus

in the Haarsteegse Wiel (Acorus does not occur in the Oude Waal and in the

Haarsteegse Wiel Nymphoides is very scarce). Furthermore a number of pupae was

found on the roots of Typha angustifolia at both localities. Some pupae have

hatched in the laboratory so that a definite identification of N. brunnipes was

possible. From the roots of Typha angustifolia also a specimen of N. dorsata

hatched.

It is clear that Nymphaea and Nuphar are important for N. brunnipes to complete

its life cycle, but also such plants as Nymphoides peltata, Acorus calamus and Typha

angustifolia.

In the Netherlands Nymphaea alba also occurs in oligotrophic, dystrophic

moorland pools. Here N. brunnipes also occurs although the above-mentioned

plants do not. Possibly Typha latifolia L. fulfills a similar function in these habitats.

Although larvae are observed having pierced their stigmata in roots of plants, it

is so far unknown whether the larvae remain attached to one root or plant-

individual only or whether they move from one plant to another. It is clear that the

larvae do not need to be always pierced with their stigmata in plant roots. They

may creep from Nymphaea or Nuphar to Nymphoides, Acorus or Typha; the distance

between these groups of plants can be several metres.

It is possible that the younger larvae can remain longer in detritus poor in

oxygen (skin-respiration) and creep larger distances than the full-grown ones

without having pierced the stigmata in plant tissue.

How long the development of the larvae takes is not exactly known; in spring

full-grown larvae can be found, so that N. brunnipes certainly hibernates as a larva.

Furthermore it is very likely that N. brunnipes has only one generation per year.

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 123

FOOD OF THE LARVAE

According to Dahl (1959) De Meijere (1940a) has written that the larvae of

N. brunnipes mine in the roots of Nymphaea alba, but, in fact, De Meijere (1940a, b)

denied this. His papers are written in Dutch, which may have caused the

misunderstanding. De Meijere (1940b) who studied the young larvae in petridishes

with detritus reports that some specimens showed a green or brownish mass in

their body and thus concluded that they had fed obviously on fresh plant material

or detritus; most of the larvae however were totally colourless, crept restlessly

around and did not feed.

To study the feeding habits of the larvae some experiments were carried out. A

number of newly hatched larvae were put in petridishes with water; green

filamentous algae (1), detritus from the Oude Waal (2), fresh submerged Nuphar

leaves (3) and decaying floating leaves of Nuphar (4), Nymphaea (5) and Nymphoides

(6) were offered as food. Only some larvae kept in the petridishes with detritus (2)

and decaying leaves (4, 5, 6) had material in their intestines after a day. Probably

the larvae feed on material from the detritus and decaying plant parts or on

organisms occurring there, such as bacteria, fungi and protozoa.

Under laboratory conditions the larvae, however, could not be kept alive for a

long time. They also tried to creep out of the petridishes via the condensation

water; lack of oxygen may have caused this behaviour.

Mortality and predation

When the larvae drop themselves in the water they have a great risk to be eaten

by fish; furthermore they can get lost when they creep in wrong directions, when

they find no suitable root for attaching and die by lack of oxygen.

PUPA

The puparia of N. brunnipes (fig. 7) have been described by De Meijere (1940b)

and Hennig (1943). As already mentioned, the puparia can be found pierced with

their stigmata in the roots, root stocks or short shoots of Nymphoides peltata, Acorus

calamus and Typha angustifolia, possibly in order to obtain oxygen from these

plants. The puparia remain attached to the same root.

Puparia were found in spring, thus N. brunnipes hibernates also as puparium.

After the hatching under water the flies must reach the water surface quickly; it is

thus favourable that the puparia occur in shallow water.

EFFECTS OF THE DRY SUMMER OF 1976

By the prolonged drought during the summer of 1976 the broad of the Oude

Waal became nearly completely dry and also the interconnections between the

ponds, so that the littoral border plants and especially Nymphoides peltata suffered

badly. In 1977 the coverage and vitality of N. peltata in the Oude Waal had strongly

diminished, in 1978 the vegetation had regained its vitality.

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

Fig. 7. Life stages of N. brunnipes. A, imago: B, eggs; C, larva; D, pupa (pierced in root).

As a result the population of N. brunnipes of 1977 remained very small in

numbers with respect to other years, which supports the suggestion that there is

only one generation per year.

GEOGRAPHICAL DISTRIBUTION

N. brunnipes has been recorded from Central Europe (Becker, 1926), Italy

(Torelli, 1922, according to Hennig, 1943), France (Séguy, 1934), Belgium

(Rousseau, 1919), The Netherlands (De Meijere, 1940a), North Germany, Silesia

(Grünberg, 1910), Great-Britain (Oldroyd, 1975) and Fennoscandia (Dahl, 1959).

Dahl (1978) gives the following rough distribution area in Limnofauna Europaea:

Iberian Peninsula, Eastern Balkans, Western lowlands, Central lowlands, Baltic

Province, England, Boreal highlands, Northern Sweden and Taiga. The second

author of the present paper observed the species also in the Republic of Ireland in

1978.

According to Dahl (1959) the species is not found north of the limes

norrlandicus (about the 16° C isotherm of July); from Finland there are only some

southern records. He writes that both the ecological and phenological distribution

is governed by the species’ affinity for Nuphar and Nymphaea, but the geographical

distribution of N. brunnipes does not appear to include the northern parts of the

Nymphaea and Nuphar area of Fennoscandia.

Our investigations have shown that N. brunnipes occurs during its life cycle in

two different habitats and is bound indeed to Nuphar and Nymphaea but also to

Typha angustifolia, Acorus calamus, Nymphoides peltata, and probably Typha

VAN DER VELDE & Brock: Life history of Notiphila brunnipes 125

latifolia. Both the Typha species, as well as Acorus and Nymphoides have a

distribution south of the limes norrlandicus (Hultén, 1950), which may explain the

similar distribution of N. brunnipes.

SUMMARY

As can be concluded from the data in this paper there are direct relations

between N. brunnipes and the nymphaeids Nuphar lutea, Nymphaea alba, Nymphaea

candida and Nymphoides peltata.

Adults of N. brunnipes visit the flowers of Nymphaea and Nuphar for:

(a) food in the form of stigmatic exudate, nectar or pollen;

(b) the deposition of eggs; the development of eggs is favoured by the

temperatures and the air-humid or alternately dry and wet environment in the

flowers;

such as Odonata;

(d) copulation site; especially first-day flowers attract many specimens and

copulation often occurs there;

(e) resting site, e.g. for warming up the flight muscles; because of the higher

temperatures their digestion probably goes faster inside the flowers.

The flowers of Nymphoides peltata have these functions only partly (not b). The

flies play an important role in the pollination of all the nymphaeids mentioned.

Adults of N. brunnipes occur on the floating leaves of Nuphar, Nymphaea and

Nymphoides for:

(a) food, viz., directly for consumption of fluids of decaying leaf parts and

indirectly to consume body fluids of dead insects;

(b) copulation site;

(d) shelter or protection under aerial leaves or leaf margins which have been

curled up and dried out;

(e) sporadical egg-deposition on sheltered sites. By their behaviour adults of

N. brunnipes can function as vectors of spores of fungi, occurring on the

floating leaves.

The larvae and puparia seem to be restricted to the roots of Typha, Acorus and

Nymphoides, and obtain oxygen by means of pointed stigmata from the

intercellular cavities. The larvae feed there most likely on detritus and decaying

plant material.

The fact that N. brunnipes is bound to at least two different plant species during

its life cycle may explain the geographical distribution of the species.

ACKNOWLEDGEMENTS

We are much indebted to Prof. Dr. C. den Hartog and Mr. J. Krikken for

critical remarks, to Dr. P. J. van Helsdingen and Dr. W. J. Kabos for their help

with the identifications of flies and to Mr. A. W. Dicke for making the scanning

photographs.

126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 5, 1980

REFERENCES

Becker, T., 1896. Dipterologische Studien IV. Ephydridae. — Berl. Ent. Ztschr. 41: 91—276, pls. 4—7.

, 1926. Ephydridae. In: E. Lindner (ed.), Die Fliegen der Paläarktischen Region 6 (1): 1—115. —

Stuttgart.

Berg, C. O., 1950. Hydrellia (Ephydridae) and some other acalyptrate Diptera reared from Potamoge-

ton. — Ann. Entomol. Soc. America 43: 374—398.

Dahl, R. G. S., 1959. Studies on Scandinavian Ephydridae (Diptera Brachycera). — Opusc. Ent., Suppl.

15: 1—225.

——, 1978. Ephydridae. In: J. Illies (ed.), Limnofauna Europaea. Eine Zusammenstellung aller die

europäischen Binnengewässer bewohnenden mehrzelligen Tierarten mit Angaben uber ihre

Verbreitung und Ökologie (2nd. revised ed.): 1—532 (pp. 482—484). — Gustav Fischer Verlag,

Stuttgart, New York. Swets and Zeitlinger, Amsterdam.

Fischer, E. & E. Gäumann, 1929. Biologie der pflanzenbewohnenden parasitischen Pilze: 1—428. —

Gustav Fischer, Jena.

Grünberg, K., 1910. Diptera, Zweiflügler. Heft 2a. In: A. Brauer (ed.), Die Susswasserfauna Deutsch-

lands, eine Exkursionsfauna: 1—312. — Gustav Fischer, Jena.

Hennig, W., 1943. Übersicht über die bisher bekannten Metamorphose-stadien der Ephydriden, mit

Neubeschreibungen nach dem Material der Deutschen Limnologischen Sunda Expedition

(Diptera: Ephydridae). — Arb. morphol. taxon. Ent., Berlin-Dahlem 10 (2/3): 105— 138, 2 pls.

Heslop-Harrison, Y. & K. R. Shivanna, 1977. The receptive surface of the Angiosperm stigma. — Ann.

Bot. 41: 1233—1258.

Hinton, H. E., 1960. How some insects, especially the egg stages avoid drowning when it rains. — Proc.

S. Lond. ent. nat. Hist. Soc. 1960: 138—154.

—, 1969. Respiratory systems of insect egg shells. — Ann. Rev. Ent. 14: 343—368.

—, 1971. Polyphyletic evolution of respiratory systems of egg shells, with a discussion of structure

and density-independent and density-dependent selective pressures. In: V. H. Heywood (ed.),

Scanning Electron Microscopy. Systematics and evolutionary applications. The Systematics

Association. Special Volume No. 4: 1—331 (pp. 17—36). — Academic Press.

Hultén, E., 1950. Atlas över växternas utbredning i Norden. Fanerogamer och ormbunksvaxter:

1—512. — Generalstabens Litografiska Anstalts Forlag, Stockholm.

De Meijere, J. C. H., 1940a. Hydromyza livens Fall. en Notiphila brunnipes Rob. Desv., twee dipteren

wier levenswijze verband houdt met Nymphaea alba L. — Ent. Ber. (Amsterdam) 10 (232):

2202922;

——, 1940b. Over de levenswijze van Notiphila brunnipes Rob. Desv. — Ent. Ber. (Amsterdam) 10

(236/237): 281—287.

——, 1944. [1946] Zevende supplement op de nieuwe naamlijst van Nederlandse Diptera van 1898

(Eerste supplement op mijne naamlijst van 1939). — Tijdschr. Ent. 87: 1—25.

Muller, G. W., 1921. Insectenlarven an Wurzeln von Wasserpflanzen. — Mitt. naturw. Ver. Neuvor-

pommern Rügen 48/49: 121.

Oldroyd, H., 1964. The natural history of flies: 1—323. — Weidenfeld and Nicholson, London.

, 1975. Ephydridae. In: G. S. Kloet & W. D. Hincks (ed.), A check list of British insects, 11 part

5. Diptera and Siphonaptera: 1—139 (pp. 83—85). — Stockport, England.

Robineau-Desvoidy, A. J. B., 1830. Essai sur les Myodaires. Inst. de France (Cl. des) Sci. Math. et

Phys., Acad. Roy. des Sci., Mém. présentés par divers Savants (ser. 2) 2: 1—813. — Paris.

Rousseau, E., 1919. Notes biologiques I. Notiphila stagnicola Sten. (Diptére). — Bull. Soc. ent. Belg. 1:

101—102.

Seguy, E., 1934. Dipteres (Brachycères) (Muscidae, Acalypterae et Scatophagidae). — Faune de Fran-

ce 28: 1—832 + pls. 1—27. Paris.

Stenhammar, C., 1844. Försök till gruppering och revision af de svenska Ephydrinae. — K. Vetensk.

Akad. Hand. (ser. 3) 1843: 75—272, | pl.

Torelli, B., 1922. La Notiphila chamaeleon Becker e sue larva rinvenute nel laghetto craterico degli

Astroni. — Ann. Mus. Zool. R. Univ. Naples (H. 2), Suppl. Fauna Astroni 9: 1—6, | pl.

Van der Velde, G., 1978. Structure and function of a nymphaeid-dominated system. — Proc. EWRS Sth

Symp. on Aquatic Weeds, 1978: 127—133.

VAN DER VELDE & BROCK: Life history of Notiphila brunnipes 127

—., Observations on the floral biology of Dutch Nymphaeaceae. — Aquat. Bot. (in prep.).

——., Th.C. M. Brock, M. Heine & P. M. P. M. Peeters, 1978. Flowers of Dutch Nymphaeaceae as a

habitat for insects. — Acta Bot. Neerl. 27 (5/6): 429—430.

Varley, G. C., 1937. Aquatic insect larvae which obtain oxygen from the roots of plants. — Proc. R.

Ent. Soc. Lond. (A) 12 (4—6): 55—60.

Wahlgren, E., 1927. Svensk Insektfauna. 11. Diptera. Cyclorapha. Schizophora. Fam. 21—26:

323—416. — Stockholm.

Waitzbauer, W., 1976. Die Insektenfauna männlicher Blütenstande von Typha angustifolia. — Zool.

Anz., Jena 196 (1/2): 9—15.

7 DLL

privata

Dir

oii TE

cen).

CELSO

| DEEL 123 AFLEVERING 6 1980

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

M. A. LIEFTINCK. — Prodrome to a monograph of the Palaearctic species of the

genus Melecta Latreille 1802 (Hymenoptera, Anthophoridae), p. 129—349,

text-figs. 1—359, pls. 1—8.

Tijdschrift voor Entomologie, deel 123, afl. 6 Gepubliceerd 31-X-1980

PRODROME TO A MONOGRAPH OF THE PALAEARCTIC

SPECIES OF THE GENUS MELECTA LATREILLE 1802

(HYMENOPTERA, ANTHOPHORIDAE)

M.A. LIEFTINCK

“Kalliste”, 3911 MS Rhenen

With 359 text-figures, 8 plates and 3 maps

SUMMARY

Melecta is a parasitic melectine bee genus of the Anthophoridae, with a Holarctic distribution. In the

Palaearctic region it is widely distributed over north temperate Eurasia, all round the Mediterranean

basin, and the Canary Islands, off the west African coast. On the African continent Melecta does not

occur south of the Sahara. So far known, all members are parasitic on Anthophora Latr., s.str. and

immediate allies. As a prelude to a monograph, a revision is presented of the taxonomy, nomenclature,

and synonymy of all Old World taxa, based on — mostly new — characters. No subgeneric names are

proposed, and only two fairly definable units are recognized, the one here termed “the M. albifrons

alliance” being the only group eventually to be segregated from the rest as a nominate subgenus. The

essential taxonomic part comprises outline drawings of external structural details, compared to

characters given in descriptive keys provided separately for both sexes, 38 species and subspecies being

included in the key to the males, 26 for the females. All previously described taxa are re-defined, their

pertinent morphological features illustrated, while at the same time the nomenclature of some better

known species had to be drastically altered. The validity of 75 nominal taxa is taken into consideration,

all names being listed in the Index at the end of this work. Of a total of 67 names which are considered

validly proposed in Melecta, 17 are described as new, the remaining 50 comprising 21 new synonyms, 22

whose validity is analysed and re-established, and seven of uncertain status, the latter being discussed

separately in an Appendix. Distribution maps are given of nine species and subspecies, while 16 types

and/or paratypes are photographed. Regardless of their former or present status, the types of 46

previously named taxa were re-examined.

The systematic part is preceded by short non-taxonomic chapters containing brief notes on the

phylogeny of the Melectini and their general distribution, followed by some biological aspects, viz. the

typically vernal occurrences and bivoltinism, host/parasite relations, orchid pollination, and the

recently established (very rare) habit of collecting earthen substance, instead of pollen grains, at the

legs, exhibited by females of four species of Melecta, — presumably an atavistic phenomenon pointing

to common ancestry.

CONTENTS

ELECT RR Ree A GRE harte SE Han 130

Sommentsionspecimensiexamined et; Los. hee EG EREN ae NEN AEN 131

“SEES SUN ed Nn ae enden align mation aline te créant re ti 131

Some non-taxonomic observations on Melectine bees ........................... 132

Einserelations,origin andidistribution ent. n ene nen ese ann 132

LEU Embden on e Ari os ee ee BAE EE 132

Melcetintand'archidipollinationi EIN ER CNE Da 133

KindscallectingifemalesiofMelectans sas bern OAS. Bus MDOT deu Na 135

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Acknowledgements) 25 ann NE ON sie eee me ate, A Ee an Sena ate a 135

Systematic Part © 4: 9 200 wos ace. erated) ete am one ME ie en Re CR ER 137

Phylogeneticiconsiderationsy Sa eN 137

Generic characters: ro MER MANN NN 138

The'type-species of Melectae st Anse NEA EEE 139

Material:and methods", 1, 2.0.0. en an RE NIN A NN 139

Museumicollectionsandnomenclature typestudies REC a: TIRI 139

Key.and group characters: a. zen et NE Se a ET TEE 140

Terminology” ne =... Gs Boete ener eere Bra LIA te eee 1 ee to à à à on - 141

Variation: Ahr RE 143

Type.designations „ror sa 00. Salon ee nn an nn a © ob CT TRE 143

Historicalidentificationse oe AR he eo eee 143

Preparation of copulatory organs, andillustrations tt ss) eee 144

Flower records 5 2 tn u. See te ae ee EEEN 145

Descriptive key to Palaearctic species of Melecta=Male TOSO 145

Descriptive key to Palaearctiespeciesiof Melecia Remales rn RIONE 172

TheM: albifrons Alliance. … as nun see SSSR ne RO RE 193

1? Group of M-albifrons (Borstet)! >.< egg oe Go et ee ER RE 193

2; GrouplofMyiduodecimmacilata (ROS) 209

IINR:emaininpiPalacaretiesspecies'andisubspecies..t ee EE 219

Appendix#Speciesincertaesediss 4.417 ni... Nee Syne esi ee gee 334

AdditionalireferenGes nyc sen wh kod Rte? dela neds koma at Ele bate NE 338

Index ma tragen hag se grees ak emee be ent dcr ogee ee 340

INTRODUCTION

It is generally understood that ever since the turn of the century, the systematics

and nomenclature of Eurasian species of Melecta had fallen into a state of utter

confusion. Nevertheless, most twentieth century writers, though possibly well

aware of what should have properly been done, failed to compare their novelties

with the (partly easily accessible) types of some 40 nominal taxa defined by earlier

authors. This has given rise to a steady increase of nondescript species,

meaningless varietal names, and an incongruously large amount of obvious errors

in nomenclature, — a burden which finally caused some students to resignedly

give up identifying and studying these attractive bees. Considering the need of

putting an end to a deplorable situation, I have ventured to undertake a revision of

this difficult genus. In order to accomplish this in a more or less satisfactory way, it

became soon clear that a critical taxonomic analysis and a revaluation of

characters were urgently needed by finding out new and more reliable features to

re-establish the identity of all regional members of the genus. At the same time an

attempt was made to unravel an astounding quantity of misidentifications, which

eventually led to a thorough check of the validity of all nominal taxa.

The work was started early in 1958, when the opportunity was given to study the

types of the earliest described taxa in European museums. Numerous

controversial specimens were recovered and borrowed on these occasions, while

valuable collections of private bee students could also be consulted. These studies

were continued, with many interruptions, and brought to an end at the close of last

year (1979). The considerable delay in completing the manuscript has been at least

somewhat advantageous to my wish for including also the yearly flow of recent

acquisitions received from friends and colleagues who had collected in little

LIEFTINCK: Palaearctic Melecta 131

explored countries. This material, particularly from the eastern Mediterranean

region and Middle East, comprised welcome additional examples of several

insufficiently known species, and even one or two novelties.

Comments on specimens examined

The revision is based on an examination of over 2,800 specimens, i.e.

presumably only 75% or even less, of all unnamed material still available for study

in museums which I have been unable to visit. Apart from that, most of the above

total investigated by me belong to common and widespread species. Not counting

old and new synonymies, approximately 43% of the remainder belong to species of

which not more than six individuals (both sexes) could be compared, whilst no less

than six species (three male & three female) are known only from unique type

specimens. Consequently, it will be clear that this review does not pretend to be in

any way exhaustive except, perhaps, regarding the synonymy of the regional taxa.

On the basis of the analytical survey now available, it will be necessary to continue

collecting on a much larger scale than has been done before, so that good series

also of the less known taxa can be placed into the hands of a future monographer.

The scarcity of Melecta in the field is well known, and there are many reasons

which have caused these bees to be relatively poorly represented in collections. In

the past, collecting was mainly done during the summer holiday months, when the

flight season of nearly all species is practically over. Scattered occurrence, short

duration of adult life as well as the difficulty of finding the nesting sites of

Anthophora, are also responsible for the rareness of Melecta as compared, to take

an example, with the species of Andrena and its cuckoo bee Nomada, both

represented throughout the Holarctic region by a multitude of species and

individuals. Taken as a whole, the latter can be found from earliest spring till late

autumn. Most Melecta, on the other hand, occur singly, skimming low over

herbage, visiting flowers, or patrolling old walls, stony roadside banks, etc., in

search of their hosts’ nest entrances.

The area surveyed

As the title implies, only the specific characters of representatives known to

occur in the eastern hemisphere are dealt with in the systematic part, that is to say,

the whole of Eurasia with the inclusion of the African countries bordering the

Mediterranean Sea north of the Sahara. The Canary Islands, an archipelago

situated some 100—300 km off the westcoast of North Africa, are usually all of

them considered to form part of the Palaearctic region; they have much in

common with the majority of other known taxa and for that reason are also

included. The same applies to a few species occurring just outside this somewhat

arbitrarily defined area, e.g. those found in the foothills on the south slopes of the

Himalayan range, and parts of eastern China as well. They are much alike north

(western) forms and exhibit features strongly suggesting Eurasian (non-tropical)

origin; their inclusion is based on the fact that the distribution area virtually

coincides with that of the host bees of the genus Anthophora s. str., and maybe

other units closely related to these.

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

SOME NON-TAXONOMIC OBSERVATIONS ON MELECTINE BEES

Host relations, origin and distribution

Thus far known, all Old World members of Melecta are parasitic on a great

variety of Anthophora species. Leaving out of account a number of smaller, mainly

subtropical Afro-Asiatic genera, each with only few species whose anthophorine

hosts are quite unknown, there are yet indications that the allied genus Habropoda

occasionally also serves as host. Apart from the records already summarized in my

former revision (1972), a few classic observations relating to this subject published

by apidologists of earlier generations (e.g. Friese, J. Pérez), will be brought up

again and briefly pointed out in the text under each species in question. Very little

of real importance can now be added to what is already known of the relation

between host and parasite in the temperate Eurasian fauna, but some recent

observations bearing on the subject are, where expedient, also indicated.

Some of the obstructions met with on trying to establish the intimate relation

between the host species of Anthophora and the Melecta parasitizing them, will be

exemplified in some detail under M. albifrons albovaria, one of the species

occurring in central Spain (p. 203). A few more instances mentioned in the text will

also illustrate the above difficulty.

Both Melecta and Anthophora have typically a Holarctic distribution, but the

described species of the latter far outnumber those of its parasite: there may be

anything up to eight times as many described Anthophora in the Palaearctic region

alone than there are known of Melecta today. Unfortunately, however, the

systematics, nomenclature, and distribution of the numerous Old World

Anthophora are still in a completely chaotic state, and there are no reliable

structural illustrations or even keys, — a situation up till now comparable with

Melecta. So, realizing the undeniable interdependence of these two genera, it

would be irrational to speculate on the origin and distribution of Melecta, not even

of its commonest species. In spite of that I have, of course, taken pains to put on

record all available data about the range of each and prepared three maps giving a

— undoubtedly preliminary — graphic impression of the distribution pattern of

nine taxa occurring round the Mediterranean basin (see maps 1—3).

For an interesting account of the behaviour pattern of the New World species

Melecta separata callura (Cockerell), for instance the mode of nest entry and

Oviposition in cells of the Californian Anthophora edwardsii Cresson, see Thorp

(1969).

Bivoltinism

It is well known to European bee students that the great majority of Eurasian

Anthophora (s. str.) as well as their suspected cleptoparasites of the genus Melecta,

are strictly vernal in occurrence, completing only one generation per annum. Yet

there appear to be some striking exceptions to this rule found in populations of M.

italica Rad., leucorhyncha taormina Strand, and also a few albifrons nigra (Spinola),

which occur in Italy, the large Tyrrhenian Isles and Israel. In October 1959, while

LIEFTINCK: Palaearctic Melecta 133

studying the rich material kept in the museum collections at Bologna, Rome and

Torino, I came across a limited number of carefully dated examples of typically

“spring species” of Melecta and several of Anthophora that had been captured also

during the late summer months and even in autumn and winter. ') Acknowledging

this, the dates of capture of all vernal species were noted down for later use. As far

as Melecta is concerned, the pertinent data were copied and suffixed ‘(sic)’ in the

census of material examined, under each species discussed in the present paper. It

seems worthwhile in the present context to summarize all late summer records,

from August on, in the following list. I noticed that most of these specimens were

in perfectly fresh condition and did not differ from conspecific individuals caught

during spring. Consequently, with the possible exception of host bees collected at

higher altitudes (Apennines, Sicilia), whose flight season evidently shifts into the

summer months and which are excluded, there is circumstantial evidence that

certain Anthophora and Melecta complete two generations in one year. Of course,

much more selective collecting will be necessary to corroborate the supposed

bivoltinism expressed in the list; but I hope that it may form a modest starting

point for future investigation and stimulate critical studies in the field by students

interested in the seasonal distibution and host-parasite relationship of these bees.

Melecta. — M. albifrons nigra: (Italy), 9, Castelvetro (Emilia), 18.viii.1885 & ¢

same loc., 7.x.1885. — M. italica: (Italy), 9, Castelvetro (Emilia), 12.viii &

7.x .1885, and ©, Rimini (Romanga), x .1893. (Israel), g Jerusalem, 10.viii. 1946.

— M. leucorhyncha taormina: (Italy), 9, Castelvetro (Emilia), 10.viii. 1886.

Anthophora. — A. canescens Dours: (Italy), ¢ 9 Pietracuta (Emilia), x .1893. —

A. crassipes Lep.: (Sicily), Mt. Etna, 1890 m, 4.viii.1948. See also Frey-Gessner

(1907), & 9, Wallis (Switzerland), g, vi-vii & 9 “Mitte October, zweite

Generation?” — A. dispar Lep.; (Sicily), g, Taormina, 200 m, 16. xii.1949 and

Torre Marica, 19.xii.1949. A notoriously early spring species, evidently of a

second generation. — A. acervorum (L.): (Sicily), g, Sicilia or., Torre Archirafi,

0—800 m, 9.ix.1948 (many also in spring of same year: evidently two

generations!).

Melectini and orchid pollination

Since about twenty years, professor Bertil Kullenberg and his staff have

reported on observations and field experiments carried out on the pollination of

the orchid genus Ophrys, chiefly in countries of the Mediterranean region

(Kullenberg, 1961). Several species in this genus, the flowers of which do not

secrete nectar, are now well known to be pollinated by sexually excited males of

various aculeate wasps and bees. Only males are involved, female aculeate

Hymenoptera never having been observed visiting Ophrys flowers. In 1976, the

first instance of a melectine bee thus attracted by these nectarless Ophrys flowers

was announced by the same authors in a well illustrated publication containing a

') Under favourable circumstances certain spring species in North Africa and Israel are recorded to be

on the wing already from mid January on.

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

picture of a male bee resting on the labellum of Ophrys reinholdii Fl. It was drawn

from a colour photograph taken in the island of Rodos and shows a pollinium

attached to the bee’s front (Kullenberg & Bergstrom, 1976, sub Melecta spec.).

This bee was later recognized as a male Eupavlovskia obscura simulatrix Lieftinck, a

near ally of Melecta and a common vernal species in Rodos and other

mediterranean islands. Among the many melectines examined by me for the

purpose of this paper, some more examples were found of Melectini having

pollinia attached to facial parts of the head, viz. one more Eupavlovskia male of the

same species, taken by myself on Rodos, and one Melecta, the holotype of M.

tuberculata spec. nov., collected in the same island and carrying a similar

adornment at its frons; and there are a few more examples of pollinia-bearing

Melecta listed in the text. Hence the participation of these parasitic bees in the

special pollination process, seems to be well established. Chemoreceptory stimuli

are undoubtedly responsible for these interactions, and we may assume that at

least in the cases presently recorded for Melecta, the olfactory sense organs on the

flagellar segments of the male antenna (unapparent, it is true, in the genus

Eupavlovskia!) are responsible for guiding the bee’s behaviour on this kind of

orchid flower. Kullenberg and his co-workers made it clear that the Ophrys flower

incites the different phases of male copulatory behaviour, from the approach flight

until the attempted copulation, although actual copulation can, of course, never

be accomplished that way. The male copulatory instinct is stimulated by the

perfume exhaled from the conspicuous labellum of the flower. The excitant males

were observed to assume such positions and attitudes on the labellum that the

pollinia are loosened and removed, so that pollination can be accomplished. As

explained by the authors, neither the approach flight nor the tactilely guided

movements of the bee on the labellum can be released without the olfactive

stimulation by the flower perfume. Evidently, several Ophrys flowers are

sometimes visited in succession by a single bee, as witness the extraordinary

specimen shown (in pl. 7 figs. 36—37) of M. tuberculata, a new species from the

island of Cyprus, which I found in the British Museum (Nat. Hist.) collection. This

bee carries a cluster of not less than eight pollinia on its forehead. The pollinia

attached to the left side of this specimen’s face were recognized by Dr. Kullenberg

as a species of Ophrys, the identity of the rest remaining uncertain. Just before the

completion of this article, he kindly sent me a male M. albifrons albovaria

(Erichson) together with an enlarged photograph of the inflorescence of Ophrys

sphecodes atrata (Lindl.) A. Meyer, with the bee in the act of “attacking’ one of

the orchid flowers. This male darted upon the flower three times in succession in

an attempt to copulate with the labellum, but just only once the attack was so

vigorous that the scene could be filmed during the whole performance and the bee

captured and named. The observations were made by Dr. Kullenberg in

Languedoc (S. France), near Montbazin (Her.), 1—5.v.1979, in an old olive and

almond-tree plantation. It is the third instance of an equal number of melectine

species attracted by the scent of an Ophrys labellum and stimulated to copulatory

attempts.

LIEFTINCK: Palaearctic Melecta 135

Mud-collecting females of Melecta

In connection with the above, attention may be drawn to a surprising discovery

made while examining the leg structure of many hundred Melecta females during

the present investigation of the morphology of these bees. In a limited number of

individuals (totalling only 14), belonging to four species, definite lumps of earthen

matter were found to be firmly attached to the external faces of the hind tibiae

and, more rarely, also at one or both of the mid tibiae, of the bees’ legs. This

substance is clayey, of a yellowish to dark grey-brown tint, and has, in the most

perfect state of conservation, a suboval or semicircular shape, closely resembling

an almost completed, though somewhat less conspicuous, pollen load at the

corbicula of a worker bumblebee. The bees dressed up in this way were collected

at random in seven countries and islands all round the Mediterranean. In one fe-

male which I took on Rodos island, all four posterior tibiae carried a conspicuous

load of these clods. In view of the solid consistence, modelling, and symmetry of

this matter, I am convinced that there can be no question of incidental defilement

of the legs resulting from digging activities inside the nest tunnels of the host. As

far as I know, this phenomenon has not so far been reported in the literature on

parasitic bees. One may well ask whether it has ever been observed also in

Psithyrus ? I venture to interpret this prodigious mode of behaviour as an atavistic

device exhibited by a melectine, thereby demonstrating its true kinship with the

non-parasitic anthophorine host: — a remains of the pollen collecting instinct of

the host bee inherited by its parasite and materialized in the form of a defunct

substitute? It is significant that no males are involved and that most females are

old, more or less worn specimens having tattered fore wings and partly rubbed off

pubescence. The nature of the earthen substance has not yet been analysed, but

this can, I hope, be done and the results published along with some photographs

showing the matter in situ.

ACKNOWLEDGEMENTS

I welcome the present opportunity to express my sincere thanks to all those who

have, since many years, provided me with specimens of Old World anthophorid

bees, — inspiring gestures, which have in no small measure contributed to the

decision of continuing a series of revisional studies on these fascinating bees. First

of all, I wish to acknowledge my gratitude for the courtesies received from

specialists and curators in charge of the Hymenoptera collections during my

visiting their museums and institutes all over Europe, initially to study types but

later mainly to compare fresh acquisitions with questionable specimens obtained

earlier. I am grateful also to many colleagues who gave me access to their personal

collections by sending me on loan specimens collected by themselves and

permitting me to retain a number of duplicates, ultimately to be deposited in the

Rijksmuseum van Natuurlijke Historie at Leiden (ML).

For the present survey I have again employed a series of abbreviations

indicating the ultimate location of the material studied, followed by the names of

all taxonomists and field collectors who have supplied material, and without whose

136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

generous assistance this study would not have been possible. To locate the

specimens examined, the same symbols are used in the text, placed in parenthesis

behind all data recorded in the opening paragraphs under the species headings

“Material”.

AID -— Beth Gordon Agriculture & Nature Study Institute, Deganya A, Israel (Y. Palmoni)

AMNH — American Museum of Natural History, New York (J. G. Rozen)

BM — British Museum (Natural History), London (G. R. Else & I. H. H. Yarrow)

CAV — Coll. E. Asensio de la Sierra, Valladolid

CB — Coil. J. de Beaumont, Auvernier

CBS — Coll. H. Bytinski-Salz, Tel Aviv

CCS _— Coll. A. Compte Sart, Madrid

CE — Coll. P. A. W. Ebmer, Linz/Donau

CFP — Coll. F. Parre, Bad Soden

CG — Coll. J. Gusenleitner, Linz/Donau

CGS — Coll. A. Giordani-Soika, Venezia

CHW — Coll. H. Wolf, Plettenberg

CJH — Coll. tJ. Heinrich, Aschaffenburg (now SMF)

CK — Coll. M. Kocourek, Vyskov, CSR

CKW — Coll. K. Warncke, Dachau

(GIL — Coll. W. Linsenmaier, Ebikon (Luzern)

CMC — Coll. M. Comba, Roma

CMS — Coll. Max. Schwarz (incl. coll. H. Priesner, CP), Ansfelden

CO — A.Z. Osychniuk, Institute of Zoology Akad. Sci. Ukr. SSR, Kiev

Cr — Coll. H. Teunissen, Oss

CTP — Coll. B. Tkalcu, Praha

CVS — Coll. F. Verges Serra, Canet de Mar (Barcelona)

CWG — Coll. W. Grünwaldt, München

CVZ — Coll. G. van der Zanden, Eindhoven

DEU — Department of Entomology, University of Uppsala (Bertil Kullenberg)

EIB — Institut f. Pflanzenschutzforschung, Eberswalde DDR (G. Morge & J. Oehlke)

FAG — Faculté des Sciences Agronomique de l'Etat, Gembloux (J. Leclercq)

IEB — Istituto di Entomologia della Università, Bologna (f G. Grandi)

IEM — Instituto Espanol de Entomologia, Madrid (Miss E. Mingo)

IEP — Istituto di Entomologia Agraria, Portici (R. Priore & E. Tremblay)

INER — Istituto Nazionale di Entomologia, Roma (M. Cerruti)

INRA — Institut National dl. Recherche Agronomique, Versailles (R. Desmier de Chenon)

IZK — Institute of Systematic & Experimental Zoology, Kraków (M. Dylewska)

MA — Instituut voor Taxonomische Zoölogie, Zoölogisch Museum, Amsterdam (J. P. Duffels)

MBUD — Hungarian Museum of Natural History, Budapest (J. Papp & L. Móczár, Szeged)

MC — Universitetets Zoologiske Museum, Kopenhagen (B. Petersen)

MCG — Museo Civico di Storia Naturale, Genova (Miss Delfa Guiglia)

MCNT — Museo Ciencias Naturales, Santa Cruz de Tenerife (P. Oromi)

MG — Musée d’Histoire Naturelle, Genéve (C. Besuchet)

MH — Zoological Museum of the University, Helsinki (P. Nuorteva & J. Teräs)

MIT — Museo Insular de Ciencias Naturales, Santa Cruz de Tenerife (M. Baez & Pedro Oromi)

MKB — Zoologisches Forschungsinstitut u. Museum Alexander Koenig, Bonn (t K. F. Buchholz)

ML — Rijksmuseum van Natuurlijke Historie, Leiden (C. van Heijningen)

MLLT — Museo Depart. Zoologia, La Laguna, Tenerife (M. Baez)

MMB — Moravské Museum, Brno, CSSR (J. Stehlik)

MNB — Museum für Naturkunde der Humboldt Universität, Berlin DDR (E. Konigsmann)

MP — Muséum National d’Histoire Naturelle, Paris (Mlle S. Kelner-Pillault)

MT — Museo ed Istituto di Zoologia Sistematica, Torino (O. Elter, Mlle M. Goss & U. Parenti)

MUC — Museo Zoologico da Universidade, Coimbra (M. de Asensio Diniz)

LIEFTINCK: Palaearctic Melecta 137

NMB — Naturhistorisches Museum, Basel (t+ F. Keizer & W. Wittmer)

NMP — Národni Museum V. Praze, Praha (Oldrich Sustera)

NMW — Naturhistorisches Museum, Wien (M. Fischer)

NRS — Naturhistoriska Riksmuseum, Stockholm (S. Erlandsson)

SMF — Natur-Museum Senckenberg, Frankfurt a.M. (D. S. Peters)

UK — University of Kansas, Lawrence (C. D. Michener)

USNM — National Museum of Natural History, Smithsonian Institution, Washington D.C. (P. D.

Hurd & K. V. Krombein)

ZIL — Zoological Institute, Academy of Sciences, Leningrad (M. N. Nikolskaja & Ju. A. Pesenko)

ZMM — Zoological Museum, Moscow (A. N. Zhelokhovtsev)

ZSM — Zoologische Sammlung des Bayerischen Staates, München (F. Bachmaier & F. Kühlhorn)

As this paper is based on about 60 private and institutional collections, I

regretfully admit not being able to express my indebtedness to any person in

particular; suffice it to say that I realise how much I owe to those who sent to me

their material for perusal and patiently awaited the completion of this paper, —

some of my correspondents, in fact, having been looking forward to the return of

their specimens for more than twenty years.

Unfortunately, two important collections containing important specimens of

melectine bees from parts of the Mediterranean basin and Middle East countries,

had to be left unstudied owing to their inaccessibility, viz., one formerly the

property of the late B. Pittioni, cursorily inspected by me several years ago but no

more available; and a second, probably substantial collection owned by a British

bee specialist who serenely left all appliances unanswered, being not prepared to

have his collection incorporated in the present work.

My best thanks are due to my good friends Dr. Max. Schwarz, of Ansfelden

(Austria), who kindly presented me with a copy of Rossi’s ‘Fauna Etrusca, 2’’ and

Dr. P. J. van Helsdingen, of the Leiden museum, for having carefully translated

some diagnoses contained in the same work; to Prof. Dr. J. T. Wiebes, of the

Leiden University, who provided the photographs of the pollinia-bearing male of

M. tuberculata in the British Museum (Nat. Hist.) collection; to Dr. I. H. H.

Yarrow, who supplied the fine picture of M. grandis in the same collection; and to

Messrs. Roman and the late Ch. Hoorn, photographers at the Leiden museum, for

the remaining images of whole insects reproduced on plates 1—8 of this article.

SYSTEMATIC PART

Phylogenetic considerations

As pointed out by J. G. Rozen (1969), the evolution of the subfamily Melectinae

is intricate and not easy to understand. In his account of the early stages of the

tribe Melectini, based on representatives of the New World, the author memorizes

first, that comparative studies on the morphology of the imagines have learned,

that the two major divisions of parasitic Anthophoridae, viz., the largest and most

complex group, Nomadinae on the one side, and the three melectine tribes,

Melectini, Ctenioschelini (Ericrocini o/im) and Rhathymini on the other, are of

138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

different origin and have evolved separately. This was confirmed by a careful

comparative study of the mature larvae of both groups. All members of the

Melectini have been found in the cells of anthophorine bees, whereas those of the

two other tribes depredate the nests of Centridini. An inquiry into the larval

morphology and anatomy of these cuckoo bees supports the view that the

Melectini evolved from the same lineage as their hosts, the Anthophorinae, or in

other words, have arisen from anthophorine ancestors, whereas the two other

parasitic tribes evolved from parental stock at the root of the Centridini. In short,

an investigation of the early stages implies a diphyletic origin of these tribes.

Rozen summarizes his thorough investigations in the following way: “The fact that

many parasitic bees have arisen from the same stock as their hosts, is true in many

instances and is a matter of significant interest to the evolutionist delving into the

relationships of bees”. At the same time he emphasizes that “It would be naive,

however, to apply this rule a priori in order to deduce the origin of parasitic bees”.

He then adds that this point is proved, for example, merely by the fact that larvae

of the nomadine genus Triepeolus (of the Epeolini) were recovered from the nests

of host bees representing four different families.

Generic characters

The difficulty of the subject and insufficient knowledge of New World forms,

prevent me from giving a full account of the generic characters. All Melecta

inhabiting the western hemisphere being excluded, I see no point in trying to

redefine the genus taken as a whole, the more so while the American taxa are

much more diversified and discontinuously distributed in the Nearctic parts of the

continent. They do not seem to have much in common with those occurring in

Eurasia, the set of characters displayed by the two units being of a different nature

and arranged according to a different pattern. Moreover, the New World

Melectini are still only summarily characterized structurally. They are far less

numerous than their Old World tribal members, comprising little more than 20

species and subspecies, which are nevertheless placed in four genera (including six

subgenera), though most taxa are assigned to the nominate subgenus. A useful

account of the Nearctic melectine fauna was published by Hurd (1953), with a

bibliography of the more important publications. Further details are summarized

in my 1972 review. The most recent and complete survey of the literature on the

American representatives of the tribe, is contained in vol. 2 (Apocrita) of the

“Catalog of Hymenoptera in America North of Mexico”, under the direction of

Karl V. Krombein and Paul D. Hurd, Jr. (1979, Smithson. Inst. Press, Washington,

D.C.: 2170—2172). In this catalogue four genera of Melectini are listed for the

United States, with a total of 18 species and subspecies, the majority (11)

pertaining to Melecta s. str. with one monotypic subgenus, the three remaining

genera comprising six species and subspecies.

Turning to the Old World group, diagnoses of the eight genera so far known are

to be found in two fairly recent revisional studies (for references, see Lieftinck,

1972, with a key to all genera and bibliography). As these definitions are not quite

complete and therefore still insufficient, it seems best for the present to leave a

LIEFTINCK: Palaearctic Melecta 139

comprehensive and really satisfactory review of all genera and subgenera of

Melecta for a future monographer, who then may be capable to deal with the whole

complex on a world basis, making use at the same time of some neglected

characters introduced in this paper for the definition of the Old World members.

The type-species of Melecta

As pointed out recently by Day & Fitton (1977), the first described and best

known Eurasian member of Melecta occupying a fairly extensive part of the

Palaearctic region is Apis albifrons Forster, 1771, a polytypic species. This

(inappropriate!) specific name should now replace the much more familiar names

applied to the same taxon, viz. Melecta punctata (Fabricius, 1775, et auct.) and M.

armata (Panzer, 1799). The first and earlier defined of the two, Fabricius’s Apis

punctata, became the type-species of the genus by designation of Latreille in 1810;

but since Apis punctata and Andrena armata are conspecific and recently turned out

to be synonymous with albifrons, the latter ipso facto takes the place of punctata as

the type-species of Melecta. Sure enough, the nominate subspecies, here re-

defined as M. albifrons albifrons (Forster), is the only well known regional member

of the genus.

MATERIAL AND METHODS

Museum collections and nomenclature, type studies

The names considered in this revision total 75. To economize space, all nominal

taxa are entered in a single alphabetical index at the end of the work.

Apart from five earlier synonymized forms, three homonyms, and 17 species

which are described as new, the identity of 50 nominal taxa is verified. Of the

latter, 38 were originally described as full species and 12 as varieties (or forms) of

either M. punctata (c.q. armata) or luctuosa (Scopoli, 1770). Three of the first

category, wrongly assigned to other genera, are transferred to Melecta. They are:

Nomada? duodecimmaculata Rossi, Pseudomelecta baeri Radoszkowski, and Crocisa

atra Jurine. Only the latter must be dropped as a synonym of M. albifrons.

Regarding the 50 specific and infraspecific (‘‘varietal’’) names, 21 are found to

be synonyms of taxa already known (marked ‘syn. nov.” in the text), and seven

species incertae sedis discussed separately in an Appendix at the end of the

descriptive part of this work. The status of the remaining taxa, amounting to 22,

was analysed in the course of the investigation, which proved all of them to be

validly proposed species and subspecies (see text). As to the fate of the “varieties”

I have, of course, taken the line that their names should be preserved as much as

possible by re-studying the types and reconsidering their validity, rather than

discarding them as so-called nomina oblita. This resulted in the recognition of only

three whose names are retained and applied to taxa given specific (two) or

subspecific rank (one), the nine others being relegated to synonymy and included

in the above total marked “‘syn. nov.”.

140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

It must be added that among the many bees described by O. I. Radoszkowski,

the types of 12 species of Melecta went to the Berlin museum (MNB). Between

1895 and 1905, these were traced and recognized by Günther Enderlein, at that

time one of the custodians in charge of the insect collections. Apparently

disinclined to inquire into the validity of Radoszkowski’s names, Enderlein merely

labelled them by copying the specific name, with the addition “rev. Dr. Enderlein.

Type”. As far as I know, a list of these has never been published. Therefore the

actual existence of these supposed types is confirmed in the present paper and

their identity substantiated by holotype and/or lectotype designations, — purely

formal statements, which can be found in the paragraphs “Type material” under

the heading of the majority of Radoszkowski’s species. Regardless of their former

status (specific of varietal), and including Enderlein’s tentative selections, 46 types

were re-examined.

Key and group characters

For the characterization of both sexes, about 45 morphological features were

listed and taken into consideration, 70% being unisexual and peculiar to the male.

Inevitably, even a number of these had to be disregarded, after all, as a means of

specific differentiation, partly by lack of sufficient comparative material of either

sex, but also because some important qualities of the integument are sometimes

hidden from view by the hairiness of the bees.

Much time and consideration has been devoted to construct workable keys. The

initial plan to offer simple dichotomic keys was given up almost from the outset.

With the discovery of new species, objectively useful external characters were also

detected, so that the keys underwent considerable alteration and repeated tests

until they were thought to be acceptable as a tolerably serviceable whole. Of

course, key characters, once established, can be arranged in various ways,

depending on subjective valuation. The best way to gain an impression of the

general appearance of each species proved to try out, for either sex, the most

characteristic features combined, thus defining them in a not too elaborate

descriptive way. The sexes are treated separately, the male being considered first

because it possesses many more reliable and easily observable characters than the

female, which is often difficult to identify even after exposing the pygidial plate or,

when necessary, dissecting out all terminal sclerites. Male and female characters

of the M. albifrons alliance are combined in the key to the males, for all other taxa

they are kept separate.

Group characters. — As with the nearly allied genus Thyreus, the characters

employed have not afforded much help in establishing more than two definite

species groups in the Palaearctic fauna. With the exception of the albifrons alliance

just mentioned, which constitutes a division of its own, the male genital organs

with their internal sternal plates, have proved to be valueless for this purpose

because shapes and setal arrangements, though always differing somewhat

interspecifically, are similar in principle, while nearly all other characters are

shared in such a haphazard way and allotted so differently in species showing a

similar pubescent pattern, that one is disposed to think that many features have

LIEFTINCK: Palaearctic Melecta 141

had an independent origin. These incongruities are well expressed in the keys, in

which the successive characteristics are not always enumerated in the same order.

This proved unavoidable, because body texture and vestiture — to take an

example — had to be at one time treated separately, but in another were

considered logically consistent and best discussed jointly.

In my last account of Old World melectines (1972: 264 —265), I called attention

to a rather aberrant group within the genus characterized by a general dullness of

the body integument caused by very dense punctation and short tomentum; in this

group also the first two abdominal segments are more nearly equal in length.

These unusual features have nevertheless been used in the key in spite of the fact

that, with new discoveries, some species are intermediate or fall outside any group.

Instances like this may be multiplied, e.g., with regard to the development of

strong bristles, impunctate areas on the vertex, depth of sutures, sculpture of

propodeum, and the like. It is obvious, therefore, that these keys are rather

artificial and only partly reflect true relationships. In no way do I pretend to have

found all potentially useful characters, but I do hope that those employed,

clarified by portraits of bees and as many sketches of structural parts as were

thought convenient, may serve as a basis for further investigation.

In the key to the males, the form of the gonocoxite of the genital bulb will be

introduced as one of the few fairly constant localized characters used as a means

of specific distinction; it needs a brief explanation. Viewed from above, the distal

halves of the gonocoxites, beyond the partly inflected blunt inner tubercle, are

somewhat divergent, their inner borders curving at first outward and then inward

in a specifically characteristic way, usually forming a broadly convex bend before

curving ventrad; this swollen part may be either rounded off, or projects more or

less distinctly inward: this is called the ‘‘gonocoxal angle” (fig. 6, g.a.). The distally

open cavity embracing the central genitalic organs bounded by the swollen

gonocoxites, is termed the “‘gonocoxal enclosure”. All remaining characters

employed in the keys will be easily understood by comparing the illustrations.

Since the keys also comprise characters not visible externally, the copulatory

organs should be dissected out and always compared with the figures. Many

species can be recognized with certainty only when fresh examples are available

whose body parts have been extended and made plainly visible. Specimens in full

dress are found only soon after emergence and then are usually easily recognized.

Aged females, however, may have lost much of their freshness and vestiture; they

are frequently met with carrying worn off mandibles, imperfect tarsal claws and

stubbed pygidial plates, — all characters of some importance reminding to careful

inspection before identifying them. Scores of females thus affected had to be left

unnamed: torn wing borders always sound a warning note.

Terminology

To avoid any misunderstanding, in the keys and further descriptions, the

antennal segments are numbered consecutively from 1 to 12 (13), thus including

the scape and pedicel, the latter being always plainly visible. For the sake of

142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

a ae eee

Figs. 1—6. Melecta, structures; 1, M. albifrons albovaria, posterior view of left antenna, showing impres-

sed sensoria (4 Montpellier, France); 2, M. luctuosa, the same, showing obliterated sensoria (4 Ne-

therlands); 3, M. albifrons albifrons, external view of transitional parts of hind tibio-basitarsus, semi-dia-

grammatic (4 Netherlands); 4, M. luctuosa, the same (4 France); 5, M. italica, dorsal view of genital

capsule, showing rounded gonocoxal angle (4 Grimaux, France); 6, M. festiva, the same, showing pro-

nounced gonocoxal angle ( same locality). Scale line 5 mm (figs. 1-2); g.a. = gonocoxal angle

simplicity I have dropped the somewhat ambiguous and rather superfluous terms

““mesosoma” (thorax + propodeum) and ‘‘metasoma” (or gaster), for the post-

propodeal part of the abdomen, and recurred to the regular practice of older

authors by using the traditional and more familiar words thorax and abdomen.

Consequently, by regarding the propodeum (= first abdominal segment in the

Apocrita) as part of, and in conjunction with, the thorax, the next abdominal

sclerites being here numbered from the first segment on. A reversion to the

original and simple terminology seems justified in purely descriptive articles like

the present one, the more so as the morphological segmentation of the thoraco-

abdominal complex is common knowledge and correctly interpreted already by

Latreille. The rudimentary 9th abdominal tergite (8th gastral) of the male, is a thin,

sub-membranous, more of less quadrangular, minute plate which, though always

present, is liable to be overlooked, for which reason its shape has been neglected

as a means of specific distinction.

All specimens enumerated, whose body parts had to be dissected out for close

examination and/or drawing purposes, are marked “(diss.)’” under the heading of

material studied.

LIEFTINCK: Palaearctic Melecta 143

Variation

Apart from the only too well known sexual dimorphism, strongly expressed in

structure, nature of pubescence and often also in wing colour and venation,

individual (infraspecific) variation is quite considerable, so much so in fact that

there is hardly any part of the body which does not participate in showing slight

differences in contour, shape or proportion, within a single population in any given

locality. As, for example, in the allied genus Thyreus (Lieftinck, 1962—1972) and

other parasitic solitary bees like the Japanese species of Nomada (Tsuneki, 1973:

5—10), this marked inconstancy even of the finer structures, such as the male

copulatory apparatus and hidden sternal plates, may greatly impede recognition,

especially where two or more near-alike species of Melecta occur together in one

locality. Added to that, the variation in the alternating black and white pubescent

pattern exhibited by both sexes, is pre-eminent and infinite, thus forming an

additional obstacle to the taxonomist in search of reliable distinctive characters.

Numerous examples of this instability will be found among the taxa treated in the

next pages, demonstrated also by outline drawings of the more eminent organs

preferably selected from a series of preparations showing the extremes (plus and

minus variants) observed in a single population of a given species or subspecies.

Wing venation also is a case in point: except colour of the fore wing membrane, no

reliance can be placed on the position and course of the cross nerves. Out of all

venational peculiarities considered, the only one at all helpful, i.e. as a group

character, is the shape of the third submarginal cell, which does seem to be fairly

constant, at any rate in the males.

Type designations

Except in a few cases, only holotypes and lectotypes are recognized. For

obvious reasons I have since many years renounced the qualification “allotype”,

also because in practice it has been too often used somewhat thoughtlessly, and as

a result of its deceptive usage led a persistent life in collections and literature in

cases of erroneous sex-associations. To avoid any misjudgement of the word

“type”, I have therefore substituted the term allotype by the non-pretentious

indication “First defined [male or female]”, which then only expresses my belief

that the newly characterized specimen of the opposite sex is conspecific with the

type. For the same reason paratype designations are avoided and selected only, for

convenience, from topotypical material, preferably of the same sex as the type. In

all other instances my own identification labels should, I hope, suffice as an

expression of my personal conception of a given taxon.

Historical identifications

Full information on localities, dates of capture, and collectors is, with very few

exceptions, given for all specimens examined, the personal names of those who

identified museum specimens in the past being also copied from the labels. These

names are printed in italics, on behalf of future students interested in the

144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

nomenclatural history of a given species, and also to make them acquainted with

the different views expressed by the older generation on the identity of their

specimens.

(It is surprising how very few entomologists seem to realise the inconvenience of

pinning identification labels in the same position to those indicating the locality

data, instead of simply attaching the name-label upside down on to the pinned

specimen. By so doing the vexation of being compelled to use one’s fingers or a

pair of forceps before succeeding to read a scientific name, can be easily avoided.)

Preparation of copulatory organs, and illustrations

The technical manipulations necessary to obtain good preparations for drawing

purpose, although often time-consuming, are relatively simple. Pinned or freshly

killed specimens can be relaxed over wet sand for six to 24 hours, old museum

specimens that were killed in cyanide bottles (a strongly dissuasive method!)

sometimes requiring up to 48 hours to soften the sclerotized parts. Thereafter the

whole male copulatory complex can be pulled out slowly as far as possible, and the

various parts separated by first clipping carefully all basal and lateral ligamentous

membranes of the minute sternal plates. After cleaning off all muscles and

connectiva still adhering to the various chitinous structures (without the use of

caustic potash), the latter are washed in diluted alcohol, transferred to glycerine,

and placed on slides protected by loosely applied cover glasses. Since most

internal abdominal sternites are usually flexible and often bent in a different way,

the shapes (contours) of the hidden plates may at times look rather different in

serial preparations of a single species collected in one locality; in any way pressure

should be avoided to prevent distortion. Component parts thus protected can be

left floating in glycerine for years, and conveniently examined at any time for

comparison with serial preparations of other specimens. The objects having been

drawn to the desired magnification, they are best demounted for permanent

safekeeping. First of all, the sclerites of each individual are washed out in 70%

alcohol with at least one change, passed to aceton for a short while to ensure

thorough cleaning, and dried on filter-paper. Then they can be kept together in

plastic microvials mounted with gum on a slip of cardboard (as for Coleoptera)

and attached to the insect’s pin. A somewhat delicate, but more convenient and

satisfactory method is to paste the objects one by one to the pointed tip and sides

of ordinary cardboard slips. For the best adhesive use Velpon (or any other glue

soluble in water) dissolved in amyl-acetate, and stir until the solution (to be kept in

a glass-topped vial) has become sufficiently syrupy. A minimal quantity of this

substance applied to any part of the slip will suffice to securely hold the separate

parts in the desired position. Care must be taken to ensure the crucial components

of each object (bristly apices of sternites, gonostyli, etc.) to project freely beyond

the card’s tip or sides, so as to simplify rapid inspection. The drawings in this paper

of male and female genital organs were all taken from preparations as described,

though not always rendered on the same scale.

All illustrations, including those showing the peculiar pubescent “pads” usually

adorning the males’ mid tibiae, are original camera lucida sketches.

LIEFTINCK: Palaearctic Melecta 145

Flower records

Only unpublished flower records are acknowledged and copied from the labels

in the lists of specimens examined, all earlier names of plants mentioned in the

literature as being frequented by the — mostly polylectic!) — species of Melecta

and their hosts, are omitted.

DESCRIPTIVE KEY TO PALAEARCTIC SPECIES OF Melecta

MALES

(Species incertae sedis are excluded but discussed in the Appendix (p. 334); males of

M. canariensis and solivaga are still unknown)

1. Gonostylus of large size, two-fifth to one-half as long as genital capsule,

incurved, hollowed out within and subparallel-sided in profile; apex obliquely

or squarely cut off, at least with part of bristly setae much longer than diameter

of stylus, these bristles characteristically curled and/or twisted, frequently

meeting or intertwining (figs. 33, 40, 49, 59). Intero-apical (mesial) angulation

of distal portion of gonocoxite drawn out and markedly protuberant in dorsal

view, directed obliquely backward and inward to form the gonocoxal angle

(g.a., fig. 6, 44). Anterior (basal) arms of sternite 7 relatively short and broad,

each expanded at about halfway length, with distinct external angulation, these

arms not nearly twice as long as the broad median plate (disk) whose apical

lobes are clothed with short setae. Sternite 8 gradually tapering, apex usually

produced, excised and also setiferous. Sides of apical extremity of tergal

(pygidial) plate 7 subparallel or slightly converging, the apex itself broad,

squarely cut off, subtruncated or, more rarely, shallowly emarginate, lacking

oblique ventral ridges; surface somewhat shiny, disk almost flat and hairless,

closely punctate. Labrum squarish but usually slightly longer than its breadth

at base or middle, with or without incomplete median ridge. Maxillary palpus

5-segmented. Distal portion of fore and mid femora clothed posteriorly with

longish erect pubescence; hind femur lacking posterior tuft, covered instead

with a compact row of fine appressed tomentum, the raised hair tips forming a

dense comb hardly projecting beyond hind border of femur; distal half or more

with partly exposed blunt or subacute median crest. Extero-apical border of

hind tibia distinctly emarginate, its lower angle produced above implantation

of spurs and furnished with a row of short, strong, spine-like setae (fig. 3), its

outer face and that of basitarsus strewn with strong, spike-like, black denticles

‘) See the excellent comprehensive review, “The ecology of solitary bees’’, published by E. Gorton

Linsley (1958, Hilgardia [Berkeley] 27: 543-599, figs. 1—3).

146

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

showing through pubescence; surface of hind tibia uneven distally, brightly

shiny in places. Hind basitarsus distinctly outbent in dorsal (posterior) view,

narrow at extreme base, then rather abruptly expanded, broadest about

halfway length, its upper border in side view straight or almost so, the lower

one convex; outer face flattened and markedly hollowed out; apex cut off

obliquely, produced dorsally to form a straight blunt process running in line

with upper border; surface finely tessellate, clothed evenly with short soft

decumbent hairs. Inner rami of mid and hind tarsal claws at most half as long as

outer and much broader than these, forming thin, axe-shaped or squarish

plates which, though often pointed, are never slender and claw-like (fig. 23).

Mid tibia somewhat expanded and swollen, its outer surface for the greater

part clothed with a compact, elongate-oval patch of short, fine and appressed,

predominantly white, silky hairs; its posterior border moreover invariably

conspicuously fringed with long, silky and (mostly) white hairs directed caudad

(fig. 7). Third submarginal cell of fore wing always longer than high, its apex

prolonged distad and more strongly and narrowly elbowed than in any other

species group, especially in male (fig. 20); yet all submarginals varying much in

size and shape. Antenna of moderate length and strength, flagellar segments

strongest in male, squarish or a little broader than long (4), or a trifle longer

than broad (9); 3 always longer than 4 but not more than 1% times the length

of this; 4 and 5 subequal. Male with well developed, distinctly impressed,

subcordate rhinaria (fig. 1). Median mesonotal line not or very shallowly

impressed; parapsidal lines short or indistinct. Suture between mesonotum and

scutellum rather deeply sulcate; scutellar tubercles short, obliquely and slightly

raised above level of scutellar disk. Propodeum coarsely rugose, rugae on

lateral edges of triangle usually placed in the long axis of body. Abdomen shiny

dorsally, punctation of tergites 1—S fine, superficial, most punctures

setiferous, the setae varying in length, semi-erect; punctures much smaller than

interspaces, finest and most widely spaced on postgradular portions.

Pubescence of head and thorax long, raised hairs concealing most of surface,

including parascutella and spines; colour variable, often more or less yellow-

brown. Antennal scape in both sexes fringed on either side with long raised

hairs, most conspicuous in male. Abdominal markings varying considerably in

size and colour, either pure white, greyish, or (more rarely) black; tergite 1

basally with longish raised tufts especially at sides, the marks on succeeding

tergites, when present, either compact, consisting of long, fine, decumbent

hairs concealing the surface, or less defined and more loosely arranged; these

spots either placed in regular row or, more frequently, constricted or twofold

on one or more segments, usually largest on 2—3, thereafter often diminishing

rapidly in size, or absent altogether. Pygidial plate of female always narrow,

long and slender, evenly and but slightly downbent; length-breadth ratio

individually variable, about twice as long as its breadth at extreme base of

lateral ridges; proximal to the latter the plate is ill-defined, extending to

(unexposed) basal margin of tergite. Surface of plate shiny, flat or almost so,

proximal portion of disk punctate, most closely so at base, for the rest minutely

LIEFTINCK: Palaearctic Melecta 147

tessellate, with rectilinear, finely raised side-margins, tapering gradually from

base to apex, the latter simply rounded; apical portion occasionally with feeble

median crest. Mid tibia in female also invariably fringed with some raised white

hairs (dark in melanistic forms). Hind basitarsus slender, much narrower and

less outbent than in male, at least four times as long as its breadth near apex, its

outer face slightly convex instead of being hollowed out, clothed densely with

longish, mostly decumbent hairs. Body pubescence in extremely melanistic

forms throughout deep black. Size very variable, length 6—20

oi rde BONE oA ONS ase M. albifrons (Forster) alliance 2

Gonostylus generally much shorter and straighter, at most one-third length of

capsule, not or less conspicuously hollowed out within, tips of styli more widely

distant, most of these parts strongly setiferous, but longest setae only very

rarely curled or twisted. Shape of gonocoxal angle variable, but distal portion

of gonocoxites in dorsal view not as described, the intero-apical (mesial)

angulation never drawn out or markedly produced obliquely caudad and

mesiad, and also less abruptly angled in side view. Anterior (basal) arms of

sternite 7 generally longer, more widely divaricate and, with few exceptions,

more slender; if broadened at about halfway length to form external

angulations, then the latter are feebly sclerotized and semitransparent; shape

of median plate very variable but generally much shorter than the limbs and

often constricted basally. Tergal (pygidial) plate 7 of male equally variable in

shape. Maxillary palpus usually 6- occasionally 5-segmented, very rarely with 4

segments only. Extero-apical border of hind tibia often oblique though

rectilinear or only shallowly concave, the lower outer angle of same not

markedly produced, but frequently spinulose and/or carrying strong setae and

bristly hairs (fig. 4). Inner rami of mid and hind tarsal claws generally longer

and more slender, more definitely claw-like. Posterior border of mid tibia

lacking conspicuous fringe of backward projecting long hairs, though often

with few longish erect bristles instead. Shape of third submarginal cell of fore

wing very variable, usually shorter and less abruptly angled distally. Abdominal

markings white, often conspicuous, those on tergites 1—S (or 6), if present,

never twofold, occasionally replaced by condensed dots of black hairs, or

HbSentaltogether Allremaining Species groups... Fee. o ieri ar. 8

Both sexes with single row of spots, one each side, on abdominal tergites 2—5

(rarely also 6) in male, and on tergites 2—4 (rarely also 5) in female. These

spots may be greatly reduced in size, either lacking on some posterior

segments, present only on tergite 2, or absent altogether, in which case

abdomen (occasionally whole body) is totally black and unspotted (some

a. nigra). Except in some female albovaria the spots are irregular in shape,

characteristically ill-defined and hollowed out anteriorly. A complete collar, or

at least a lateral tuft, varying in colour, conspicuous also on tergite 1. If spots

on tergite 3 are small, tending to become obliterated or interrupted, then all

light hairs (especially on head and thorax) are distinctly brownish yellow or

grey-brown in fresh examples, most tergal spots then being ill-limited (all

nominotypical albifrons). Sternal graduli 3—5 or 4—5 of male with

148

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

inconspicuous fringe of longish, decumbent, dark (rarely whitish) hairs.

ot Kent antiek Grenen hee NGE 1. Group of M. albifrons (Forster) 3

Both sexes invariably with two, usually pure white, spots on each side of

abdominal tergite 3 (frequently also paired on 2), and male occasionally also

with twin-spot on 4 and even 5, in which case there are two complete rows of

white lateral spots. Also a basal collar, or at least a single tuft of white (or

faintly brownish) hairs, at sides of tergite 1. Sternal graduli 3—5 or 4—5 of male

usually with distinct comb-like fringe of long, decumbent or suberect, bristly

hairs, often white and band-like laterally; these bristles in female longer than in

male, evenly and more sparsely distributed ver dt Rea

Vane ROME placa. 2. Group of M. duodecimmaculata (Rossi) 5

Head, including long fringes at antennal scape, thorax and legs predominantly

brownish yellow to grey-brown on dark brown ground, the thoracic

pubescence rather long and dense but nowhere completely obscuring the

surface. Suberect, mostly brownish black pile covering abdominal tergites

much shorter though easily discernible at low magnification (x 10); many long

raised bristles also at graduli and sides of tergites, | moreover with transverse,

ill-defined collar of long raised and fluffy brownish hairs, more crowded

laterally to form tufts of more or less curved and undulated hairs directed

sideways and partly projecting beyond hind margin; tergites 2—5 (4) or 2—4

(2) with row of whitish or pale brownish white lateral spots varying much in

size, shape and number, but nowhere sharply outlined: usually conspicuous,

irregular and tufty on 2, much smaller, placed more inward, depressed and

spot-like on next tergites, the one on 3 frequently placed transversely and

constricted, or even twofold, in which case the parts are unequal in size; spots

3—4 (5) in northern populations often vestigial or wanting in both sexes.

Venation brown: fore wing membrane never entirely hyaline, lightly infuscated

except marginal cell, centres of submarginals and papillate border, which are

slightly (though not at all contrastingly) darker brown. For structural details,

see additional descriptive notes and illustrations (figs. 3, 19—25). Body size and

markings extremely variable, length 9—17.5 mm. Hab.: Temperate NW

Europe, from Britain eastward farintoW Asia ........ a. albifrons (p. 193)

Head, thorax and legs alternately black and pure white to slightly greyish on

deep black ground, but consistence of pubescence not differing from

a. albifrons. Light hairs covering thorax rarely with distinct pale brown hue.

Venation generally darker than in nominotype but colour of membrane

variable, darkest in melanistic populations of a. nigra. Lateral spot on tergite 3,

when present, very rarely constricted or twofold. Body size and markings

equally variable: partition of white pilosity specified in next couplet. Two, not

sharply differentiated subspecies of more southern and eastern range. Varieties

intermediate between populations of typical albifrons and albovaria on the one

hand, and between albifrons and nigra on the other, are of frequent occurrence

and discussed under each subspecies, the latter being roughly distinguished as

follows „in „seek Gaus ease titan Mode alé initie RSR ee 4

Light areas on head and thorax greyish white; abdominal spots white, either

LIEFTINCK: Palaearctic Melecta 149

unequal and much reduced in size, or absent altogether. No white hairs

between scutellar tubercles. — White extremes of &: clypeus dorsally,

antennal scape partly, and whole occipital region; mesonotum as far back as

scutellum (except around tegulae); sides and venter of thorax; almost complete

external patch and posterior fringe of mid tibia; basal one-fifth of hind tibia,

and vestiges at tarsi; almost complete collar on tergite 1 and large lateral

subtriangular patch on 2; more or less transverse spots on 3—4 (5); sternites all

black. Dark extremes of ¢: white hair obscured to grey-brown on occipital

area, mesonotum anteriorly, and centre of mesepisterna; external patch of mid

tibia and spots at base of hind legs also obscured and small; body for the rest

entirely black. — White extremes of Q: head in front and occipital area (in

part); anterior collar of mesonotum and isolated lateral patch; small tuft

behind wings; mid and hind legs as in male; lateral tufts on tergite | and small

lateral spot on 2; vestiges on 3—4; or body (inclusive of tergite 1) totally black

except well defined spots on 2—4 only. Dark extremes of 9: body entirely

black. Size less variable, length 11.5—15 mm. Hab.: Discontinuously, from SW

England through western France; Italy and Tyrrhenian Is. . . . a. nigra (p. 204)

Both sexes with light areas on head and thorax pure white, on mesonotum

anteriorly occasionally faintly tinged light brown. White extremes: abdominal

spots conspicuous, placed in regular row, but often ill-limited on tergites 1—5

(3) or 1-4 (9), becoming smaller from before backward, those on

intermediate segments distinctly transverse, usually largest on 2, irregular in

shape, tapering inward, often excised anteriorly, the anterolateral angle

occasionally reaching compact tuft at sides of 1. A pair of more or less isolated

black dots slightly in advance upon middle of mesonotum rarely present in

male, more frequent in female, the latter in addition with pair of larger white

tufts to the inside and just beyond halfway level of tegulae, and small tuft of

white between scutellar tubercles. Mid tibial pad of 4 complete, often with

tiny rusty brown streak along anterior border. Dark bristly hairs fringing

sternal graduli 3—5 of ¢ frequently pure white instead of black laterally. Both

sexes of darkest populations in southern France and Spain having vestigial

tergal spots 3—4 (5), are practically inseparable from light extremes of a. nigra.

Body size very variable, length 7—19 mm. Hab.: Discontinuously (see above)

from central France and Iberia throughout the whole Mediterranean as far as

REIN BE Pe EIN OB RE. a. albovaria (p. 196)

. Size small, length usually 10—12 mm, not exceeding 13 mm. Head and thorax

clothed all over with greyish white to light brown hairs, small dark brown areas

occasionally only low down at temples, on basal portion of scutellum and,

narrowly, at sides of thorax above mid coxae. Integument dark brown, never

deep black; legs often lighter brown. Vestiture rather long and dense,

scutellum with conspicuous tuft of recurved brown hairs between short spines.

Mesonotum of female without any trace of anterior dark spots. Outer faces of

mid and hind tibiae and tarsi for the greater part clothed evenly with short,

mostly decumbent, pale brownish to white hairs, sparsely distributed at both

ends without forming definite patches or spots; hinder pair in both sexes

150

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

moreover beset with setiferous punctures of different sizes mixed with few

short, raised, pale bristles and strong flattened, suberect, black spicules on

somewhat shiny ground, the latter most numerous in female, longest and

present also upon distal portion of basitarsus. Wings infuscated except at

extreme base; fore wing, from slightly beyond cells to extreme tip, broadly

bordered smoky brown. Abdominal pubescence rather long, especially the

collar at base of 1, which in male is broad, only slightly arched behind and but

little condensed laterally, usually pale brown. All spots on next tergites

isolated, decumbent, smallish, but varying in shape and size, always paired on

2—3, often also on 4—5 (4) or 4 (Q); bristly sternal pubescence long and

dense, pregradular fringes oblique, present in both sexes, hairs suberect, most

conspicuous and longest on 3—S, colour silvery to light brown. For further

details, see text and figs. 57—59. Hab.: E China ......... chinensis (p. 218)

Size variable, length 11—18, frequently exceeding 14 mm. Head and thorax at

least with some black at sides and underneath, markings pure white, strongly

contrasting, hair length variable. Integument black. Mesonotum of female

often with pair of isolated black spots anteriorly; scutellum and mid-posterior

sclerites of thorax predominantly black, the former often with tuft of pure

white recurved hairs between short spines. Outer faces of mid and hind tibiae

and basitarsi contrastingly white-spotted: at least extreme bases and apices of

mid tibiae and distal portion of hinder pair remaining black. Pregradular

suberect bristles covering sternites 2—5 of female less numerous and all

blackout z hess waters aste Dita tele as ae 6

Body pubescence in both sexes throughout long and rather dense, hairs

covering dorsum of thorax partly concealing surface and even exceeding

length of tegulae and scutellar tubercles; all black hairs on disk of abdominal

tergites suberect, well visible with the naked eye. Basal half or more of tergite |

with uninterrupted band of long raised white hairs most closely set and tufty at

sides, but with its posterior limit almost (or perfectly) rectilinear. Marks on

succeeding tergites composed of thickset hairs raised above surface level

(male, profile!), the tips recurved or turning sideways, projecting beyond hind

border of tergites; spots depressed, less bushy and more sharply outlined in

female. All spots, though placed in regular rows, varying much in size and, on

terminal segments, also in number: on 2 and 3 invariably paired and completely

isolated. Inner spots on 2 always smaller, on 3 a little larger, than outer spots,

the interspaces variable though frequently broader than diameter of inner spot.

Scutellum completely black, rarely a few white hairs midway between

tubercles; propodeum black in middle; thoracic sides predominantly, though

never completely, white. Punctation on outer faces of mid and hind tibiae

dense, punctures of different sizes, mixed with longish bristles and interspersed

with few short, broadly triangular, spike-like setae arising from much larger

punctures, most of these spicules in Mediterranean populations hidden under

pubescence; interspaces polished though small. Wings only little obscured,

membrane of fore wing gradually and only slightly infuscated from base to tip,

darkest in marginal cell, but papillated area not contrastingly coloured or

LIEFTINCK: Palaearctic Melecta 151

band-like. Male with sternal plates 2—5 clothed all over with suberect hairs,

3—5 moreover with subapical brush of more raised hairs (profile!), longest and

partly white at sides. Apical plate of sternite 7 longer than its width at base (fig.

38); distal portion of 8 also relatively long, sides lacking distinct convexities

(figs. 38, 42). Gonocoxal angle well pronounced though bluntly rounded,

usually less incurved than in next taxa (see, however, fig. 44). Size extremely

variable. Hab.: Mediterranean region (typical), through N Asia as far as

RN IEN COOPER aah Pe DINO d. duodecimmaculata (p. 209)

Body less hirsute, the pubescence shorter, especially so in female; longish hairs

on dorsum of thorax mostly shorter than length of tegula. White hair band

covering basal half of tergite 1 forming an arched collar, narrowest at middle or

even interrupted in the median line. Marks on succeeding tergites composed of

compact hairs which are not, however, raised above surface level. Wings

darker, fore wing membrane more contrastingly coloured beyond cells. Strong

spicules covering outer face of hind tibia well visible. Apical plate of sternite 7

subequal in length to or shorter than its width at base (fig. 43); distal portion of

8 (unknown in excelsa) also comparatively short, sides usually with prominent

convexities (fig. 43). Gonocoxal angle often more prominent and markedly

OE UOC SS RENTE AUT, Karr ROUE TIR RE A 7

Size large, length 15.5—17 mm. General appearance similar to nominotype but

pile shorter, abdominal spots of much larger size and fore wing membrane with

broad (ca 2 mm) though not contrasting, grey-brown band bordering distal

margin beyond cells. Thoracic pubescence of both sexes shorter but more

variegated with snow white: well defined mesonotal spots to the inside of

tegulae, conspicuous tufts on metapleurae behind wing bases and propodeum

laterally, the scutellum moreover with triangular white spot between tubercles,

this isolated tuft consisting of curiously depressed radiating hairs directed

cephalad and laterad, female in addition with pair of black dots on mesonotum

anteriorly. Legs shaped much as in nominotype, but setiferous punctures on

outer faces of mid and hind tibiae (particularly on hinder pair) less numerous

and more widely spaced on rather irregular ground, leaving distinctly more

shiny areas. Length-breadth ratio of male hind basitarsus = 100: 30. All tergal

spots of large size, sharply defined, 2 invariably larger than in nominotype (see

descriptive notes). Apical plate of sternite 7 shorter than its breadth at base

(fig. 48). Hab.: Transcaspian region and Iraq ......... d. jakovlewi (p. 215)

General appearance as in pl. 1 fig. 4. Also of large size, length 16.5—18 mm.

Differs from previous taxa as follows. Vestiture of whole body still shorter and

more even. Alternating black and white thoracic pattern distinctly more

contrasting, the black predominating; white spots between tegulae and

scutellar tubercles in both sexes small, standing out clearly, the white

mesepisternal patch completely isolated. Fore and hind wing membrane

subhyaline, but fore wing with costal vein and whole area beyond cells

distinctly more contrastingly tinted, the apical band and tips of hinder pair

broader and darker brown. Black pile covering abdominal tergites and

scattered bristles at sides visible only at some magnification. All tergal spots

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

relatively small, more regular and sharply outlined. Collar-like band at base of

1 markedly concave behind, complete in male, subinterrupted in female, hairs

long and thin, partly raised at base, broadest laterally though less defined and

composed of longer hairs than in jakovlewi; spots twofold and completely

isolated on 2 and 3, single on 4—6 (4) or 4—5 (Q), the latter diminishing

gradually in size posteriorly; appressed hairs closely set, not exceeding hind

margin of tergites. Sternal plates 3—5 of male very closely punctate, the black

setae of minute size, the longer subapical fringes black in middle, slightly

raised, small, spot-like and white laterally; raised black bristles much shorter

than usual; female with all hairs and bristles more sparsely distributed. Legs

more robust, outer faces of mid and hind tibiae more sparsely and rugosely

punctate, the surface (especially of hinder pair) being very uneven, with

smooth and distinctly shiny areas isolating many strong raised, spike-like setae

placed on elevated rugae, especially conspicuous in female. Hind basitarsus

more markedly outbent and hollowed out externally and also more expanded

at middle, length-breadth ratio = 100 : 34.6. Genital structures of male as in

figs. 43—44, the pygidial plate of female shaped and sculptured as in other taxa

of the group, but broader, length-breadth ratio = 100 : 65. Hab.:

Afghanistan: 4 ores eed yearn dineti Cat EE Ge excelsa (p. 217)

Antennal rhinaria on flagellar segments 3—13 absent, replaced on 4—12 by

large, poorly defined, oval or subrectangular, rather flattened areas occupying

nearly the whole posterior surface of each (fig. 2), extreme bases of these areas

occasionally with a minute, subtriangular, impressed spot (rutenica only):

accordingly clearly discernible, distinctly impressed and shiny sensoria are

invariably wanting. Mid tibia unmodified, neither flattened nor broadened,

nature of pilosity covering its outer face not or hardly differing from that on

hind tibia and lacking a well defined oval or subcircular patch of short,

compact, felt-like, snow-white (rarely black) tomentum, a white hair spot,

when present, being restricted to basal portion of tibia, ill-defined distally and

consisting of decumbent hairs, the latter either short or of normal length (fig.

8). Hind basitarsus slightly but distinctly outbent in posterior view. Inner rami

of mid and hind tarsal claws slender, not broader than outer, from 2/5 to 3/4 as

long as main branch. Antennal scape fringed laterally with long, raised, mostly

white hairs. Size moderate, length not exceeding 13.5mm ............ 9

Antenna variable but often stronger, usually with shorter, more swollen

flagellar segments, the posterior faces of 3—13 or 3—12 always provided with

distinct, mostly deep, pit-like and shining rhinaria, placed slightly in advance of

the middle (fig. 277), the one on 3 elongate-oval, those on succeeding segments

subtriangular or partly linear, often becoming smaller and more rounded

distally; if antenna is thin and more slender, then these impressed sensoria are

occasionally more superficial and less defined, though invariably present (e.g.

fulgida). Mid tibia generally slightly expanded and swollen, consistence of

pilosity covering outer face markedly different from that of fore and hind tibiae

by the presence of an elongate-oval or subcircular flattened patch of short,

compact, felt-like and usually white tomentum; long backward directed hairs

LIEFTINCK: Palaearctic Melecta 153

fringing posterior margin of mid tibia inconspicuous: sparsely distributed or

NEEN ine rp lean US ice RA ALS 11

. Antennal rhinaria placed at base of flattened areas of segments 4—12

extremely minute and superficial, as indicated above, absent on 3 and 13; each

of the flattened areas bounded posteriorly by a low ridge which, though poorly

indicated, is a little convex posteriorly, giving the antennal joints a somewhat

swollen appearance. Whole antenna moderately strong, rather short and

hardly reaching tegulae; scape clothed with long erect brownish black hairs

roundabout; 3 little longer than its breadth at apex (100: 80) but slightly longer

than almost square following segments. Labrum slightly longer than its

(greatest) breadth at base, evenly closely punctate lacking median carina,

anterior border convex, little upturned. Body texture and nature of

pubescence much as in luctuosa, but pile on head, thorax and legs

predominantly black, the light hairs, inclusive of a long decumbent patch

covering clypeus, more greyish instead of pure silvery white; head for the rest

black, as is a long fringe at occipital border, save only a small thin tuft of grey

on each side of the latter; anterior mesothoracic collar, a tuft at upper half of

mesopleurae, and a still smaller spot behind wings, whitish; remaining parts all

black. Scutellar tubercles shorter than and partly concealed by long raised

hairs, triangular and shiny. Legs more closely and finely punctate, less lustrous,

than in /uctuosa, the rather long pubescence much the same, but long white

hairs restricted to a posterior fringe at articulation of fore femur-tibia, and

vestigial subbasal spots at outer faces of mid and hind tibiae. Outer face of mid

tibia beyond white spot with decumbent, dullish hairs, pile more compact and

longer than on the normally hairy and partly exposed hind tibia, thus differing

from luctuosa and brevipila. Hind basitarsus little or not hollowed out

externally; with at least some long bristles fringing posterior border. Fore wing

membrane darker than in /uctuosa. White collar of raised hairs at base of tergite

1 thin, intermingled with black medially, the still longer loose tuft on either side

replaced by black at the bend; lateral spots on 2—5 transverse, smaller and

more rapidly diminishing in size posteriorly than in next species, hairs also

shorter. Tergal (pygidial) plate 7 as in fig. 97. Sternites 7—8 as in fig. 98.

Genital capsule squarish, more parallel-sided than in luctuosa, but gonostylus

and dorsobasal process shaped similarly to quite some luctuosa. Size moderate,

length 12 mm, fore wing 10.5 mm. Hab.: S European Russia; Turkey (see map

LO OAT? So. SARAS lv BA ee rutenica (p. 233)

Flattened areas replacing impressed antennal rhinaria simple, without any

indication of minute centro-basal impressions, these areas not bounded by a

low convex ridge posteriorly. Antenna a little longer and distinctly more

slender than in rutenica. Body pubescence lighter, predominantly white on

antero-dorsal part, sides and ventral sclerites of thorax; long white tufts at least

posteriorly at fore tibiae, behind wings, and on either side of scutellar

tubercles; patches of appressed white hairs at outer faces of all tibiae and tarsi;

a band of long raised or suberect hairs, broadest laterally, also on tergite 1, and

compact lateral spots on 2—5. Middle of scutellum and sclerites behind this

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Figs. 7—12. External view of & right mid tibia, showing white pubescent pad; 7, M. albifrons albovaria

(Montpellier, France); 8, M. luctuosa (Baarn, Netherlands); 9, M. grandis (Oran, Algeria); 10, M. trans-

caspica (lectotype Turkmenia); 11, M. aegyptiaca (Egypt); 12, M. baeri (Krasnowodsk, Turcomania)

LIEFTINCK: Palaearctic Melecta 155

EN EN AEEA css, SY 10

10. Antenna normal, reaching tegula, 3 less than twice as long as its width at apex,

but distinctly longer than succeeding segments which, though always slightly

longer than broad, vary somewhat in length, the distalia occasionally almost

square. Body relatively short and broad, apical segments more rapidly

tapered than in next species; pubescence longer and more conspicuous, all

raised hairs on thorax directed straight up. Legs posteriorly and externally

with many longish erect black bristles, long tufts of white on fore and mid

femora, rather long and dense black (or white) fringes also on hinder pair.

Outer faces of tibiae and tarsi reticulate-punctate, sculpture partly visible

under pubescence; hind tibia distal to white hair-spot rather shiny, with short

spine-like setae sparsely intermixed, but fringed with long bristles. Shape and

vestiture of hind basitarsus much as in rutenica, subparallel-sided in side view

and about four times longer than broad, not or scarcely expanded about

halfway length, fringed with distinct thick marginal setae and long bristles.

Seutellar tubercles greatly variable in shape and length, frequently very short

or barely indicated though often straight and more or less pointed, directed

obliquely upward and backward; even if spines are long then nevertheless

much shorter than, and partly concealed by, surrounding pubescence.

Abdominal tergites smooth and shiny, finely superficially and not very closely

punctate, all punctures setiferous, much smaller than interspaces, smallest

and least impressed on apical portion of tergite 1, but successively more

widely spaced on next segments. White lateral spots compact, placed

transversely in regular row, composed of long, depressed, finely branched

hairs, all spots much broader than long except those on tergite 5, which may

be smaller and almost circular. Remaining dorsal pubescence black, hairs

suberect and partly branched, with longish black bristles sparsely intermixed

at sides and on last two segments. Sternites dull, punctation variable, usually

close, except rather broad hind borders of 1—4 more finely so and partly

impunctate, all clothed fairly densely with mostly decumbent black hairs, like

tergal pile clearly visible even at low magnification in side view; posterior

borders impunctate. Tergal plate 7 tapering rapidly, the apex itself narrow,

subparallel-sided and hairless; surface shiny, reticulate-punctate on each side

of a broad, shallow, impunctate median sulcus; apex straight or slightly

upturned, distinctly excised but depth of emargination variable, the side-

angles either rounded or almost pointed; usually reddish brown or even paler.

Lateral ridges of sternite 6 very low, invisible in side view. Shape of hidden

sternites 7 and 8 variable, arms of 7 rather broad and straight (figs. 82, 85, 88,

92). Genital capsule of small size, 1.2 — 1.5 mm (incl. stylus); inner margins of

gonocoxites divergent dorsally, the gonocoxal angle evenly rounded (fig. 90).

Length 11 — 13.5 mm, fore wing 9.5 — 11 mm. Hab.: Central Europe, W. Asia

EE CE eh ae NS cuits Te A digital, di luctuosa (p. 227)

— Antenna still longer and more slender, reaching back to beyond tegula; 3

almost twice as long as its width at apex, about one and one-fifth as long as next

segments, which are all markedly longer than broad. Body relatively long and

156

—

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

narrow, general appearance, pl. 2 fig. 7; abdomen elongate-oval, gradually

diminishing in breadth, end segments less abruptly pointed than in /uctuosa and

many species resembling it; pubescence generally thinner and much shorter,

raised hairs on dorsum and sides of thorax more inclined backward. Scutellar

spines strong, straight or slightly downcurved, directed caudad, more exposed

than in luctuosa though not exceeding tufts of surrounding long pubescence.

Legs dull, more closely punctate than in /uctuosa, reticulation finer but

sculpture remaining partly visible under pubescence; most parts, including

trochanters, with minute appressed hairs not quite concealing somewhat shiny

surface; light tufts present on all coxae; conspicuous, long and more compact

white and black hairs only at posterior ridges of fore and mid femora, those

covering hind femora extremely short, dense and appressed, and hairs fringing

blunt longitudinal ridge suberect, though visible only under a strong lens. Pile

covering mid tibiae and tarsi short and depressed, concealing most of surface;

outer face of hind tibia distally with few short erect spine-like denticles. Hind

basitarsus visibly hollowed out externally, distinctly expanded in side view

about halfway length, hence less than four times longer than broad, devoid of

any long bristles at posterior border, only apical margin fringed with longish

thick bristles. Abdominal tergites dullish, finely superficially and closely

punctate all over, basal portions with smooth and rather shiny surface, but

postgradular areas finely transversely wrinkled with more closely set

punctures, those on apical tergites larger and still more crowded; hind margins

narrowly impunctate. White lateral spots relatively small, varying in size and

shape, placed in regular row, smallest in holotype (pl. 2 fig. 7); hairs

decumbent, much shorter than in /uctuosa. Remaining dorsal pubescence

black; setae very short, suberect, partly branched; no longish black bristles,

except few suberect hairs at sides of tergites and near apex on dorsum of 6.

Sternites dull, evenly very closely punctate, all punctures setiferous and almost

contiguous on distal sternites, the setae minute and in profile visible only at

some magnification on two basal sternites; posterior margins not or very

narrowly impunctate. Tergal plate 7 tapering gradually toward end; surface

dull, flat or very slightly concave lacking median sulcus, clothed densely with

elongate setiferous punctures, all setae depressed; lateral rims of apical portion

not sharply pronounced, tip subtruncated, more or less translucent, the

emargination crescent-shaped with rounded angles (holotype and one

paratype, fig. 99), or only very shallow (one paratype); oblique submarginal

ridges on ventral face of tergite distinct. Lateral tubercles of sternite 6 distinct

though low and visible in side view. Sternites 7 and 8, fig. 100. Genital capsule

small, 1.7 mm (type, incl. stylus); inner margins of coxites evenly curved

dorsally, at first diverging, then parallel and finally slightly converging, the

gonocoxal angle a little prominent and incurved though evenly rounded;

gonostylus, fig. 101. Length 14 — 17 mm, fore wing 10.5 — 11 mm. Hab.:

Turkestan st: as Ghee, Teese ME SU brevipila (p. 235)

. Mid tibial and outer hind tibial spurs reduced in size, both thick and spine-like,

the former in profile less than one-third as long as apical breadth of tibia, the

LIEFTINCK: Palaearctic Melecta 157

latter shorter than half length of inner spur (which is slender and normal) and

hardly half as long as apical breadth of hind tibia in side view. Hind femur

posteriorly with dense fringe of erect black hairs much shorter than diameter

of femur (see description). Outer face of hind tibia strewn with strong suberect

black denticles showing through the pubescence. Vertex without glabrous area

just outside each of the lateral ocelli and no impunctate areas on propodeum.

Labrum squarish. Antenna strong, reaching to a little beyond tegula, flagellar

segments 4—13 thick, scarcely longer than broad; rhinaria well developed,

deeply impressed. Non-pubescent parts of abdominal tergites shiny, very finely

superficially punctate. Body, inclusive of outer faces of legs, extensively white,

head and thorax almost entirely so; this pubescence long and dense, forming

large-sized depressed lateral marks on abdomen; sternites 2—5 on either side

with transverse fringe of longish white submarginal hairs; 6 with pair of distinct

low tubercles. Tergal plate 7 rather broad and flat, apex shallowly emarginate

(fig. 164). Gonocoxal enclosure V-shaped, i.e. margins nearly straight and

divergent, but angles (though rounded off) rather suddenly curving inward;

gonostylus parallel-sided, apex almost squarely cut off, dorsobasal process

setiferous, figs. 165—166. Size moderate, length 12 mm, fore wing 9.5 mm,

wing expanse 22.5 mm. Hab.: Uzbekistan and Turkmenia ... . nivosa (p. 257)

Mid tibial and outer hind tibial spurs of normal size and length, i.e. the former

subequal to apical breadth of tibia as seen in profile, the latter more than half

as long as inner spur and at least % apical breadth of hind tibia in side view;

spurs generally somewhat undulated, feebly angulated or S-shaped and/or

slightly twisted. Other characters combined not as above ............ 12

. Labrum long and narrow, in frontal view almost twice as long as its breadth at

base (100 : 51—53) almost parallel-sided (fig. 150). Face markedly porrect,

clypeus squarish in dorsal view. Antenna slender, all flagellar segments longer

than broad, 3 and 4 subequal; rhinaria distinctly impressed. Median mesonotal

line slightly but distinctly sulcate. Upper surface of clypeus entirely concealed

from view by a thick layer of very long and dense, appressed silvery hairs; sides

at least partly white. Long and dense white posterior fringes at fore and mid

femora, replaced on hinder pair by a comb of very short black hairs fringing

median ridge posteriorly. Wings almost hyaline, all veins yellowish brown,

marginal cell of fore wing and a cloudy streak along anterior border distal to it,

ferruginous. Pubescence on head, thorax, legs and abdominal tergite 1

extensively white, hairs long, raised and (on mesonotum) dense,

characteristically dowdy. White postero-lateral spots 2—5 large, placed

transversely, most conspicuous on 3, all much broader than their interspaces,

except on 2 abbreviated and rather narrowed inward; remaining pubescence

black, all hairs somewhat raised, intermixed with longish white and black

bristles at sides and on apical segments. Terminalia as in figs. 151—153. Small,

sexually heterochromatic species: Q almost wholly black! Length 9 — 11.5

mm, fore wing 7.5 — 8.5 mm. Hab.: Egypt ......... angustilabris (p. 251)

Labrum in frontal view not nearly twice as long as its greatest breadth at base,

usually only little longer than broad, sometimes squarish or even slightly

158

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

broader than long. White tergal spots on 3—4, when present, not considerably

broader than distance separating them and if so, then spots are not three or

more times broader than high (see, however, baeri and prophanta). Often larger,

not‘sexually heterochromatic, (species ja) Asia) OS NE ee) ee 13

. Body black (both sexes!), black-haired without any light markings, except a

narrow, dirty brownish area centred on outer face of flattened mid tibial pad,

and lateral tufts behind wings, the latter exceptionally dark: fore wing for the

greater part dark brown with low purplish and blue reflections; veins almost

black. M + Cu in hind wing coincident with cu-v, or nearly so. Vestiture of

body and legs generally much shorter than usual, the raised hairs covering

thorax, though dense anteriorly and at sides, hardly longer than diameter of

tegula, and not much exceeding length of scutellar tubercles in perfectly fresh

examples (holotype); tufts of longer hair present only on antennocular, genal

and occipital areas, laterally behind wings, and on lower parts of thoracic sides.

Raised hairs on post-scutellar sclerites sparsely distributed; suberect bristles on

mid and hind legs and abdomen remarkably short. Large species, length 17

mm, fore wing 13.5 mm. For further details, see description, figs. 146—148 &

pli fig: Hab ABeypE Arendt ere RITA: fumipennis (p. 248)

Body at least with some white pubescence anteriorly on mesonotum, tufts on

either side of scutellum behind wings, at outer faces of mid and hind tibiae,

and/or laterally on one or more tergites of abdomen. Fore wing membrane

often strongly infuscated but not strikingly dark brown. If body is wholly black,

then fore wings are not exceptionally dark, legs and abdomen are partly

clothed with longish hairs, and the size is smaller: length not exceeding 15

DT A OE Se rn En A EE Se 14

. Antennal segment 3 almost or fully twice as long as 4. Integument at outer face

of distal portions of mid and hind tibiae brilliantly shining, surface uneven but

strikingly polished, with few large, setiferous punctures and strong, triangularly

flattened denticles fitting into somewhat raised sockets. Basitarsi more closely

punctate but, like tibiae, punctures are strewn on more shiny ground than

usual. Preapical tooth on pollex of mandible undeveloped, replaced by a low

convexity (fig. 74). Antenna slender with distinct though feebly impressed

rhinaria, which fade away distally and are absent on last three segments. Legs

also thin and slender; compact patch of white covering outer face of mid tibia

less conspicuous and sharply defined than in most other species; apex of hind

tibia (fig. 63); inner rami of mid and hind tarsal claws broader and more

flattened than usual (fig. 64). Fore wing membrane much obscured, mottled

grey-brown with ill-defined subhyaline areas; part of fore wing venation, fig.

65; nervellus antefurcal. Abdomen rather glossy, punctation variable. Body

pubescence generally long and fine, though rather fluffy and not very dense.

Hidden apical sternites of characteristic form, especially sternite 7 (fig. 67);

gonocoxal angle rounded (fig. 68). Pygidial plate of 9 narrow and slender (fig.

70), resembling in shape that seen in the albifrons group. Length variable,

8.5—14 mm. For further details, see specific description. Hab.: From S France

through the northern Mediterranean as far east as Iran ..... fulgida (p. 219)

LIEFTINCK: Palaearctic Melecta 159

— Antennal segment 3 less than 1% times as long as 4 or even shorter. Integuinent

at outer face of distal portion of mid and hind tibiae lacking strikingly polished

and impunctate areas; if partly smooth and shiny, then surface is either finely

tessellate or fairly closely punctate with or without interspersion of strong,

tooth-like setae or spicules; apex of hind tibia not strongly produced

externally. Plate of sternite 7 not as described and figured for fulgida, apex

usually emarginate or bilobate and strongly setiferous .............. 15

. No raised black or white hairs of any length in the centre upon the bend at base

of abdominal tergite 1. The latter relatively long, i.e., up to 4/5 length of 2 in

dorsal view (average ratio 79 : 100), its subhorizontal part sloping rather

abruptly into the declivous anterior face, but angulation rounded. All tergites

noticeably dull (except occasionally slightly shiny on disk of basal and upon

middle along hind margin of apical segments), integument exceptionally

closely and finely punctate, all punctures in fresh examples bearing minute,

decumbent or suberect dark setae, which are finely branched or plumose, most

of the surface thus producing the effect of being bare and almost lustreless.

Integument of mesonotum posteriorly, disk and sides of scutellum, as well as

median area of postscutellar sclerites, sparsely black-haired or naked, the

scutellar tubercles remaining well visible. White pubescence covering anterior

portion of mesonotum shorter than usual, hairs scarcely longer than diameter

of tegulae. Longer white hairs present on clypeus, behind antennae, at

occipital border, on posterior angles of mesonotum in front of parascutella,

behind wing bases, and on lateral thoracic sclerites; tufts of long white hairs

often also around and/or below scutellar spines. Legs not at all hirsute: clothed

with short hairs, hence lacking long bristles, except thin tufts at coxae and the

usual white posterior fringes at fore and mid femora. Hind femur evenly closely

punctate, laterally clothed with minute decumbent pubescence in addition to a

posterior brush or comb of extremely short, closely set hairs. Black denticles at

outer face of hind tibia not very strong or conspicuous, though frequently just

visible amid the white pubescence. Outer faces of all tibiae and tarsi with

appressed, predominantly white, tomentum; dense felt-like pad on mid tibia

distinct, complete and well defined. Hind basitarsus invariably a little outbent.

White abdominal spots conspicuous, sharply outlined, composed of matted

hairs, as shown in the photographs (pl. 3 figs. 15—18). Labrum longer than

broad or squarish. Antenna relatively slender, scape with dense lateral fringes

of raised hairs that are longer than diameter of scape; rhinaria present,

distinctly impressed. Paraocular region invariably with smooth impunctate

area adjoining outer side of lateral ocellus. At least distal portion of fore wing

membrane obscured. Tergal plate 7 broad, subtruncate, hind border variously

emarginate; surface uneven, disk somewhat hollowed out between low rims.

Paired ventral ridges or small tubercles present at apex of sternite 6, forming

low wart-like projections, not prominently visible in profile. Gonocoxal

enclosure U-shaped dorsally, the distal angle obtuse-angulate, either broadly

rounded or slightly more suddenly deflecting, the angle itself always rounded

onasizerrathenlarge length 418mm ton PO UCR MENU 16

160

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

At least a few long raised white or black hairs present in the centre upon the

bend at base of tergite 1. The latter shorter, viz., at most slightly over 3/4

median length of 4 (ratio 66.8 : 100 or less), the surface generally less convex

and sloping gradually into declivous anterior face of tergite, the transition

accordingly more gradual. All tergites invariably somewhat glossy; integument

often closely finely punctate but if so, then most microsetae are simple or less

finely branched, frequently somewhat raised and not concealing the surface;

this vestiture usually interspersed with longer setae or bristles, especially

abundant at sides and gradular lines as well as upon disks of apical segments.

Integument of posterior thoracic sclerites often partly exposed but most parts

clothed with raised hairs exceeding diameter of tegulae in length, those

surrounding scutellar spines usually concealing them from view. Tomentum

covering posterior two pairs of tibiae and basitarsi invariably also interspersed

with some longish:hairs'or bristles;Size-variable Ka Ya) SON 18

. General appearance and body pattern, pl. 3 fig. 15. Mesonotum slightly

biconvex, integument with median and parapsidal lines markedly sulcate, the

impressed median line almost reaching scutellum, the parapsidal grooves

subequal in length to transverse diameter of tegula. Disk of mesonotum and

scutellum coarsely rugosely punctate, most punctures deep, confluent with

irregular and shiny rugae; anteriorly and on either side of parapsidae punctures

become smaller, circular and contiguous. No median scutellar line; tubercles

short, robust, placed transversely, shape somewhat variable: either straight,

forming an isosceles triangle, or more slender, compressed and spike-like, their

surface punctate. White tufts below scutellar tubercles at least three times as

long as these. Propodeal triangle dull, irregularly punctate, with vestigial

impunctate area at its base; side angles filled out with 3—4 obliquely placed

ribs. Raised pubescence covering head and thorax relatively long, the

somewhat depressed hairs on mesonotum anteriorly concealing most of the

surface; white at temples and low down on thoracic sides becoming black,

hairs on posterior part of mesonotum and scutellum predominantly black.

Parascutella hidden from view by a compact brush of deep black hairs. Distal

two-fifth of outer faces of hind tibia and bases of mid and hind basitarsi black.

Fore wing membrane beyond cells infuscated. White lateral spots at base of

abdominal tergite | relatively small, placed far apart, subtriangular in outline,

broadest basally, mainly composed of short appressed hairs but laterally at

extreme base distinctly longer, tufty and suberect; spot on 2 broadly oval or

subcircular, those on 3—5 more transverse and narrower, though broader than

interspaces, becoming smaller posteriorly, apparently wanting on 6. Length

14.5—18.5 mm, fore wing 13—13.5 mm, wing expanse 30 mm (pl. 3 fig. 15). For

further characters, see descriptions and figs. 168—172. Hab.: Uzbekistan to

Pakistan: Turkey-(?)Sedsraely ie ia tin 6008981 corpulenta (p. 260)

Combined characters not as above, but sculpture on dorsum of thoracic

sclerites much as in corpulenta. Median mesonotal and parapsidal lines less

distinctly impressed. Raised pubescence covering head and thorax shorter,

white predominating, hair on mesonotum posteriorly and sides of thorax all

LIEFTINCK: Palaearctic Melecta 161

white. Outer faces of mid and hind tibiae as well as the tarsi mostly white. All

lateral abdominal spots large and sharply outlined, those at base of tergite |

subrectangular, occupying most of the dorsal surface but abruptly narrowed

inward to form pointed basal prolongations nearly meeting in the median line;

spots on succeeding tergites also large and conspicuous though varying in

EEA asl ll BO DENE tee. Bt eR SES DIET EFT, 17

. Labrum square. Stature slender, with elongate abdomen and slim legs; all

femora and tibiae rather thin, the outer faces of mid and hind tibiae not

noticeably convex. Fore wing membrane, subhyaline basal and anal cells

excepted, abruptly and contrastingly dark smoky brown. Impunctate area on

either side of lateral ocelli somewhat impressed, polished; a crescentic area

along anterior border of median ocellus also impunctate though not impressed,

small circular spot upon middle at some distance behind median ocellus. A

small, somewhat shiny and finely wrinkled median impunctate area also at base

of propodeal triangle, the latter closely punctate except that lateral angles are

traversed by 3—4 irregular rugae placed in the long axis of body, less marked

than in corpulenta. Clypeus above and paraclypeal area with silvery patch of

mostly decumbent hairs not reaching anterior border medially; raised white

pubescence covering rest of head, thoracic sides and underneath, short and

rather sparse, hairs on mesonotum directed caudad, not exceeding length of

tegulae and nowhere quite concealing surface. Facies and body pattern as in

pl. 3 fig. 16. Length 15 mm approx., fore wing 10 mm, wing expanse 24.6 mm

(holotype). Further details, see description and figs. 173—178. Hab.: Turkey

MEERN pi) Egypt, Israeliand'Libyai 10 Poe 344 hl as honesta (p. 264)

Labrum a little longer than broad. Stature more robust, with a larger head,

broader abdomen and more swollen legs. Outer faces of mid and hind tibiae

distinctly convex and more densely pubescent. Fore wing paler, membrane

tinged much as in corpulenta, the distal half gradually becoming light brown.

Impunctate areas surrounding ocelli as in honesta, except that there is no

smooth spot behind median ocellus. Propodeal triangle closely punctate

lacking definite median impunctate area, lateral angles distinctly rimmed but

enclosed space devoid of rugae. Raised white pubescence covering head and

thorax longer and more compact, hairs upon mesonotum raised but concealing

most of surface, more abundant and much longer also at sides and underneath.

Length 14 mm approx., fore wing 12 mm, wing expanse 28 mm. For further

details, see description, pl. 3 fig. 17 & figs. 179—184. Hab.: Turkestan (terr.

SARE PENNEN TE NEE AI PL LEDEN a amanda (p. 266)

. White tergal spots on 2—4 distinctly transverse, much broader than deep,

shallowly emarginate anteriorly, boundary not sharply delimited and

contrasting at all sides. Clypeus above clothed with long, little diverging,

suberect white hairs, as also the midfrontal and parorbital areas; clypeal hair

not definitely spot-like and matted centrally but more fluffy and of unequal

length, not concealing all of the surface anteriorly; sides of clypeus and malar

area with long raised black hairs. Thorax clothed with long, not very dense,

white pubescence, but lower parts of pleurae and ventral sclerites black. Tergal

162

TIIDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

spots 1—5 conspicuous, hairs on 1 very long and partly raised, on 2—$

transverse, depressed and band-like. Sternites partly with thin fringes of

longish decumbent silvery hairs. Dense pad of short felt-like white pubescence

covering outer faces of mid tibia incomplete and ill-defined: hairs thinning

gradually toward base so as to leave integument partly exposed. Hind femur

posteriorly clothed evenly with dense brush of erect brown hairs becoming

increasingly longer toward base. Few long raised black bristles bordering hind

tibia and basitarsus posteriorly. Inner rami of mid and hind tarsal claws only

half as long as outer. Tergal plate 7 slender, sharply defined, perfectly straight

in profile, bounded by a pair of subacute rectilinear ridges (fig. 155). Basal

arms of sternite 7 unusually broad and flat (fig. 156). Genital capsule slender,

enclosure U-shaped, gonocoxal angle evenly and broadly rounded, the

diverging borders following an almost straight course toward lower edge of

gonostylus, the latter short and broad, hollowed out within (fig. 158). Small

species, length 10.5—11 mm, fore wing 8.7 mm. Hab.: Balearics, Romania, and

from the Caucasus south into Turkey, Lebanon and Israel . guichardi (p. 254)

White tergal spots on 2—4 varying in shape and size but nearly always sharply

outlined anteriorly. Clypeus — if not entirely black above — with conspicuous

silky patch (varying in size) of depressed, long and dense, mostly silvery white

pubescence concealing the surface; these hairs, implanted from extreme base

to beyond halfway length of clypeus, are straight and usually arranged fan-

wise, either lying flat upon surface with tips pointing forward, or directed

slightly upward, whether or not surpassing anterior border; long hairs covering

sides of clypeus, malar space and lower portion of parocular area usually black

and distinctly raised. Vestiture of body variable, but hairs at sides of thorax

generally all white. Dense pad of short, felt-like pubescence covering outer

face of mid tibia varying in size but invariably more sharply outlined, though

occasionally obscured. Inner rami of mid and hind tarsal claws nearly always

more than half as long as outer. Basal arms of sternite 7 longer and more

slender; if short and rather broad, then outer borders of these limbs are more

sinuous. Tergal plate 7 and other characters combined not asabove ..... 19

. Gonocoxal angles in dorsal view prominent: the inflected inner margins of

gonocoxites, which together form the boundary of the enclosure, at first

divergent, rather V-shaped and almost rectilinear (festiva fig. 6), or else, more

outbent and U-shaped, curved markedly inward round the bend, but in any

case forming distinctly pronounced, occasionally subacute, gonocoxal angles

(which themselves are rounded), the margins finally diverging again abruptly to

reach the dorsobasal process of gonostyli in a straight or undulated line (fig.

344). If gonocoxal angles are only slightly incurved, viz. in alcestis and alecto,

then hind femur is devoid of longish hairs. Antennal scape invariably clothed

on either side with long raised hairs. Hind basitarsus in dorsal view always

more or less outbent, the outer face at least slightly hollowed out (concave) in

side view. Body pubescence variable, but black pile on abdomen usually well

visible with a pocketlens (x 10); 2—4(5) conspicuous, compact, usually placed

transversely though paired white tergal spots varying in size and shape, made

Liertinck: Palaearctic Melecta 163

up of long hairs, which on disk of 1 are loose and raised, becoming gradually

more crowded together and spot-like at sides, the tips of these tufts reaching or

eeceeding posterior-Dorder of tergite | 4.1.12 pi 20

Gonocoxal angles in dorsal view not incurved, usually simply rounded:

curvature of inner margins of gonocoxites embracing central cavity variable,

the borders diverging more markedly before the bend of the open enclosure,

but angles are at best subrectangular or, more frequently, broadly rounded,

deflecting gradually in a broad curve toward dorsobasal process of gonostyli,

either directly (as in fig. 134) or following an undulated course to become “‘bi-

Bate as eg inileucorhyncha, fig, 329 roa eee oo oe BS 24

‚Hind femur closely, rather strongly punctate and short-haired, the vestiture

quite inconspicuous and visible only with a powerful lens: short and decumbent

at posterior face and along ventral longitudinal ridge, on which punctures are

contiguous, beset with a brush, or comb, of extremely minute hairs hardly

projecting beyond hind border in side view. Hind tibia and basitarsus entirely

devoid of long, widely spaced, erect marginal bristles posteriorly, but on outer

face of tibia the soft white and black hairs are intermixed with many strong,

tooth-like black spicules. Base of hind basitarsus, just outside ventral ridge,

with fairly well pronounced, finely punctate, flattened and shiny area,

occupying from one-sixth to almost one-fourth of total length. Antenna much

as in next species, intermediate segments slightly but distinctly longer than

broad in frontal view. Fore wing membrane smoky grey-brown especially

toward apex and in cell centres (wings in old worn males apparently clear)

ee no pda AO a ah + ores. pS oe 21

Hind femur similarly punctate but posteriorly devoid of short dense tomentum,

clothed instead at all sides with much longer raised hairs which project caudad

well beyond ventral (posterior) ridge, most hairs being at least one-fourth as

long as diameter of femur, frequently much longer, especially toward apex. In

lateral view the upper gonocoxal margins, just after the angle, follow an

(almost) straight course toward lower distal edge of gonocoxite (fig. 228) .. 22

. Gonocoxal enclosure widely V-shaped in dorsal view, but borders at first gently

concave, then curving a little inward round the bend, from which point, after a

short and slight concavity, they form a second obtuse angle (hence are “‘bi-

angulated’), until finally diverging again, pursuing a long, shallowly incurved

course (profile!) above the dorsobasal process of gonostylus, before reaching

base of the latter (fig. 194). Impunctate area on either side close to lateral ocelli

not impressed but very shiny and equalling ocelli in size. Legs more robust;

hind basitarsus relatively long (100 : 27.7—30.1), nearly parallel-sided in side

view. No row of conspicuous strong spines, just above shortest (external) spur,

at apical border of hind tibia (fig. 190). Tergal plate 7 rather broad, sides

converging with swollen border, tip subtruncated, shallowly emarginate, not

upturned; disk flat, striato-punctate and hairy (fig. 191). Long raised black

hairs at sides of clypeus and anterior porton of parocular area. Conspicuous

tufts of long white hairs on posterolateral sclerites of thorax and roundabout

scutellar tubercles, only narrowly separated by black on middle of scutellum.

164

22)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Abdominal tergites rather shiny, all punctures larger and more deeply

impressed than in next species, the black hairs longer and transverse white

spots on 1—5 composed of longer hairs, while spots on 2—3 are oblique and

not sharply delimited anteriorly. Hind margins of sternites only narrowly

impunctate; gradular lines of 2(3)—5 conspicuously decorated with compact

white spots on each side. Long soft pubescence covering sides of thorax and

sternal faces all white. Body pattern, pl. 2 fig. 11. Size moderate, length 13—14,

fore wing 10—10.5 mm, wing expanse 23.5—24.4 mm. For further details, see

description and illustrations. Hab.: South European Russia and Israel

Ya Re RAL DER GRA ye alcestis (p. 270)

Gonocoxal enclosure U-shaped in dorsal view, inner margins at first gently

concave, then almost straight though curving a little inward round the bend,

from which point the margins diverge again, following an almost straight

course toward ventral edge of gonostylus. Impunctate areas close to ocelli

dullish, not pitted, unapparent or vestigial. Legs distinctly more slender than in

alcestis, but hind basitarsus shorter (100 : 32—34.3) and more expanded in side

view, its greatest breadth slightly beyond halfway length (fig. 195). Apical

border of hind tibia, just above shortest (external) spur, with row of 5—6 robust

black spines, which are longer than the strong tooth-like black spicules

interspersed between the soft white and black pubescence covering its outer

face. Tergal plate 7 a little broader, more finely and less distinctly rimmed than

in alcestis; disk flat, hairless and distinctly more shiny, finely superficially

striato-punctate, tips a little upturned (fig. 196). Only sides of clypeus black,

parocular area white. Tufts of long white hairs covering posterolateral sclerites

of thorax and roundabout scutellar tubercles shorter and less conspicuous, the

latter small and broadly separated by black on middle of scutellum. Abdominal

tergites more glossy, finely superficially punctate, the black hairs shorter than

in alcestis and unapparent at low magnification; white spots on tergites 2—5

compact, composed of shorter hairs, those on 2—3 oblique anteriorly though

sharply delimited at all sides. Hind margins of sternites broadly impunctate; no

white fringes at gradular lines. Tubercles near apex of sternite 6 small, but

more prominent than usual, just visible in lateral aspect (fig. 197). Long soft

pubescence covering sides of thorax white upon hypoepimeral area and upper

portion of mesepisternum, lower parts mixed white and black, or wholly black.

Size moderate, length 14 mm approx., fore wing 10—10.5 mm. For further

details, see description and illustrations. Hab.: From Turkey and Israel to Iraq

and E’Furkestane ver Redon eue bas ngi eee non alecto (p. 273)

Large species, length 16—21 mm, fore wing 12.5—14 mm (pl. 5 figs. 29—30).

Tergal plate 7 of abdomen relatively narrow, sides strongly converging, very

hairy and lacking a blunt carina, apex bifid, ending in a pair of closely

approximated rounded tips, which are hairless, light coloured and separated by

a deep crescentic V- or U-shaped emargination; surface slightly concave,

punctate and densely hairy on each side of a complete elongate, somewhat

shiny, naked and variably sculptured area (fig. 225, ventral side!). Tubercles at

sternite 6 elongate, varying in size but always small, rather low and hairy, not

23.

LIEFTINCK: Palaearctic Melecta 165

or only just visible in profile. Dorsobasal process of gonostylus with both angles

exposed and rounded off in side view (fig. 229). Propodeum coarsely punctate,

but triangle invariably with distinct dullish median impunctate area varying in

shape, usually with short impressed median line. Posterior faces of mid and

hind femora throughout evenly closely punctate lacking impunctate spaces

near apex. Inner face of hind basitarsus sulcate at full length, black hairs

covering sides of depression suberect and longer than decumbent pile covering

sulcus; no well defined smooth basal concavity of any length at ventral face just

inside lower border, the latter evenly convex in oblique side view. Inner rami of

mid and hind tarsal claws at least 4/5 length of outer. Body pubescence

generally shorter, legs more slender and less hairy, than in next species, but

mid and hind tibiae and basitarsi with admixture of longish raised marginal

bristles; outer face of hind tibia throughout closely punctate, the tips of short,

strong, black spicules clearly visible amid the soft hair. Fore wing membrane

distinctly burnt brown beyond cells and, especially, toward apex of marginal

cell and further. White tergal spots widely separated, placed far laterad, the

hairs shorter, more compactly set, and spots on 2—3 better defined anteriorly,

than in next species. Hair covering sternites all black. Antenna strong,

intermediate segments slightly but distinctly longer than broad, more elongate

than in next species. Silky white patch covering clypeus lying flush with surface

and restricted to its dorsum, the hair tips just reaching anterior border. Hab.:

SL SEG Se Mer COCO 00), 8 ER grandis (p. 286)

Size smaller, length not exceeding 17 mm, usually less. Tergal plate 7 of

abdomen variable, but usually broader, sides less strongly converging, bluntly

carinate and hairy, apex straight cut off or shallowly (occasionally more

deeply) emarginate, tips more apart and broadly rounded; surface flat or

slightly hollowed out, disk sparsely, more broadly, striato-punctate, hairless

and rather shiny (figs. 241, 250, 257, 259, 261 & 268, dorsal and ventral sides).

Only distal angle of dorsobasal process of gonostylus exposed in side view (figs.

261, 273). Propodeum as in grandis, but sculpture of median area of triangle

very variable in both species. Abdominal sternites with hind margins not or

very narrowly impunctate; gradular lines often fringed with mixture of

decumbent, bristle-like, white and black hairs, or with more condensed white

lateral patches. Remaining characters combined notasabove ........ 23

Sternite 6 ventrally with pair of robust, elongate, subapical tubercles on each

side of a suboval, less sclerotized area, these bosses either partly naked or hairy

(fresh examples), but always plainly visible in lateral aspect (fig. 267). Ventral

(inferior) face of hind basitarsus, just inside lower border near base, with

distinct elongate-oval concavity, this sulcate area occupying from 1/4 to 1/3 of

basitarsal length (occasionally hidden from view by overlying inner tibial spur),

its surface slightly shiny, covered with minute setiferous punctures, the hairs

minute and depressed; lower border in oblique lateral view shallowly but

noticeably concave for about same distance as length of sulcate area (this

concavity unapparent if basitarsus is looked at in straight lateral position!).

Disk of tergal plate 7 slightly concave, clothed with short appressed brownish

166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

hairs. Gonocoxal angle less abruptly hooked inward than in festiva, apex of

gonostylus bluntly rounded. Outer face of hind tibia closely reticulate-

punctate, basal 1/5 to 4/5 clothed densely with appressed, silvery white, finely

branched hairs concealing surface, rest of tibia black; outer faces of hind tibia

and basitarsus devoid of short, erect, spine-like black setae between soft, white

Figs. 13—18. External view of 3 right mid tibia, showing white pubescent pad; 13, M. leucorhyncha leu-

corhyncha (‘‘ebusana’’) (Mallorca, Baleares); 14, M. leucorhyncha leucorhyncha (Rodos 1.); 15, M. italica

(Italia mer., Zeller); 16, M. festiva (Vesperterminen, Switzerland); 17, M. candida (holotype Quetta, Pa-

kistan); 18, M. prophanta (holotype, Lanzarote, Canary Is.)

24.

LIEFTINCK: Palaearctic Melecta 167

and black pubescence (profile!). Raised white and black bristles at lower

border of hind tibia and outer ridge of hind basitarsus much shorter and less

numerous than in next species. Inner rami of mid and hind tarsal claws more

than half as long as outer (fig. 266). Propodeal triangle usually with small,

dullish, median impunctate area. Silvery patch covering clypeus made up of

decumbent hairs, as described for grandis. For further characters, see

description, pl. 6 fig. 35 & pl. 7 figs. 36—37, and illustrations. Hab.: see map

EEE CROP RES ers Ee cer LEES tuberculata (p. 300)

Tubercles at ventral side of sternite 6 not conspicuously prominent, much as

described for grandis, occasionally just visible in side view. No well marked

elongate-oval concavity at base of ventral (inferior) face of hind basitarsus, a

sulcate area, when present, unapparent, very short, the lower border of same

rectilinear in oblique lateral aspect, or very shallowly concave. Disk of tergal

plate 7 coarsely striatopunctate on slightly shiny ground, almost hairless (figs.

241, 261). Gonocoxal angle sharply pronounced, subacute (figs. 245—246,

264); apex of gonostylus usually truncated, its dorsobasal process

subtriangular, mostly fringed with long, often finely branched bristly setae

(figs. 246—247). Outer face of hind tibia (and/or basitarsus) somewhat more

coarsely and less densely reticulate-punctate than in tuberculata, white

pubescence more extensive, covering at least basal half; this external

pubescence moreover with admixture of short, suberect, spine-like black setae

whose tips are quite conspicuous (profile!). Broadly spaced white and black

bristles fringing inner (lower) border of hind tibia and outer ridge of hind

basitarsus, raised, more numerous and of greater length than in tuberculata (fig.

249). Inner rami of mid and hind tarsal claws generally shorter (fig. 237).

Propodeal triangle usually without dullish median impunctate area. Silvery

patch covering clypeus made up of long, slightly raised hairs directed straight

forward, leaving clypeal surface partly exposed in front but surpassing a little

the anterior border. For further characters, see description, pl. 6 figs. 31—32,

asehiliestrationspHabxiseemap 2 am. ae ee mann: festiva (p. 293)

Hind femur clothed at all sides with short hair, much as described above for

alcestis and alecto (couplet 21), i.e. visible only at some magnification; sides

occasionally more sparsely beset with longish raised hairs which do not,

however, exceed one-third breadth of femur; posteriorly, the pilosity is dense,

composed of very short and fine hairs of equal length, sometimes extremely

closely set, appressed and directed sideways (aegyptiaca), apparent only with a

strong lens; midventral ridge usually with brushy comb of equally short hairs

not or hardly projecting beyond hind border in side view. If hairs at sides are

somewhat longer (NW African aegyptiaca), then tergite 7 is elongate, quite flat

and pubescent above, its apex truncated. Black pubescence generally short,

especially on abdomen and most of legs, with longish raised black bristles only

sparingly distributed at margins of hind tibia and basitarsus, and at gradular

lines of sternites. Gonocoxal angle evenly rounded: borders of enclosure distal

to the bend not undulated (bi-angulate). Antenna rather thick and strong,

reaching back as far as halfway or full length of tegula, length-breadth ratios of

168

25.

26.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

intermediate segments variable (figs, 128,231 & 277) 552.410 a 3 25

Hind femur more hirsute; midventral ridge often with comb of short, closely

set raised hairs, as before, but pile at sides and along hind border of femur

much longer, invariably plainly visible with a hand lens (x 10), projecting

caudad well beyond the crest, becoming longer toward base, often quite

conspicuously so and exceeding greatest breadth of femur. Black pubescence

generally longer at all body parts and legs, hairs on abdominal tergites usually

well visible(same magnification): Lr ts Waan sd dn CRS 29

Long raised pubescence covering sides and ventral sclerites of thorax at least

partly black, often predominantly so, subcircular patch of white restricted to

upper portion of mesepisternum, usually also a tuft of black just in front of hind

wing base. Tufts of white below wings and on each side of scutellum

surrounding the tubercles forming transverse bands only narrowly interrupted

by black at middle. Wings strongly darkened, smoky brownish black, fore wing

darkest brown at forked main veins, at centre of submarginals and in marginal

cell, membrane beyond cells with slight purplish reflex. Tergal plate 7 very

broad, covering whole exposed surface of segment, sides converging,

somewhat raised and swollen toward apex, which is blunt, very slightly

emarginate with rounded angles; disk concave, surface striato-punctate and

hairy only at sides, central part with low, broad, finely tessellate median ridge,

the latter smooth and sulcate, often again narrowly divided lengthwise (fig.

210); ventral tubercles oblique, placed near inner margin (fig. 207). Swollen

ridge of sternite 6 hairy, very low and unapparent. Apex of sternite 8 deeply

triangularly excised (figs. 203, 208, 211—213). Labrum square, broadest at

middle, disk almost flat. Maxillary palpi 6-segmented, but last joint usually

vestigial or absent. Silvery patch covering clypeus lying flush upon surface,

most hairs directed straight forward with tips not reaching anterior border.

White mid tibial pad broad and sharply defined, with black anterior streak,

composed of very compactly set short hairs lacking admixture of short black

spicules (fig. 10). White tergal spots large, compact, sharply outlined, except

condensed lateral tufts of white forming arched collar at base of tergite 1; spots

on 2 more or less curved and crescentic, tapering inward, their anterior limit

nearly always distinctly concave (pl. 4 fig. 22). Sternites black. Large species,

length 15.5—19.5 mm, fore wing 12.5—13.5 mm, expanse 30 mm (pl. 4 fig. 22).

Habs scedescnriptionyast Ans 1e aide alt Br: SR transcaspica (p. 275)

Long raised pubescence covering sides and ventral sclerites of thorax all white,

except black on metepisternum and posterior parts. Wings much lighter, bases

clear or subhyaline, fore wing gradually growing darker, grey to grey-brown,

toward apex, light brown (or more contrasting) streaks or spots only at end of

radial and marginal cells, and centres of submarginals. Tergal plate 7 more

rapidly tapered, disk never bisulcate medially. Apex of sternite 8 not so deeply

excised. Maxillary palpus 6-segmented (unknown in candida). Abdominal

tergites less closely, more finely punctate and shiny ............... 26

Tergal plate 7 slender, tapering gradually toward end, sides rectilinear, densely

brown-haired mixed with longish depressed bristles, apex truncated or

LIEFTINCK: Palaearctic Melecta 169

shallowly emarginate, sides and rounded angles swollen, often distinctly

upturned, yellow-brown; disk flat or slightly concave, surface dull, reticulate-

punctate but in fresh examples covered with golden brown, decumbent hairs

completely hiding the surface, only the tip remaining bare (fig. 132), lower

border of tergite below plate with fringe of rather long dark brown bristles.

Body more elongate than in baeri and transcaspica. White tergal spots 2—5

relatively small, placed wide apart in regular row, diminishing gradually in size,

spot on 2 often somewhat ill-limited and concave anteriorly, the one on 5

nearly always present, though small, subrectangular (pl. 4 figs. 19—21). Legs

slender. Posterior (inner) faces of hind trochanter and femur, in addition to

short black pile, clothed all along midventral ridge with regular row of

exceedingly dense and fine decumbent hairs concealing the surface; these

matted hairs are of even length, directed obliquely caudad and apicad, the tips

not or scarcely projecting beyond hind border of femur; colour dark brown in

certain lights. Hind basitarsus hardly expanded, subparallel-sided in side view,

length-breadth ratio 100 : 30—30.6, scarcely to moderately outbent in dorsal

aspect. White mid tibial pad rather narrow, some black spicules interspersed

and visibly projecting at least toward apex of tibia (fig. 11). Silvery patch

covering clypeus fan-like, hairs slightly raised, tips exceeding anterior border;

long raised hair at sides of clypeus upon paraocular area black up to level of

antennal base. Body pubescence generally longer than in baeri and candida.

Structural features, figs. 127—145. Size moderate, length 13—16.5, fore wing

11—12 mm. Hab.: see map 1 and also Canary Island Lanzarote (sub lindbergi

Mietiesy nano.) (pli figs. 1921) sens remontant sers aegyptiaca (p. 244)

— Tergal plate 7 not as above, more distinctly sulcate: disk narrower, median area

—I

at least partly exposed, shiny and hairless, the apex pale, shallowly emarginate

between slightly upturned tips (figs. 186 & 233). Tergal spots 2—5 conspicuous,

much larger and made up of more closely set hairs, than in aegyptiaca. Hind

basitarsus more expanded at about halfway length ................ 21,

. General appearance, pl. 3 fig. 18. Stature more slender and size smaller than in

next two species, length 14 mm, fore wing 10 mm. Median mesonotal line

distinctly impressed, obliterated, irregular and punctate, punctures on

middorsum on either side of line strong and deep, in some places larger than

somewhat shiny interspaces. Distance separating ocelli about equal to their

own diameter; a conspicuous, slightly impressed shiny impunctate area outside

and just behind each lateral ocellus. White collar at base of tergite |

subinterrupted medially, forming well defined lateral spots composed of

condensed, mostly appressed fine hairs, spots on 2 almost squarely cut off

inward. Short hairs covering sternites mainly black, but 2—5 each with pair of

small, well defined, submarginal white spots. Raised hairs on paraocular area

shorter than in next two species, black upon antero-lateral part of clypeus but

soon becoming white upward along eye margin. Short vestiture of hind femur

much as described for aegyptiaca, but matted stripe bordering midventral ridge

less conspicuous, darker, and composed of still shorter hair, the ridge itself

better defined toward apex. Mid and hind tibiae and tarsi almost wholly white

170 PU DSCHRIET VOOR ENTOMOLOGIE, DEEL 123, APL, 6, 1980

externally (fig, 185), mid tibial pad (fig. 17) convex, composed of very short and

dense, silky and shiny tomentum, the curved apical process partly hidden and

disk almost lacking black spicules shining through the pubescence; a dark

streak present along anterior border of pad. Lower margin of hind basitarsus

more evenly convex than in baeri, but length-breadth ratio similar, Fore wing

membrane more evenly and deeply brown on outer half than in baeri and

prophanta, Punctation of abdomen rather dense, punctures much smaller than

interspaces, Tergal pattern as shown in pl. 3 fig. 18. Median area of tergal plate

7 at first convex with few elongate punctures, then flat and almost impunctate

(fig. 186), Terminalia, figs. 186—187. Capsule smaller than in baeri and

prophanta, 1.6 mm long; dorsobasal process of gonostylus twofold, but distal

branch much narrower in side view (fig. 188). Wing expanse 24 mm. Hab.:

Baluchistan "9 IRRE EEE USER candida (p. 268)

Stature more robust and size larger: length 16—18 mm, fore wing 11.5—13 mm.

White collar at base of tergite | complete and broad but concave behind, hairs

condensed and spot-like, longest and raised at sides. Shiny area outside and

just behind each lateral ocellus unapparent and small. Short pile covering

sternites all black, Mid tibial pad distinctly more flattened, composed of

somewhat longer and less closely set hairs, Hind basitarsus a little narrower

basally than in candida, length-breadth ratio 100 : 33—36. Body pubescence

generally longer, the white tufts behind mesonotum conspicuous, only

narrowly interrupted by black upon middle of scutellum. White tergal marks

shaped differentiy tu OU vtt Ot vene RS CAA ES 28

28, White tergal spots 3 and 4 exceptionally transverse, broader than the distance

separating them, those on 2 and 5 subequal in breadth to the interspaces.

Mesonotum not sulcate medially, dorsum evenly convex but median line

distinct, somewhat raised, its course irregular, Labrum devoid of an

incomplete median carina, Apical segment of maxillary palpus longer and

more slender than usual (fig. 232). Ocelli closely approximated, separated by a

space less than one-half their own diameter. Legs more slender than in baeri;

short vestiture of hind femur as described for candida but all hairs raised and

black, lacking matted stripe bordering midventral ridge, the latter subacute

only at distal one-fourth or less. Subbasal one-fifth of fore tibia and basal half

of hinder pair white externally, rest black, All basitarsi entirely black, the

distalia partly white; hind basitarsus less markedly outbent than in baeri,

covered all over with setiferous punctures, Mid tibial pad narrow, abbreviated

basally, dark anterior streak poorly indicated (fig. 18). Wing bases hyaline,

veins very dark; fore wing with ends of distal cells, centres of submarginals as

well as whole marginal cell and papillated border, dark grey, more

contrastingly variegated than in aegyptiaca, baeri and candida; distal side of

third submarginal cell slightly more angulated than in aegyptiaca. For more

details, see description and illustrations, figs. 231—236. Hab.: Canary Is.

(Lanzarote) I Toe UAE NEER CRON FORT prophanta (p. 290)

White tergal spots placed in regular row, less transverse and also slightly

deeper than in prophanta, those on 2 and 3 subrectangular, squarely cut off

29,

Liertinck: Palaearctic Melecta 171

inward and on 2 distinctly less broad than the distance separating them.

General appearance and pattern, pl. 2 fig. 12. Median mesonotal line

indistinctly sulcate, but integument slightly depressed midposteriorly. Labrum

somewhat longer than broad with subacute median carina, broadest at apex,

occupying distal one-fourth; anterior border a little upturned (fig. 216). Apical

joint of maxillary palpus shorter than preceding segments. Ocelli farther apart,

distance separating them subequal to their own diameter. Legs more robust,

tibiae and basitarsi more expanded, mostly white, including distitarsi; short pile

covering hind femur as described for candida, with distinct matted stripe

bordering midventral ridge, the latter blunt. Nearly whole outer face of fore

tibia and basal three-fourths or more of hinder pair white; hind basitarsus

distinctly more outbent and a little broader than in prophanta, punctation

similar but entirely hidden under appressed white hairs. Mid tibial pad

occupying almost whole outer face, with few bristles forming thin black streak

along anterior border; few black spicules only along posterior border (fig. 12).

Distal half of fore wing membrane only little lighter and more evenly tinged

with brown, than in candida, not dark grey and much less contrasting, than in

prophanta. Disk of tergal plate 7 finely transversely wrinkled with few

punctures at extreme base; tips hairless, light brown, directed straight back,

gently rounded (fig. 220). Terminalia, figs. 220—221; sternite 7 with deep, V-

shaped incision. Capsule of large size, 2 mm long, dorsobasal process of

gonostylus twofold, the distal branch largest, abruptly outbent, oval in side

View, Babe Kazakhstan; Turkmenistan une, een baeri (p. 282)

Median mesonotal line impressed, markedly sulcate middorsally, sulcus closely

punctate and partly replacing the line, concealed anteriorly under long and

fine, grey-white raised pubescence. General appearance, pl.8 fig. 40, facies very

similar to italica (pl. 8 figs. 38—39). Antenna strong, rhinaria as in italica (fig.

277), length of 3 slightly exceeding 4, all segments 4—13 a trifle longer than

broad (fig. 295). Clypeus with broad patch of long, suberect silvery hairs

occupying whole dorsal surface, the tips exceeding anterior border; hairs at

sides long, raised, black. Scutellum black, a few white hairs only at side of

tubercles. Thoracic pleurae all white. Hind femur clothed densely all over with

setiferous punctures, but hairs noticeably shorter than in italica. Hind

basitarsus even more expanded and outcurved than in italica, broadest slightly

beyond halfway length. Fore wing membrane somewhat darker beyond cells.

Body pubescence rather long, but all hairs shorter than in italica. Abdomen

distinctly less shiny, more closely punctate, all punctures larger and deeper,

than in italica. Base of tergite with thin collar of long raised white hairs, similar

to that species; spots on 2—4 more defined, those on 3 a little broader and

more transverse than on 2, slightly indented by black posteriorly, 5—6

unmarked. Fore more characters and structural details, see description and

BEE aD, Egypt Dea DPR, assimilis (p. 318)

Median mesonotal line not impressed, impunctate, usually well visible though

Dien concealed trom view by long piibescence “ue se IN 30

30. Species of eastern range (Transcaspian). Wings almost or entirely hyaline.

172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Sides of clypeus as well as paraclypeal area clothed with long raised black

hairs. Head and thorax (including sides of latter) predominantly white. White

tufts behind wings and surrounding scutellar tubercles conspicuous, coalescent

on each side of a narrow black-haired area on middle of scutellum, much

longer than tubercles. Legs white-spotted and abdomen also with fairly regular

row of white spots on each side of tergites I—5, or those on 1 dispersed,

forming a thinner collar of much longer raised white hairs. Pile fringing hind

femur posteriorly also erect, black, shorter than diameter of same, no

subapical impunctate area at lower face along posterior ridge ......... 31

— Species of more western distribution (Atlanto-Mediterranean to Iran). Wing

membrane frequently much darkened beyond cells or wholly obscured, more

rarely almost or entirely hyaline. Body marks very variable. Pubescence

occasionally entirely black tonni. has Ruine sorti SNN 32

. White pubescence long and decidedly fluffy, concealing much of the surface

on mesonotum and sides of thorax, the short raised hairs covering abdominal

segments black, well visible with a hand lens (profile !). Punctures on vertex as

well as on dorsal and posterior sclerites of thorax separated by narrow

lustreless ridges; no apparent impunctate areas in front of median and beside

lateral ocelli. Body compact, thick-set, thorax and abdomen broad, the latter

more shiny than in sibirica, cordiform, rapidly tapered, length-breadth ratio

100 : 83 approx. (pl. 2 fig. 8). Antenna thick and strong with deeply impressed

recessed rhinaria, but flagellar segments a trifle longer than broad. Legs more

expanded, hind basitarsus relatively broad, lower border distinctly convex,

length-breadth ratio about 100 : 36. Tergal spots comparatively large, well

defined, composed of rather long hairs, placed in regular row, spots on tergite 5

a little broader than deep. Disk of tergite 7 shallowly concave, rather shiny,

clothed sparsely with short hairs, surface striato-punctate, except at extreme

tip (fig. 118). Body length variable, from 12 (holotype) — 16 mm (pl. 2 figs.

8—9). Hab.: Turkestan (Uzbekistan) and “Alai” ...... turkestanica (p. 239)

— White pubescence markedly shorter, the white hairs on mesonotum and

thoracic pleurae less dense, more even and mostly of equal length, nowhere

entirely hiding the surface, short erect black hairs covering abdomen (except

on tergite 1) less easily perceptible. Punctures on vertex and mesonotum-

scutellum separated by narrow, somewhat shiny ridges, small areas just in front

of median and beside lateral ocelli remaining impunctate, or beset with few

scattered punctures. Body elongate and of more slender form, abdomen

distinctly less shiny and closely punctate, gradually narrowed toward apex,

length-breadth ratio 100 : 70 approx. Antenna and legs thinner, rhinaria less

deeply impressed, but flagellar segments of antenna scarcely longer than

broad. Hind basitarsus more nearly parallel-sided, length-breadth ratio about

100 : 28. Tergal spots of much smaller size, composed of shorter hairs, those on

tergite 5 very small, subcircular. Distal side of third submarginal cell less

abruptly angled than in turkestanica. Disk of tergite 7 more distinctly hollowed

out and more shiny, at most with few scattered punctures, surface minutely

transversely striated. Structural details, figs. 103—111. Size small, length

—

32.

33.

LIEFTINCK: Palaearctic Melecta 173

12—13 mm. Hab.: Tadzhikistan, E Kazakhstan .......... sibirica (p. 236)

Scutellar spines distinctly downcurved, strong and cylindrical, only little

shorter than midlength of scutellum, at least three times as long as their width

at base and usually a little longer than the distance separating them; spines

parallel, well visible in profile but shorter than surrounding pubescence, at first

directed obliquely upward and backward, then downbent and gently tapering

to a blunt point; laterobasal edges of scutellum usually a little swollen or even

tubercular (fig. 303). Pile roundabout hind femur of moderate length, hairs in

side view shorter, of equal length and more closely set, rather comb-like along

posterior ridge. Antenna thick and strong, flagellar segments scarcely longer

than broad, 3—4 as in fig. 302; scape clothed densely all around with raised

hairs, longest at sides and white on internal, black on external faces. Patch of

silvery hair covering clypeus flat, fan-like, though all hairs are directed

obliquely upward with tips distinctly surpassing anterior border. White tufts

below scutellar spines conspicuous, almost continuous with white behind wing

bases, but separated across middle by a much narrower black-haired space

between spines. Fore wing membrane distinctly infuscated especially beyond

cells and (usually) in cell-centres. White pubescence on thorax at sides and

underneath divided by a zone of black hairs slightly below midway down.

Outer faces of hind tibia and basitarsus punctate, interspaces rather shiny,

tibial pilosity mixed with black spicules; dorsal margin of both fringed with

longish bristles. No hairless suboval groove or pit near base on lower face of

hind basitarsus, the latter slightly outbent and concave externally, little

expanded about middle; white external pad covering mid tibia elongate, not

reaching apex of segment. Thorax strongly convex above, abdomen broad and

compact. Raised white hairs at base of tergite 1 long, collar-like, lateral spots

of 1—5 large, made up of rather long hairs, 1—3(4) transverse but far apart,

subquadrangular in shape, thick and compact. Sternites invariably black.

Tergal plate 7 variable but median area usually smooth and shiny with

scattered punctures on each side of an (occasional) low median ridge; apex

shallowly (rarely more distinctly) emarginate, the tips a little upturned (fig. 304,

ventral aspect). Gonocoxal angle broadly rounded or feebly bi-angulate, the

dorsobasal process of gonostylus either as in fig. 308, or distinctly longer and

more tapered. Sternites 7 and 8 as in figs. 306—307. Size small, length 9—13.5,

fore wing 8—10.5 mm. Hab.: Canary Is.: Tenerife (terr. typ.), Gran Canaria &

EAM ee STS crie dada lentes dude thse curvispina (p. 320)

Scutellar tubercles varying in shape and size, usually shorter and only very

rarely somewhat downcurved. Remaining characters combined not as

ENE irene 2208 wet LE bl SSA Re 33

Body compact (most characteristic “facies”, pl. 8 fig. 38), rather like grandis

and assimilis, but all parts more hairy. Thorax high, abdomen short, broadly

cordate, more flattened than usual, widest at end of segment 2, then rapidly

tapered, length-breadth ratio 100 : 90 approx. Antenna not very long but

unusually thick and strong, median segments square or even slightly broader

than long, with deeply impressed rhinaria (fig. 277). Scutellum mostly black:

174

34.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

white tufts when present usually small and placed just beside or behind

tubercles. Hind femur punctate also along posterior (inner) faces, lacking

smooth and shiny area extending alongside posterior ridge and hairy all over,

especially posterior fringes long and raised: the vestiture nearest base

frequently longer than diameter of femur. Mid tibia expanded, felt-like pad

well defined, broad and flat, the hairs very dense, usually pure white (fig. 15).

Third submarginal cell relatively long, distal side not sharply angulated, but

cell rarely longer than high. Very long, erect, mostly white hairs all along base

of tergite 1; spots on 2—4(6) also consisting of longish hairs, their shape and

size irregular and very variable, becoming either smaller or larger caudad, but

when all spots are present usually largest on 2 and often not sharply outlined,

frequently vestigial or absent on posterior tergites in melanistic populations.

Sternites unspotted. Tergal plate 7 narrow, sides converging, disk nearly always

hollowed out, broadly sulcate and smooth medially (occasionally dull and

short-haired!), apex deeply emarginate between closely approximated slightly

upturned tips (figs. 278, 288). Gonocoxal angle evenly, broadly rounded, very

rarely feebly bi-angulate. Size very variable, 10.5—17 mm, fore wing 9.5—13

mm. See further notes in description, and also /eucorhyncha and assimilis (p.

318). Hab.: From Portugal in the west throughout the whole Mediterranean

excluding mostof Turkey (See map Jeet ee 8 SRO italica (p. 309)

Characters combined not as above: antenna less thick and swollen, body

usually more slender and less hairy, end of abdomen less abruptly tapered;

vestiture generally shorter, especially on dorsum of thorax and at hind

fermôrarsr AEN a MENEER ARTS DR ee 34

Closely similar in general appearance, size and antennal structure to

turkestanica from Alai (pl. 2 fig. 9); length 14— 16 mm, fore wing 11—12.5 mm.

Differences are: white body pubescence, especially on summit of head,

mesonotum and abdominal tergite 1 shorter, denser and more even, white and

black parts more contrasting. Whole dorsal surface of head closely punctate,

but all punctures larger, not quite contiguous in front of median ocellus, a

small area just beside each lateral ocellus moreover impunctate, the

interspaces, where present, finely tessellate and slightly shiny. Punctures on

mesonotum correspondingly less crowded than in turkestanica. Wings darker,

fore wing membrane distinctly infuscated beyond cells; third submarginal

slightly shorter and less strongly angled. Sculpture and shape of legs much the

same, but hind femur more densely pubescent at all sides, raised hairs fringing

posterior border markedly longer, some almost equaling breadth of femur in

side view. Upper border of hind basitarsus (length-breadth ratio 100: 34) nearly

rectilinear and beset with long black bristles much as in italica (fig. 123), no

distinct ventrobasal concavity. Abdomen somewhat more glossy, tergal

punctures more widely spaced and superficial; also raised black hairs more

finely branched, spots 2—5 a trifle larger and more transverse, than in

turkestanica. Tergal plate 7 distinctly broader, more trapezoidal, than in that

species, surface quite flat with more pronounced and somewhat convex lateral

ridges, apex subtruncate or shallowly emarginate, edges almost rectangular,

35.

36.

LIEFTINCK: Palaearctic Melecta 175

tips little or not upturned (fig. 124); sculpture as in turkestanica. Sternal plates 7

and 8 variable, though generally as in last species, limbs of 7 distinctly more

slender and curved. Gonostylus and its dorsobasal process of characteristic

shape (fig. 125). Hab.: Cyprus (terr. typ.), Turkey, Israel, Iran .megaera (p. 243)

Size smaller, body length (abdominal segments in natural, non-extended,

position) never exceeding 13 mm. Shape of tergite 7 very variable but more

narrowly tapered, apex usually distinctly emarginate. White tergal spots, when

present, generally smaller, those covering sides of 2—3 frequently somewhat

broader than deep but more often subcircular. Whole propodeal area very

closely, coarsely punctate. All abdominal sternites black; graduli beset with

longish bristles at sides of tergites and along full breadth of sternal plates.

mimemeharactersicombined:notvasibefore: :is. Ale slau!) gels Kreis ln: 35

Apex of abdominal tergite 7 strongly and abruptly tapered, disk almost flat,

shiny, finely tessellate with few deep, scattered, striato-punctures on each side

of narrow median area; lateral rims conspicuously swollen, ending in a pair of

blunt tubercles, the hind margin deeply emarginate, U-shaped (figs. 316, 321).

Antenna comparatively long and slender, relative lengths of 3—4 as in fig. 313

(holotype), succeeding segments slightly but distinctly longer than broad;

rhinaria on 3—12, oval, moderately deeply impressed. Median mesonotal line

extremely fine, incomplete posteriorly. Labrum square, or a little broader than

long, not produced medially, distal portion without well defined median ridge.

Fan-like silvery patch covering clypeus consisting of long, decumbent, silky

hairs projecting markedly beyond anterior border. Body pubescence

predominantly black; head, thoracic sclerites and legs variably tufted with

white, but abdomen (except some white on either side of tergite 1) nearly

always totally black. Hind tibia and basitarsus normally proportioned, much as

in leucorhyncha (figs. 330, 347). Wing membrane strongly infuscated. Structural

details (figs. 313—319, 321). Hab.: Lanzarote (terr. typ.) and Fuerteventura

NEER ere eee dovr dé au she Sols no ARAM caroli (323)

Apex of abdominal tergite 7 broader and less tapered, lateral rims not

markedly raised or swollen, tip not so deeply excised, the emargination shallow

(rarely more deeply scalloped: leucorhyncha from Libya, fig. 327), more nearly

crescent-shaped; disk finely transversely wrinkled, usually distinctly striato-

punctate on each side of broader median area. Antenna less slender, length of

segments variable, but 3—12 at most a trifle longer than broad, squarish, or

even a little broader than long. Pubescence on head and thorax much as in

caroli, but wing membrane invariably lighter, and white lateral spots on tergites

2—4 only rarely wanting (gracilipes from Menorca) ............... 36

Legs very slender, all femora, tibiae and basitarsi noticeably more elongated

than in allied species (see description and cf. figs. 330 & 341); hind basitarsus

more outcurved and also more distinctly hollowed out externally, than in all

leucorhyncha populations (i.e., incl. “ebusana’ and subspec. taormina); all

black, inclusive of very long fringes at posterior ridges of fore and mid femora,

which are twice as long as greatest diameter of same, those on hinder pair

much shorter. Antenna stronger than in caroli, 4—12 hardly longer than broad,

176

317;

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

almost square. Body pubescence long and tufted, hair upon clypeus semierect,

pointing straight forward, white only upon middle, the rest black; long lateral

fringes at antennal scape black anteriorly, those at occipital ridge mixed black

and white. Abdomen throughout black, inclusive of nearly all thin raised hairs

upon basal portion of tergite 1. For further details, see description and figs.

341343 )Hab:Mienorea( Balearics.) Marene gracilipes (p. 332)

Legs not unusually slender, hind tibia and basitarsus broader (figs. 330, 347).

Antenna still somewhat stronger than in gracilipes, usually square but

occasionally a trifle broader than long (/. taormina) ................ 37

White-haired areas on whole body extensive, /uctuosa-like: fan-shaped silvery

patch upon clypeus broad, covering whole upper surface, composed of long

hairs lying almost flush with clypeal surface, tips projecting beyond anterior

border; frons, lower part of temples, long fringes on either side of antennal

scape and along occipital ridge; whole mesonotum and most of thoracic sides;

conspicuous tufts behind wing bases, those beside (and partly surrounding)

scutellar tubercles spot-like, the processes varying much in shape and length

but always shorter than white hairs concealing them. Legs black: inner face of

hind femur punctate on shiny ground, punctures more widely spaced toward

end; hair short, black. Outer faces of all limbs white in places: long (partly

black) fringe at base of fore femur and tibia, and spots covering all tarsal

distalia, the basitarsus remaining black; long posterior fringe at mid femur and

large, elongate, felty pad at tibia of same (figs. 13—14), and all tarsal distalia;

hind tibia with oval patch covering its basal one-third, an oblique subtriangular

area on lower one-third to four-fifth of basitarsus as well as all distalia, likewise

white. Wings subhyaline, membrane of fore wing gradually changing to pale

brown toward apex; third submarginal cell varying in length, outer side

moderately angled. At least tergites I—4 of abdomen with broadly spaced

white tergal spots, often 5 also with subcircular dots or points, all placed in

regular row; spots on I—2 made up of much longer hairs than on succeeding

tergites, at base of 1 forming a distinct broad collar of tufted, semierect hairs,

condensed at sides and equalling length of whole tergite. Distance separating

spots on 2—4(5) averaging little less than four times their transverse diameter,

the spot on 2 usually largest. Colour and structural details (figs. 13—14,

322—332 & 344—357). Hab.: Discontinuously, on Balearic Is. (Mallorca & ?

Ibiza), the S and E Mediterranean (except Italy, Tyrrhenian Is., and Malta);

parts of Greece, the Ukraine and all over Asia minor (see map 3). For

comments on eastern forms, see description and illustrations ...........

bolero Ltd ala ot AE Pr leucorhyncha leucorhyncha (p. 325)

Structural characters and — remarkably — also wing colour and venation, as

before, but vestiture generally much darker than nominate subspecies: silvery

patch upon middle of clypeus narrower, though conspicuous, in sharp contrast

with otherwise deep black pubescence covering rest of head; hairs fringing

antennal scape and occipital ridge also black (occasionally a little white

interspersed). White pile on mesonotum anteriorly and thoracic pleurae

stopping short at level of hind margin of tegulae, white tufts behind wing bases

LIEFTINCK: Palaearctic Melecta 7a

and around scutellar tubercles almost all replaced by black; posterior fringes

and spots at outer faces of legs likewise much less extensive, mixed black and

white, or wholly black; felty pad at mid tibia narrower, completely surrounded

by black. Abdominal tergites black, collar-like band of white at base of 1 thin,

almost linear, interrupted by black medially, the longish hairs at sides white-

tipped only; remaining tergites all black, or 2—3 with mere vestiges of white

only at sides of 2(3). Colour design and structural features (figs. 333— 340 & pl.

8 fig. 41). Hab.: Italy, Tyrrhenian Is. (terr.typ. Sicily), and Malta (¢ Sardinia

and Malta unknown!). For remarks on intermediate forms, see description

EO ran eenander tiende a Eh las leucorhyncha taormina (p. 330)

FEMALES

(Characters of the two groups of the M. albifrons alliance are incorporated in the

key to the males; females of the new species M. alcestis, candida, gracilipes and pro-

phanta are still unknown)

Labrum long and narrow, as in male, only little less than twice as long as its

breadth at base (fig. 150). Body structure generally as described for male, but

vestiture wholly different: Body entirely black, very exceptionally partly light-

haired (see description). Antenna more slender than in male, all flagellar

segments relatively longer. Wing membrane considerably darker, only bases

subhyaline, for the rest dark smoky brown, especially in and beyond marginal

cell, cell centres and whole papillate distal area of fore wing; length and depth

of third submarginal cell equal. Pygidial plate (fig. 154) slender, not downbent,

broad at base, sides of proximal portion converging, the distal half narrow,

subparallel-sided, with distinctly convex median ridge. Size small, length

9—11.5 mm, only one specimen (from Wadi Riched) being much larger, 14.5

NAE OVO. 1: Tesini co res dits ak ae angustilabris (p. 251)

Labrum in frontal view not nearly twice as long as its greatest breadth at base

or about halfway length, usually only little longer than broad, sometimes

squarish or even slightly broader than long. Pubescent body markings not

slkinelwvedafesentansboth SEXES! 2 2.40 Ftsthbatnbedigags dk eu verk een: 2

Large species, length 16.5—19.5 mm, fore wing 13—14.5 mm. Similar to male

(pl. 3 fig. 13). Body black, entirely clothed with short, sooty black pubescence,

lacking any light coloured markings, except occasional greyish hairs on either

side of mesonotum in front of tegulae. Fore wing membrane still darker brown

than in male, with slight purple and bluish reflections; lighter coloured areas as

in male. Body texture also similar, no obvious impunctate areas on vertex and

propodeum, except smallish spot in front of median ocellus; median mesonotal

line feebly sulcate posteriorly. Vestiture of whole body much shorter than

usual, the raised pubescence covering mesonotum and scutellum hardly longer

than tegula. Legs slender, sparsely clothed with longish hairs; hind femur

throughout closely, finely punctate, except narrow, somewhat shiny area to the

outside near apex of blunt median ridge. Spine-like setae covering outer face of

178

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

mid and hind tibiae short, not conspicuous, as little as the suberect bristles,

though the latter are longer than in male; inner rami of mid and hind tarsal

claws about half as long as outer. Antenna much slenderer than in male; all

flagellar segments distinctly longer than broad. Pygidial plate (fig. 149) brown,

evenly and slightly downbent, the distal one-third obscured; surface extremely

finely, transversely reticulated, with or without few small punctures. Hab.:

Eeypt HE lk En RTE NE 2008 fumipennis (p. 248)

Combined charaetersinot'as'aboveni) i. omen eet A 3

. Antennal segment 3 fully twice as long as its width at apex and up to twice as

long as 4, the latter subequal to next segments, which are a trifle longer than

broad. Wings relatively longer and broader than usual, membrane strongly

infuscated, third submarginal cell usually longer than high, its distal side

markedly projecting and strongly angled. Pubescence covering head and

thorax rather fluffy, as in male. Scutellar sclerites invariably all black, lacking

long white tufts around short tubercles; white tufts behind wings isolated.

Pygidial plate (fig. 70) slender, long and narrow, up to twice as long as its

breadth at base, not or scarcely downbent; surface flat and shiny. Legs slender,

outer faces of mid and hind tibiae and basitarsi not very hairy, beset with short

thick spicules on uneven, smooth and very shiny ground, especially notable on

distal half of hind tibia, the latter closely tessellate-punctate and white-spotted

only on basal half. Inner rami of mid and hind tarsal claws about half as long as

outer, though broader than usual, resembling those of male and recalling

albifrons in this respect (fig. 64). Labrum squarish, clypeus broader than long,

both clothed with longish raised hairs. Postgradular areas of tergites very

broadly impunctate or almost devoid of fine punctures. Median mesonotal line

not impressed. Stature, body size and shape of white abdominal marks

extremely variable, tergal spots frequently vestigial on one or more segments

though never completely wanting. Hab.: From southern France through the

northern Mediterranean as far east as Iran, see map 1 ...... fulgida (p. 219)

Combined characters not as above. Antennal segment 3 always much less than

twice as long as 4. Distal half of hind tibia not noticeably polished and shiny

externally, but if so, then only toward apex I... N MER +

. Stature slender, size small, length 11—12 mm. Resembles fulgida in some

respects: pygidial plate shaped similarly, dark brown tipped with black, plate

nearly twice as long as its breadth at base, apex with or without feebly

pronounced median ridge (fig. 163). White hair on clypeus above and frontal

area likewise long, suberect and fluffy, only partly concealing surface, not

exceeding anterior border of clypeus. Very similar to male, differing from

fulgida as follows. Abdomen narrower, more rapidly tapered and pointed. Body

with rich white markings. Hairs fringing both sides of antennal scape long and

dense; conspicuous white tufts behind wings fused together with those

covering sides of propodeum and also nearly meeting well defined tufts just

behind each of the strong spike-like or markedly downcurved scutellar spines,

which are shorter than the spot-like tufts (the latter absent in fulgida). Lateral

marks on tergites 1—4 transverse, except on 1 forming subcircular patches

LIEFTINCK: Palaearctic Melecta 179

narrowly connected across dorsum with thin collar of equally long raised hairs;

marks on 2—4 band-like, as in male, characteristically concave and not sharply

defined anteriorly; distance separating spots on 3—4 much shorter middorsally

than their own breadth; short dark hairs covering tergites suberect. Body

punctation closer than in fu/gida: interspaces of punctures on mesonotum less

than one puncture width, less shiny. Raised hairs covering thoracic dorsum

anteriorly more sparsely intermixed with black. Legs slender, dark brown; hind

femur glossy, throughout closely punctate, except narrow, almost impunctate

stripe bordering median ridge distally, hairs scanty, suberect. Outer faces of

mid and hind tibiae and tarsi almost wholly white, but sometimes becoming

black toward apex of tibiae without forming well marked boundary line; hairs

short, decumbent, tibiae with conspicuous admixture of robust dark spines;

inner rami of mid and hind tarsal claws little less than half as long as outer and

scarcely broader than these, never definitely axe-shaped. Postgradular areas of

all tergites 2—5 distinctly but very finely punctate. Appressed sternal

pubescence and raised bristles on 2—S silvery in certain lights, as in male.

Balearics, Romania, and from the Caucasus south into Turkey, Lebanon and

IRE RETE DADI ZIE RUE, >; guichardi (p. 254)

Siemaicversicombined not as described 7777.11 ER EEE I BE! 5

Whole body clothed with moderately long, snow-white pubescence; admixture

of dark hairs, black bristles and spike-like setae only in places: see below and

original description. Labrum longer than in male, widest slightly before

halfway length, then somewhat narrowing toward end (length-breadth ratios 40

: 30 : 26), the apical border shallowly emarginate with rounded side-angles;

surface at first convex, then little concave, apex slightly upturned; no distinct

median ridge; disk coarsely rugosely punctate on shiny ground; discal bristles

and apical fringe dark brown. Antenna thin and slender, 3 in frontal view

longer than 4 (100: 74), this equal to next segments, all distinctly longer than

broad; scape on both sides with long and dense fringe of pure white. Hair patch

upon clypeus dense, depressed, not nearly reaching anterior border, surface

shiny, punctation as in male. Non-pubescent parts of abdominal tergites as

described for male; band-like lateral markings on 2—6 (sic) occupying most or

all of the surface, the distance separating spots on 2 equal to their transverse

diameter, on 3 less so, and spots on 4—S confluent across middle, occupying

entire surface. Sternites I—5, except polished postgradular hairless areas, very

closely and finely punctate and clothed all over with short, appressed silvery

white tomentum. Pygidial plate, see description and fig. 167. Length 13—14

mimeniabs Uzbekistan and’Turkmenia 9. 2.8 Saisset. out nivosa (p. 257)

Body markings less extensive: abdominal spots not fused together, tergites 4

and 5 not entirely white; pygidial plate shaped otherwise ............. 6

Pygidial plate (fig. 215) of characteristic shape and texture, with conspicuous,

subacute median crest, highest and most swollen just before apex. Median

mesonotal line distinctly sulcate, impression deepest at level of tegulae,

dorsum on either side of median line convex, polished and hairless, with few

scattered deep punctures. Wing membrane very dark brown, except basally.

180

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Face and frons black, thorax, legs and abdominal tergites I—4 marked with

white; pubescence generally short; integument of tergites very finely

reticulated. Large species, length 17 mm, fore wing 12.5 mm (holotype). Male

tinknhownHabs Algetia' ur MP Mae solivaga (p. 279)

Pygidial plate not shaped as above. Strongly fluted median mesonotal line and

partly hairless dorsal thoracic sclerites not always in combination with very

darkforewing membrane. dusig Ata beenen sees Sets 7

Vestiture of whole body recalling Paracrocisa: anterior portion of clypeus,

temples, upper frons, most of the vertex, mesonotum-scutellum, as well as

propodeal area, for the greater part exposed to view, bare, or clothed with

short black hairs which are usually hardly longer or even shorter than scutellar

tubercles. White markings conspicuous, sharply outlined. Pile covering

antennal scape partly suberect, usually white and shorter than diameter of

scape about halfway its length. Whole distal half to one-third of clypeus

exposed, hairless and closely punctate, most of frons, paraocular and ante-

ocellar areas with dense patch of longer, partly suberect, snow-white

pubescence; raised collar-like hairs fringing occipital ridge conspicuous, pure

white. Surface of mesonotum uneven: median line markedly impressed, the

sulcus especially developed anteriorly, partly filled up with an elongate,

compact patch (ms) of decumbent or slightly raised snow-white hairs directed

sideways on each side of line; parapsidal lines usually also indicated by a

longitudinal depression. Vertex, immediately beside and slightly distal to each

lateral ocellus, with impunctate, smooth and shiny (or finely chagreened) and

frequently distinctly concave or pitted, area. Propodeal triangle with small

impunctate median area, varying in size and shape, traversed lengthwise by a

fine impressed line, more rarely devoid of the latter and punctate like rest of

propodeum. Punctation of mesonotum diverse: punctures on anterior surface,

as far back as a little before front margin of tegulae, very close and mostly

contiguous, covered with short, dense depressed hair, behind this surface is

more coarsely, partly confluently punctate, adorned with tufts of suberect

white hairs (/pn-als, small mls, plsa-t, and large pls). Outer faces of mid and hind

tibiae beset with strong black spicules and few longish erect bristles; at least

the basal one-third of each marked with white. Abdominal tergites noticeably

dull, or at most faintly lustrous (less shiny than in males!), all very closely and

finely reticulate-punctate, especially postgradular areas with punctures even

more crowded and larger than interspaces, and all setiferous, the non-white

spotted areas covered with minute black decumbent hairs; tergal graduli with

or without thin fringe of dark bristles, those on the sternal ones usually

suberect, more numerous and longer. Sternites invariably black, lacking white

hairs. Fore wing membrane usually strongly infuscated, never entirely

hyalinesr. han amas us oats Seid vanta OS SE 8

Vestiture of whole body generally longer. Hair fringing both sides of antennal

scape longer, sometimes exceeding greatest diameter of scape. Sculpture of

mesonotum and disposition of white body markings variable, but not as

described. Dorsal surface of abdomen distinctly somewhat glossy, often very

LIEFTINCK: Palaearctic Melecta 181

brilliantly luminous, punctation frequently dense but punctures always smaller

than interspaces, and not necessarily setiferous, though unspotted areas often

clothed with longer, suberect dark hairs and bristles ............... 11

. Pygidial plate very markedly downbent (profile!), its distal portion moreover

distinctly constricted before the end, the apex itself noticeably expanded and

broadly rounded (figs. 172, 178, 184). First abdominal tergite relatively long,

transition between vertical (anterior) and horizontal faces rather abruptly

ROGEN SEREN EM AES LENA AANGE DIEETEN Gee 9

Pygidial plate either gradually downcurved, or almost flat, distal portion

variably expanded, but usually more evenly tapering toward apex, which itself

is rounded off (e.g., figs. 136, 214, 224). First abdominal tergite shorter,

transition between vertical (anterior) and horizontal face less marked, the

dorsal one sloping gradually into declivous anterior face (see key to males) 11

. Silky white spots at sides of tergites I—4 differing in shape and size; those on 1

variable: either tapering inward, pointing toward each other along base, with

very oblique hind border and only narrowly interrupted in the median line

(holotype); or smaller, less than half as broad as the shortest distance

separating them, hence shaped much as in male (pl. 3 fig. 15), but composed of

short appressed pubescence; spots on 2 larger, subrotundate, as in male, on 3

and 4 transverse with rounded angles, separated by a distance slightly less than

their own breadth; all spots lying flush with surface, lacking admixture of

raised hairs. Maxillary palpi slender, 6-segmented, the ultimate joint small

(proboscis retracted in type). Antenna slender, 3 about twice as long as its

apical breadth and 1% times longer than 4 (100: 65), this and next segments

subequal and only little longer than broad. Impunctate area at base of

propodeal triangle vestigial, the lateral edges (with few exceptions)

longitudinally rugose. Scutellar tubercles short, subtriangular, directed straight

back; white tuft projecting from beneath tubercles at least twice as long as

these. Inner ramus of mid and hind tarsal claws only one-fourth length of main

branch or even shorter. Fore wing membrane infuscated much as in amanda

and candida, but much lighter, especially near base, than in honesta and

transcaspica; third submarginal cell a little longer than high, its distal side

variably angulated. Pygidial plate dark brown, slender, fully twice as long as its

breadth at base, more abruptly downbent beyond halfway length than in

honesta but slightly less so than in amanda and candida, apex distinctly, though

smoothly ridged medially (figs. 172, 189); disk flat with raised margins, surface

finely transversely wrinkled. Length not exceeding 14 mm. Hab.: From

Uzbekistan through Turkestan to Pakistan; doubtfully, also Turkey and

TEN rida Kaak a a RUA A Se EEN corpulenta (p. 260)

Silky white tergal marks 1—4 very conspicuous, resembling those of male, even

larger and more angular than in corpulenta and transcaspica, especially those on

1—2 (pl. 3 figs. 16—17). Pygidial plate slender, apex convex medially and

rimmed; disk flat, very finely transversely chagreened, colour brown, apex

obscured (figs. 178, 183—184). Black hairs covering disk of scutellum and

parascutella short, generally shorter than tubercles; brush-like tuft of black

182

-—

—

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

covering parascutella not raised above level of surrounding hairs (profile !).

Maxillary palpus 6-segmented. Antennal segment 3 slightly but distinctly

longer than 4, this and next segments somewhat longer than broad (100: 80) 10

. Tergal spots 1—4 situated in a regular row, | parallel-sided, forming a slightly

oblique quadrangle, diameter of each slightly or more markedly less than the

black area separating them; 2—4 little more transverse with convex inner

margins, those on 4 smallest. Scutellum entirely black, lacking tuft of white

hairs projecting from beneath scutellar tubercles, which are short and directed

straight back. Wings very dark, except basally, fore wing membrane almost as

dark brown as in transcaspica, with low dark bluish and coppery reflex; third

submarginal cell usually a little higher than long, with distal side more bluntly

rounded. Inner face of hind femur irregularly coarsely punctate, punctures

more scattered on shiny ground on area adjoining blunt median ridge. Inner

ramus of mid and hind tarsal claws only about one-third as long as main

branch. Pygidial plate varying in shape, little less than twice as long as its width

between base of lateral ridges, the latter at first outcurved in dorsal view, then

incurved at the constriction, and finally again expanded (fig. 178). Length

14—=18 mm: Habi: Egyptyisrachand:Libyavn. Maartense A honesta (p. 264)

Tergal spots 1—4 less regularly arranged, inner boundary of 1 distinctly more

oblique than outer, acute-angulate midbasally, inner side straight or slightly

convex; spots on 2—3 less high and more transverse than in honesta, those on 4

more approximated. A conspicuous tuft of long white hairs projects from

beneath tubercles, which are very short and slightly raised. Wings much

lighter, fore wing membrane light brown, this colour deepening gradually in

tint toward apex, darkest at end of marginal cell; length of third submarginal

cell about equal to its height, the distal side somewhat more prominent and

abruptly angled. Inner face of hind femur more superficially and finely

punctate, the area adjoining median ridge almost impunctate. Inner ramus of

mid and hind tarsal claws about half as long as main branch and also broader,

more compressed and sharply acuminate, than in honesta. Pygidial plate shaped

similarly, proportions equally variable, but apex always deflected more

abruptly ventrad (figs. 183—184). Length 14—15 mm. Hab.: Turkestan

(terr:typ:);“Sarepta’ vaar ke ak adel la ee amanda (p. 266)

. General appearance, pl. 4 fig. 23. Silky white tergal spots 1—4 quite similar to

those of male but at sides of | spots are farther apart, better defined, narrower

than the space separating them at base, placed transversely and pointing

inward, with some suberect hairs at the bend but lacking raised hairs across

middle of tergite; spots on 2—3 transverse, 2 usually also pointing inward, and

nearly always slightly concave anteriorly, all lying flush with surface (pl. 4 figs.

23—24). Raised black and white hairs covering head and thorax much shorter

than in male, resembling the design of female corpulenta, amanda and honesta

much more closely, though all hairs a little longer than these; black hair

covering disk of scutellum and parascutella also longer and more closely set,

about equal in length to tubercles; parascutella clothed with dense compact

brush of deep black hairs raised above surface level (profile); large transverse

12,

LIEFTINCK: Palaearctic Melecta 183

white tufts behind wing bases and also longish white hairs beside scutellar

tubercles. Hair on antennal scape white and decumbent, though mixed with

some raised black pile equalling diameter of scape. Maxillary palpi 5-

segmented. Fore wing membrane very dark brown with slight greenish, blue

and purplish lustre. Outer faces of mid and hind femora closely punctate on

shiny ground, punctures covering inner faces irregular, more superficial, with

almost glabrous glossy areas beside median ridges; basal half or more of outer

faces of mid and hind tibiae white, for the rest black, only last tarsal segment

with thin appressed white pubescence. Pygidial plate gradually and evenly

downbent, very broad at base, less than twice as long as its basal width, varying

in shape but narrowing gradually distad, apex simply rounded, not or scarcely

swollen or expanded, lacking median ridge; disk brown, feebly convex, very

finely transversely wrinkled (fig. 214). Size moderate to large, length 13.5—18

mm; fore wing 12—13 mm, wing expanse 31 mm (pl. 4 fig. 23). Hab.: see

PIO, DER. renna) AL IAsG Msn se transcaspica (p. 275)

These characters combined not as described above... 12

Pygidial plate short and broad, tongue-shaped with broadly rounded apex (figs.

126, 136, 139), side margins straight or slightly convex in dorsal view, thin and

upturned in dorsolateral aspect; plate gradually and very little downbent from

base to apex; colour reddish to dark brown, the side margins and apex more

obscured, whole surface dull, throughout very finely reticulated, with

occasional interspersion of some minute punctures at base only. Maxillary

palpi 6-segmented. Pubescent pattern much as shown for turkestanica (pl. 2 fig.

9) and many other species. White are: a patch upon clypeus (often partly

replaced by black!); raised tufts behind antennae and bordering occiput;

thorax dorsally as far up as slightly beyond tegulae, with pair of deep black

dots, more or less confluent anteriorly, upon anterior part of mesonotum;

denser tufts to the inside of and just behind tegulae; white mesepisternal patch,

and transverse band-like tufts, interrupted by black laterally, at base of tergite

1; and regular row of compact, depressed spots on either side of 2—4; sternites

ne Mito Ms Ruth ls, honor sud postare 13

Pygidial plate shaped otherwise, narrower, relatively longer and more slender,

Beele pnactateni! mes seamen abe bo sesame 14

. Antennal scape clothed sparsely on each side with short, suberect and

depressed black and white hairs, most of which being not longer than diameter

of scape at apex. Only basal portion of clypeus above clothed with central

patch of white depressed hairs, for the rest black; distal part of surface bare

and densely punctate. Body pubescence relatively short, hair covering head

above, mesonotum and scutellum not longer than transverse diameter of tegula

or exceeding tubercles, except white tufts below the latter which are much

longer than these. Scutellum medially and posterior sclerites closely punctate,

clothed with raised black hairs, but surface well exposed to view; tubercles

rather strong, triangular, longer than broad at base, slightly raised, more rarely

a little downcurved. Legs with short black hair, outer faces of basal one-third

of tibiae and last tarsal segment white; tibiae with distinct admixture of thick

184

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

black spinulose setae. Outer faces of mid and hind femora closely punctate on

shiny ground, punctures on inner faces of different sizes and more sparsely

distributed, with shiny impunctate area extending along median ridge;

posterior fringe of suberect black bristles thin, much shorter than diameter of

femur; bristles at basitarsi also relatively short; inner rami of mid and hind

tarsal claws from one-third to about half length of main branch. Fore wing

membrane strongly infuscated, except at extreme base; third submarginal cell

a little higher than long, distal side moderately angulated (fig. 131). Labrum

distinctly longer than its breadth at base, slightly so about midway length,

usually with incomplete median crest or impunctate line, apical border

projecting a little anterad, ending in an upturned tubercle. Abdominal tergites

and pregradular portions of sternites closely finely punctate and setose on non-

white spotted areas, the hairs minute and decumbent on rather shiny ground,

though visible only at strong magnification; sides of tergites and postgraduli of

sternal surfaces sparsely intermixed (respectively fringed) with somewhat

longer, suberect bristles. Pygidial plate relatively slender, average length-

breadth ratio 100: 64; basal 2/3 of disk slightly convex with well pronounced

raised borders, for the rest quite flat, distal portion lacking median ridge or

with merest indication of same (figs. 136, 139). Length 13.5—17 mm. Hab.: see

mialekandimapalleanr claret er Ae ee ae ee aegyptiaca (p. 244)

Antennal scape on either side with dense fringe of long white hairs, which are

markedly longer than greatest width of scape at apex. Clypeus above with

patch of partly raised white hairs much longer than in aegyptiaca, the tips often

reaching anterior border. Body pubescence longer, many hairs covering head

above, mesonotum and scutellum even exceeding length of tegula and

tubercles, especially long and dense on parascutella, completely hiding

surface. Setiferous punctures on abdomen sclerites less superficial and more

closely set, the black hairs longer and suberect on less shiny ground, well

visible with a hand lens in fresh specimens. Sides of tergites and postgraduli

conspicuously bristly. Legs more hairy; white marks at tibiae and tarsi as in

aegyptiaca, but interspersed black spinules mostly concealed among longer

white and black hairs. Femora more closely punctate, scattered punctures also

along inner face of median ridge, at the same time much more densely hairy,

the posterior fringe at hinder pair at least as long as diameter of femur; raised

black bristles at tibiae and basitarsi of great length. Shape of tarsal claws not

differing from aegyptiaca, equally variable. Fore wing membrane much lighter,

pale brownish, base proximal to forking of main veins hyaline; third

submarginal cell more obviously higher than long, but distal side more abruptly

angulated. Labrum as in aegyptiaca, but apical border straight or even slightly

emarginate, disk not or scarcely developed (if constant?). Punctation of

abdominal tergites and pregradular sternal plates less dense and also more

superficial than in aegyptiaca, the short black setae longer, suberect, giving the

tergites a more glossy appearance; lateral gradular bristles also longer and

more numerous. White tergal spots more compact, made up of longer hairs

expecially those forming arched collar at sides of 1 conspicuous. Pygidial plate

13.

16.

LIEFTINCK: Palaearctic Melecta 185

very similar to aegyptiaca but a little broader, average length-breadth ratio

100: 70; disk a little more convex with upturned and sharply pronounced

borders, apex somewhat more swollen and downbent, median ridge distinct,

though low, nearly always with some scattered punctures at extreme base of

disk (fig. 126). Length 13—17 mm. Hab.: Turkey; Cyprus (terr. typ.); Israel;

ENSENADA AOL ERGs REN VE: megaera (p. 243)

. Pygidial plate distinctly constricted subapically, thereafter expanded, apex

more or less swollen with evenly rounded tip; sides usually strongly rimmed

(figs. 201, 224, 228). Median mesonotal line not or scarcely impressed. At least

a few raised white hairs immediately below scutellar tubercles. Sternites

EA ela, REPETITIE RA SN AP RAEC 15

Pygidial plate without distinct subapical constriction, though often strongly

narrowed after its middle, the apical portion — if slightly narrowed — longer

and more tapered, subparallel-sided with evenly rounded or almost truncated

OENE NORDEN MANE EHER LIG STE ale REN, 17

Large species, length 16—19 mm, general appearance, pl. 5 fig. 28. Fore wing

membrane for the greater part very dark brown with low bluish bronze reflex.

Propodeal triangle (enclosure) with distinct median impunctate area extending

from end to end, usually broadest at base and pointing apicad. Inner face of

hind femur closely coarsely punctate, clothed densely with short hair and long

bristles, impunctate areas unapparent. Tergal spots 1—4 relatively small,

placed widely apart in regular row, only spots on 2—3 distinctly transverse.

Antennal scape fringed on each side with longish, partly white hairs. Clypeus

either wholly black or only with small central dot of decumbent silvery hairs.

Pygidial plate slender, apex expanded (fig. 224). For further details, see

pont HD see map 2 Lijs LA he EELKE VA grandis (p. 286)

Size smaller, body not exceeding 16.5 mm, less robust, abdomen more

elongated. Fore wing membrane infuscated but not conspicuously darkened.

Propodeal triangle lacking median impunctate area. Hind femur not very

densely hairy; inner face scatteredly punctate on somewhat shiny ground only

Mradvance ol-and alongside medianiridpe 35.0. 20. AT Aleen ANNE, 16

Moderate-sized species, length 12.5—16.5 mm (average 14 mm, see, however,

description). Fore wing membrane smoky grey to grey-brown; third

submarginal cell shorter than in male, length about equal to its height. No

distinct juxta-ocellar areas, though few punctures are frequently not

contiguous. White hairs fringing both sides of antennal scape conspicuous,

longer than its diameter. Clypeus above with large, somewhat fan-shaped patch

of decumbent white hairs occupying basal half or more of surface, anterior

portion naked and closely punctate. Raised pubescence covering mesonotum

rather short and thin, nowhere entirely concealing surface, except beneath ill-

limited anterior black dots and deep black tufts covering parascutella. Pygidial

plate (fig. 248) of characteristic shape: fully twice as long as its breadth at base;

margins definitely raised as far as the constriction, at which point the

transverse reticulation of the disk ends, the apex first becoming smooth and

swollen, then slopes down with the expanded rims surrounding the centre

186

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

markedly depressed; in side view the plate bends down gradually, as in grandis;

colour black upon middle at base, covered with setiferous punctures (unlike

grandis), the setae being short and depressed, for the rest dark reddish brown,

borders and apex obscured. For more characters, see description. Hab.: see

MAP Bron IA VEL ees Oe RSE festiva (p. 293)

Resembles festiva in form of pygidial plate but differs considerably from the

two preceding species, as follows. Body vestiture much shorter, corresponding

with that of male; tergites more dullish, punctation fine and dense, punctures

smaller than interspaces. Antenna distinctly longer and more slender, scape

longer, not conspicuously hairy, 3 also more attenuated, 4—12 all markedly

longer than broad. Labrum longer than broad, widest before middle. Clypeus

marked similarly to festiva. Scutellar spines short, pricker-shaped. Legs

noticeably more slender, all limbs longer and narrower; femora sparsely hairy,

inner faces scatteredly punctate on shiny ground; armature at outer faces of

mid and hind tibiae and basitarsi as in previous species, with similar anterior

and posterior fringes of long bristles, respectively; inner rami of mid and hind

tarsal claws about 1/3 length of outer. Wings infuscated, expecially along distal

border. White tergal spots compact, arranged similarly but composed of

shorter, more depressed hairs, those at sides of 1 wide apart and better defined,

broadest basally, the hairs suberect only at sides near base; spots on 2—3

transverse, broadest on 3, almost equalling width of interspaces and removed

more inward than preceding spots, the pair on 4 in line with 3, but much

smaller. Sternites all black, except tiny midbasal streak of white at gradulus of

5. Pygidial plate, fig. 201, almost straight, only apex slightly downcurved

(profile!), lateral rims strong, apex distinctly swollen medially, but surrounding

margin less flattened than in festiva. Hab.: From Turkey and Israel to Iraq and

E. Turkestan ausm aha ae alecto (supposition) (p. 273)

. Median mesonotal line impressed: sulcus distinct though not deep and at least

partly hidden under long pubescence, the line itself extremely fine; parapsidal

lines also distinct and feebly impressed but not longer than half diameter of

tegula. Antennal scape on either side with rather dense fringe of long raised,

mostly white, hairs almost or fully as long as apical diameter of scape. Pygidial

plate evenly and but slightly downcurved, sides converging, apex subparallel-

sided and obtusely rounded. Tergites 1—4 white-spotted. Length 13—16

mmiiusatt< ates. ana Maen a. At A sE 18

Median mesonotal line not sulcate, but if feebly impressed, then the white

tergal spots are not placed in regular row, differing among themselves in shape,

sometimes greatly reduced in size, or absent altogether ............. 19

. Body predominantly black: head with thin tuft of white only behind antennae

and conspicuous fringe bordering occiput; long fringe at inner (anterior) face

of antennal scape also white, the outer rows black. Antenna generally shorter,

all segments broader, 3 slightly less than twice as long as its width at apex,

much less than 1% times length of 4—12, the latter little longer than broad.

Anterior part of mesonotum alternately white and black: a subrectangular

black dot on either side of (slightly narrower) elongate median white patch;

LIEFTINCK: Palaearctic Melecta 187

tufts behind wing bases also white; posterior thoracic sclerites and pleurae all

black. Wings strongly infuscated, fore wing brown, extreme base and along

submarginal cross veins narrowly lighter; third submarginal cell much shorter

than high, proximal and distal sides moderately angulated; hind wing lighter

brown. Legs black, rather slender; hind femur with narrow impunctate anterior

streak along midposterior ridge, pubescence not very long or dense; outer

faces of mid and hind tibiae with small, isolated, subbasal white spot, many fine

spicules at hinder pair, and not very numerous longish raised bristles. Inner

rami of mid and hind tarsal claws scarcely half as long as outer. Shape and

markings of abdomen greatly resembling the type male. Pygidial plate twice as

long as its breadth at origin of lateral ridges (fig. 301), dark brown; whole

surface very finely reticulated on somewhat shiny ground; disk flat but apex

almost parallel-sided with distinctly swollen, low median ridge. Hab.:

EAR Entire daher ont any assimilis (supposition) (p. 318)

Body profusely marked with white. Resembles the male (pl. 2 fig. 12); average

size smaller than grandis (pl. 5 fig. 28), length 16 mm (holotype), 14.5— 16 mm.

Stature much less robust, abdomen more elongate and less expanded, but head

and thorax pattern much like grandis, though white tergal spots, including tufty

patches at sides of 1, occupying more of the surface, spots on 2—3

subrectangular, broad, overlapping side angles, transverse diameter of those on

3 equal to distance separating them, spots on 4 about half their own breadth

apart. Differs further from grandis and several other species (characters

following sequence of supposed importance): inner surface of hind femur

sparsely punctate, smooth and shiny toward median ridge; long bristles few in

number. Wings much lighter, basal portion of fore wing hyaline, only lightly

infuscated distally, darkest (light brown) beyond cells, inclusive of marginal

cell; third submarginal cell somewhat higher than long. Pygidial plate coloured

as in grandis but hardly constricted subapically, more distinctly downbent

(profile), the median ridge restricted to apex and only weakly indicated (figs.

218—219); basal part distinctly punctate. Labrum shorter, subequal to its

breadth at base (fig. 216), but narrowing anteriorly, as in grandis (ratios

100:98:76 approx.). Clypeus above clothed with shorter, silky white

decumbent hairs, its distal one-third or more remaining bare, closely punctate.

Antenna more slender, 3 fully twice as long as its width at apex, 4 only 3/5

length of 3, but 4—12 subequal to one another and more markedly longer than

broad; pile similar to grandis. Scutellar spines robust, oblique and flattened,

slightly raised, but directed straight back. Hab.: Kazakhstan; Turk-

EEE toernee rent Lens. al ansi iui baeri (p. 282)

. Pygidial plate coloured as in grandis but hardly contricted subapically, the

median ridge restricted to apex and only feebly indicated (fig. 274); surface of

basal part distinctly punctate. Stature robust, average size 15.5 mm; thorax

broad and bulky, subglobular, though dorsum but little convex, scutellum

anteriorly almost in line with mesoscutum, tubercles very short, conical or

nipple-shaped, tips hardly or not overhanging subvertically inclined posterior

sclerites. Propodeum coarsely contiguously punctate, triangle with dullish,

188

20.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

impunctate, subrectangular or cordiform median area, usually divided

lengthwise by a fine line in front of propodeal pit; side angles of triangle

lacking longitudinal rugae. Median mesonotal line a narrow, smooth, often

finely grooved ridge, at least partly exposed among the relatively short, raised

hairs. Body pubescence generally shorter and darker than in male. Antenna

slender, 3 almost twice as long as its width at apex, about 1% times longer than

next segments, which are all slightly longer than broad; scape not very densely

hairy roundabout, hairs shorter than diameter of scape. Clypeal patch never

reaching ‘anterior border, extending up to 3/4 distance from base, hairs

decumbent, either entirely silvery white (typical series), or reduced to a central

spot varying in size, more rarely hairs are entirely black. Wings much darker

than in male, fore wing membrane deepest in colour in melanistic populations

from Bulgaria, Macedonia and mainland Greece, whose overall pubescent

pattern is also much darker than in typical (insular) populations. Scutellum

invariably wholly black. White lateral spots on tergite 1 completely isolated,

widely separated by black midbasally; spots in 2—4 in regular row but all

smaller than in male. Labrum not longer than its greatest breadth at middle,

distal portion usually with distinct smooth median ridge. Pygidial plate evenly

and but slightly downbent, frequently flat or almost so, but sides converging

with raised, somewhat incurved margins, apex neither distinctly expanded nor

swollen, flat or feebly convex distally and scarcely rimmed; disk reddish to

dark brown, finely transversely wrinkled (fig. 274). Superficially, rather similar

to extensively white-spotted individuals of italica (= “meridionalis” Gribodo),

inhabiting S France, Iberia and NW Africa (vide postea). These can be

distinguished from tuberculata and previously defined taxa of equal size: (1)

body pubescence generally longer and denser; (2) raised hairs fringing both

sides of antennal scape more abundant; (3) hind femur closely punctate all

round and more densely hairy, the raised hairs especially abundant posteriorly

at median ridge, some being even longer than greatest diameter of femur; (4)

pygidial plate less tapered and more flattened, its apex devoid of a distinct

medianıconvexity, or ridge. Hab. see map 2°. TEE: tuberculata (p. 300)

Stature generally more slender, size often smaller, and membrane of fore wing

frequently less strongly obscured. Combined characters not as above .... 20

General appearance, pl. 2 fig. 10, closely resembling male from same locality

and of equal size (length 16 mm, wing expanse 28.5 mm approx.). Wings

subhyaline, only broad area beyond cells of fore wing slightly infuscated. Body

pubescence long and dense. White tuft surrounding scutellar tubercles

composed of long, radially arranged hairs, only narrowly divided by black;

tubercles moderate, conical, considerably shorter than surrounding tufts.

Labrum rather long, only little broadened halfway length (midlength-greatest

breadth ratio 100 : 58), apex slightly narrower than width at base, border

shallowly emarginate; apical 3/4 with distinct shiny median ridge. Silvery patch

upon clypeus longer than broad, restricted to middorsum, depressed hairs not

quite reaching anterior border. Antenna slender, 3 almost twice as long as its

apical breadth, 4—12 all distinctly longer than broad; raised hair fringing both

21.

22;

LIEFTINCK: Palaearctic Melecta 189

sides of scape white, long and dense. Summit of head closely punctate lacking

distinct shiny or impressed juxta-ocellar areas. About upper half of thoracic

sides white, rest and beneath black, as also middle of metanotum and

propodeum. Black anterior mesonotal spots isolated. Legs rather densely

hairy; inner faces of mid and hind femora irregularly punctate, punctures near

apex of hinder pair few in number on shiny ground. Inner branches of mid and

hind tarsal claws relatively long and slender, exceeding half length of outer

(much shorter in sibirica!). Tergites moderately shiny, setiferous punctures fine

and crowded, much smaller than interspaces. White collar of raised hairs at

base of tergite | complete, condensed and spot-like laterally, though smaller

than very compact, sharply defined lateral spots 2—4, which are depressed,

raised above surface level. Shape and texture of pygidial plate normal, i.e., as

in /uctuosa, the convex median ridge at apex distinct, rather long and high, well

visible in profile. Hab.: Turkestan (Uzbekistan) and “Alai” ............

EE NA re aos Sus bear i eee tes A turkestanica (p. 239)

Facies and combined characters not as above. Fore wing membrane usually

darker and body pubescence often lesswhite. int. seren Suge es 21

Scutellar processes (fig. 303) parallel, 0.4—0.5 mm long, broad at base,

directed slightly upward but soon tapering and distinctly downbent to form

cylindrical, bluntly rounded, shiny hooks with scattered punctures near base,

partly hidden among surrounding black hairs, but much shorter than

conspicuous white (more rarely black) tufts emanating from beneath each

spine. Body small, 10—11.5 mm, compact, luctuosa-like; pubescent pattern

similar, but white spots at outer faces of all tibiae and subrectangular patches

at sides of abdominal tergites I—4 generally larger, composed of more closely

set and somewhat longer decumbent hairs, the spots raised above level of

surface, hair tips downcurved, exceeding hind border of tergites. Setiferous

punctures covering tergites much finer and more widely spaced on shiny

ground, than in most species of equal body size discussed hereafter. Antennal

scape fringed with long hairs, 4—12 all slightly longer than broad, increasing

gradually in length toward end. Pygidial plate, fig. 309. Hab.: Canary Is. (in

Pane MIND Re NEBr ao RN EE curvispina (p. 320)

Scutellar processes very variable in shape and length, occasionally tubercular

or wanting, never distinctly downcurved and hook-like ............. 22

Body pubescence much obscured: head (including antennal scape), thorax and

legs wholly black, only basal half of clypeus with ill-limited dorsal patch of

decumbent dark golden brown hairs, all the rest of clypeus remaining hairless,

closely punctate on shiny ground. Sides of abdominal tergite | with tuft of long,

raised, entirely black hairs, but lateral spots on 2—4 white, depressed, of

irregular small size, just visible from above, the pair on 2 transverse, about

twice as large as on 3—4. Fore wing membrane not much darkened, only

lightly suffused with grey-brown. Pygidial plate shaped similarly to many

luctuosa: slender, almost twice as long as its breadth at base, scarcely

downbent; disk chestnut brown, impunctate, finely transversely wrinkled, apex

smooth, subparallel-sided with rather long convex median ridge clearly visible

190

23:

24.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

in profile. Body more slender and less hairy than /uctuosa, abdomen distinctly

more strongly convex, resembling brevipila and leucorhyncha much more

closely in form than the broader and flat-bodied italica and luctuosa. Labrum

only little longer than its widest point near base, narrowing gradually toward

distal border, which is a little upturned, subtruncated, very shallowly

emarginate, lacking complete (apical) median ridge. Dorsal punctation of

ocellocular area and on disk of mesonotum much as in /uctuosa, punctures

rather deep but slightly smaller than shiny interspaces, those on abdominal

tergites much finer and more widely spaced. Only a single specimen known.

Size small, 11.5 mm, fore wing 9.5 mm. Hab.: Kiev area, S. Russia; Turkey (see

MAP) MP ALTE, CIDRE STONE AES Sate CIONI rutenica (p. 233)

Combined:characters not as above. ws. seo eats TI oe eee eee 23

Vestiture of whole body totally black (minus variants) or almost so (plus

variants), the latter having (1) a tiny white spot at outer faces of mid and hind

tibiae at some distance from base, (2) vestiges of white in front of tegulae and

more inward posterior to the latter, (3) tufts behind hind wing base and at sides

of tergite 1, and (4) a speckle at sides of tergites 2 and 3. Wing membrane

throughout dark brown with slight bronze lustre in certain lights, this colour

deepest along costa, in whole marginal cell and centres of submarginals; third

submarginal somewhat higher than long, outer side not strongly angled;

nervellus (hind wing) well proximal to fork. Pygidial plate gradually

downcurved (profile), broad at base (length-breadth ratio 100 : 70), converging

as far as about 3/4 length from base with rather outbent (fig. 320) or almost

rectilinear sides, the long slender apex almost parallel-sided, with distinct blunt

median ridge (fig. 320); disk flat, finely transversely tessellated. Short suberect

black pubescence covering tergites 2—4 evenly distributed, without forming

condensed dots replacing white hair spots. Size small, 10—13 mm, fore wing

8150 mm. Habe Canary ds. (Gnipart) na Se ee ee ee caroli (p. 323)

Vestiture of body more extensively marked or white- spotted; or, if all or most

parts are black, then lateral spots on tergites 2—4 form condensed dots of

appressed black pubescence replacing white hairs. Wing membrane usually

much lighter (darker brown only in leucorhyncha taormina from Italy/Sicily).

Pygidial plate variable but usually not as described, tapering more gradually; if

plate is as slenderly drawn out as in caroli, then compact white spots are

present at sides of tergites (1) 2—4, while head, antennal scape, thorax and legs

are also tufted and spotted with white (certain /. leucorhyncha and plus variants

Of ttalica) ats setta LA ARSE EVES. IOM ee 24

Stature robust, abdomen broadest at base of tergite 2, then rather flat,

subtriangular, tapering rapidly toward apex. Vestiture throughout long and

dense, raised hair covering hind femur posteriorly conspicuous, at least basally

longer than greatest diameter of femur. Inner faces of mid and hind femora

anterior to median ridge closely punctate, but distal portion lacking distinct

impunctate glossy strip. Mandible with flattened, rounded, submedian interior

tooth (fig. 78). At least distal one-fifth of labrum with smooth median carina.

Antenna strong, median segments only little longer than broad; scape densely

LIEFTINCK: Palaearctic Melecta 191

fringed with raised hairs laterally. No distinct impunctate areas beside ocelli;

propodeum throughout closely rugoso-punctate. Scutellum black, no longish

tufts of white surrounding short, slightly curved, blunt processes. Inner rami of

mid and hind tarsal claws almost or fully half as long as outer. Wing membrane

evenly suffused with brown, much as in /uctuosa; third submarginal cell

relatively long, only little shorter than high, distal side not markedly more

angled than proximal side. White body marks extremely variable in shape and

size (see descriptive comments in text), sometimes much reduced with hairs

largely replaced by black. Pygidial plate relatively broad, not or only very

gradually downcurved, disk almost flat, brown, very finely transversely

reticulate or chagreened; sides almost rectilinear, margins raised, obscured,

less strongly converging than in next species, not or feebly constricted before

apex, the latter short, gradually tapered or subparallel-sided, with low, short

megan ridge (figs. 2835287) 1Hab;: See:map 3) a. rw... 29% italica (p. 309)

Characters combined not as above. Shape of pygidial plate variable but, if as

described for italica, then body pubescence is shorter and less dense, hairs

covering hind femur posteriorly more sparsely distributed and shorter than

greatest diameter of same. Inner rami of mid and hind tarsal claws distinctly

less than half as long as outer, often very short. Scutellar processes often

surrounded by tufts of white. Distal portion of posterior faces of hind femora

with shiny, almost impunctate area, or with scattered punctures only. Stature

generally more slender and average size smaller. (Includes brevipila, which has

the inner rami of mid and hind tarsal claws fully half as long as the outer, and

Migusiolick-haired thoracic sides!) ran SCE m RENTE BR RN 25

. Apex of pygidial plate in dorsal view slender, narrow and parallel-sided, with

well marked blunt median ridge, the much longer proximal part (ratio of

lengths 100 : 35.7) suddenly broadening toward base of tergite, the sides feebly

convex; in profile the plate gradually but noticeably curves ventrad, the

swollen convex median ridge being distinctly visible (fig. 102). Labrum shorter

than in male, with distinct, smooth median ridge, incomplete basally. All

flagellar segments of antenna longer than broad. Silvery patch of appressed

hair upon middle of clypeus short and thin, not nearly reaching anterior

border. Scutellar tubercles strong, pricker-shaped. Body pattern /uctuosa-like,

but all white marks smaller; thoracic sides all black. Wing membrane not much

obscured. White tergal spots 2—4 subcircular, placed in regular row. Length 11

approx Labs Turkestan 4 ma EIER brevipila (p. 235)

Apex of pygidial plate in dorsal view less drawn out, usually passing

imperceptibly into broadening proximal portion of plate; if apex is longer and

more slender (e.g. some leucorhyncha, fig. 324), then the plate is less downbent

with much lower mid-apical ridge not (or hardly) apparent in profile, and the

fore wing membrane is strongly infuscated (brown). Vestiture of all body parts

shorter than italica (see couplet 24). Inner face of hind femur, alongside and

upon its median ridge, with ill-defined, elongate, shiny impunctate area

extending from about midway length to almost as far as the end (sometimes a

IEMSCat er CU DUNCEUrES present). AN MENT A LL ent EEE, 26

192

26.

27.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Head, thoracic sclerites and legs all jet black, much as in caroli, only about 20

per cent or less of any population with tiny white spot near base at outer face of

mid and hind tibiae, and a small tuft of white upon parascutella and/or behind

base of hind wings. Fore wing membrane smoky brown, darkest distally and

toward apex of anterior main veins and cell centres. Abdomen either entirely

black (most ex. from Sicily), or with small lateral dots of unequal and variable

sizes only on one or more of the tergites (typical /. taormina and its varieties); or

else, with fairly regular row of sharply pronounced, compact, subcircular dots

on either side of (1) 2—4, diminishing gradually in size posteriorly (most ex.

from Italy, Corsica, Sardinia, some from Sicily, and Malta). These tergal spots,

when present, never raised above surface level, those at sides of 1 composed of

much longer, partly raised hairs (always replaced by black upon middle at

base). Raised bristles placed at graduli of tergites not very long. Hair fringing

antennal scape shorter than greatest diameter of same; segment 3 from % to

fully % times longer than next segments, all of which being a trifle longer than

broad. Pygidial plate gradually tapering, disk flat or almost so, side margins

somewhat raised, with or without short, weak midapical ridge, apex of plate

bluntly rounded; surface dull, impunctate, very finely transversely wrinkled;

colour chestnut to almost black. Size small to medium, 10-14 mm (average 11.5

mm). Hab:: Italy and Tyrrhenian Islands (see map 3) rar zen oe ene

ns Peat fare ee ee dea ssd ch leucorhyncha taormina (p. 330)

Head, thoracic sclerites and legs variegated black and white or greyish white:

mesonotum white anteriorly, frequently with pair of black dots, and sides at

least partly white. Fore wing membrane often distinctly infuscated but much

less obscured than in taormina. Relative lengths of antennal segments and

shape of pygidial plate greatly varying among local populations of each taxon,

but pygidial plate usually slenderer, more abruptly narrowed toward apex. Size

small,as before, length notexceeding, 13 mm, often less in gee

nente REN okee dated: MWE Group of luctuosa ... 27

Besides /uctuosa, the earliest described species of the group, this cluster of

near-alike Melecta females includes the nominate subspecies /. leucorhyncha

Gribodo, and sibirica Radoszkowski, three taxa whose males are quite distinct

and fairly well characterized in the specific key. The four remaining species

are: the little known canariensis Lieftinck, and probably also some of those

discussed in the Appendix as species of doubtful status, viz. ashabadensis

Radoszkowski, eversmanni Radoszkowski, and kashmirensis Nurse, all validly

proposed names for species whose males have remained unknown. Of these,

the female eversmanni is the only one that can be recognized by the rather

characteristic shape of its pygidial plate (fig. 112), all others are variable in this

and other respects, being practically indistinguishable. Brief descriptions of

luctuosa (figs. 95—96) are to be found only in a number of faunistic publications

dealing with bees occurring in Great Britain and NW. Europe, one of the most

recent being contained in a paper on Canarian Melecta (Lieftinck, 1958). For

illustrations and notes on most others, see the text: ashabadensis (fig. 113),

canariensis (fig. 312), and I. leucorhyncha (figs. 324—326 and 350).

LIEFTINCK: Palaearctic Melecta 193

I. THE Melecta albifrons ALLIANCE

1. Group of M. albifrons (Forster)

Melecta albifrons albifrons (Forster)

(figs. 3, 19—25, 71)

Selected references:

Apis albifrons Forster, 1771, Novae species insectorum. Centuria I: 94 (pars: 3 England only, lectotype

coll. Linn. Soc. London).

Apis punctata Fabricius, 1775, Syst. Ent.: 385 (type series 3 9 England, lectotype ¢ Univ. Zool. Mus.

Copenhagen).

Andrena armata Panzer, 1799, Fauna Insect. Germ. init. 6 Heft 70: 22—23, pl. 22 (insect, & ? type lost).

— Saunders, 1888, Index to Panzer’s Fauna Insect. Germ. London: 8 (cat.).

Melecta armata: Lepeletier, 1841, Hist. Nat. Ins. Hym. 2: 444—445 (9 ¢ “Environs de Paris, Pyrénées.

Commune en France’). — Pérez, 1883, Actes Soc. linn. Bordeaux 37 (ser. 4, vol. 7): 302—305 (par-

tim). — Gribodo, 1893 (f. typica, exclus. vars.!), Bull. Soc. Ent. Ital. 25: 413 (Europe, various locs.).

— Friese, 1895, Bienen Europa’s I: 156—159 (partim). — Saunders, 1896, Hym. acul. Brit. Is. Lon-

don; 345 (key), 346—347, pl. 45 fig. 3, 8, 11 (Gr. Britain). — Peets, 1910, Jahresber. Nat. Ges. Han-

nover: 42 & 53 (note on identity of A. armata Panzer). — Frey-Gessner, 1910, Hym. Apidae in Fauna

insect. Helv. 2: 143—144 (39 various locs.). — Jorgensen, 1921, Bier in Danmarks Fauna

Kgbenhavn, 25 (Danish recds). — Semichon, 1922, Bull. Soc. Ent. France, 14: 192—194, 2 fig. (lar-

va); 1925, ibid. 18: 305—306 (larva). — Schmiedeknecht, 1930, Hym Nord u. Mitteleuropas: 830. —

Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21: fig. 14 (fore wing, sex?). — Osychniuk, 1970,

Fauna Ukraini, 12 Apoidea 4: pl. 23 fig. 8 & pl. 24 fig. 1 (struct. 9, Ukraine).

Melecta punctata: Friese, 1893, Bienenfauna Deutschl. u. Ungarn: 37 (“Mecklenburg, an Hyacinthen;

Thüringen; Elsasz: Strassbg., with Anthophora personata; Süddeutschland’), 61 (Budapest). — Mori-

ce & Durrant, 1915, Trans. Ent. Soc. Lond. 2: 424 (note). — Richards, 1935, Trans. R. Ent. Soc.

Lond. 83: 172—173 (notes on ¢ 9 Fabrician types: A. punctata considered synonymous with Andrena

armata Panzer; M. punctata (F.) reinstated). — Lieftinck, 1958, Comment. Biol. Helsingfors 18.5:

20—22 (general notes); 1972, Tijdschr. Ent. 115: 256 (host relation).

Melecta armata var. gigantea Friese, 1925, Konowia 4: 27 (g 9 Elsasz, various locs). Syn. nov.

Melecta albifrons: Day & Fitton, 1977, Biol. J. Linnean Soc. London 9: 33, 40 (notes, designation of lec-

totype & (not © as stated!) A. albifrons Forster, in coll. Linn. Soc. London; A. punctata F., syn. nov.).

Type and syntypic material. — Great Britain: 1 ¢ (abd. loose in cellophane

capsule), labelled “Apis albifrons var. Anglicana” (handwritten), purple disk,

“Lectotype”, and additional identification label “lectotype A. albifrons Forster”,

both by M. C. Day (Linn. Soc. London). — [Great Britain]: 1 & (diss, figs. 21, 25

& 27), labelled “A. punc/tata’’, with red lectotype label A. punctata Fabricius, by M.

A. Lieftinck, and 1 © (syntype A. punctata Fabricius), both with O.W. Richards’

labels ‘‘Melecta punctata (= M. armata Pz.) (MC). — W.Germany | ©, labelled

“Strassbg 16.6.87, Friese, M. armata var. gigantea Friese” (pencil writing),

lectotype M. armata var. gigantea Friese by present selection (MNB); 1 &,

“Strassb. III.1887, Friese, M. armata & var.’’, det. Friese 1900, syntype var.

gigantea Friese (SMF); 2 6, “Elsasz, 13.v.1887, Friese”, and ‘Strassburg,

21.v.1887, Friese”, both unidentified but obviously syntypes of gigantea Friese

(MCG).

For the oldest and correct name of this well-known bee, see Day & Fitton

(1977), whose lectotype selection should be accepted. It is clear that M. albifrons

now becomes the type species of Melecta Latreille. Forster’s type, probably

194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

collected near Warrington (N Cheshire), in central England, is a male which

differs in no way from continental European examples of that sex. The abdominal

spots on tergite 2 are comparatively of large size, resembling those of the lectotype

of punctata, which was also described from Britain. It must be said that albifrons is

a veritable misnomer, applied to a bee in a nondescript diagnosis. Although the

hairs in front of the head and other parts of the body may with much fantasy be

called ‘“whitish” in deference to the describer, the overall impression of the long

pubescence is distinctly brownish. Forster’s diagnosis is based on English, Spanish

and German specimens, the selected lectotype standing over his label “A. albifrons

var. Anglicana”, the one labelled “A. albifrons var. Hispanica” being a 9 Thyreus

histrionicus (Illiger), while the second (unlabelled) example is a 7. orbatus (Lep.).

My descriptive notes on the typical examples of Apis punctata Fabricius, written

down while studying both sexes in 1958, need not be reproduced here, but outline

sketches of some structural details made at that time may still prove useful. The

somewhat aberrantly shaped 7th tergite of the lectotype is shown in fig. 21: instead

of being more or less squarely cut off — the normal condition, shown in other

figures — the exposed apical part is more hollowed out dorsally and bears a

crescentic incision. The form of this sclerite varies, however, within certain limits,

in the entire albifrons complex, as is the case with the contour of the last visible

(6th) sternal plate (figs. 25 & 27).

The synonymy of Apis punctata Fabricius, 1775, and Andrena armata Panzer et

auct., was first established by O. W. Richards (1935). The male of Fabricius type

specimens (1 & 1 9) was selected lectotype and labelled as such by me in 1958.

This was left unpublished so far but should now be confirmed.

With regard to Panzer’s ‘Bewafnete Waldbiene”, Andrena armata Panzer, 1799,

some doubt has arisen as to whether Alfken’s assumption (in Peets, 1910) was right

that it stands for a male of Melecta armata auct. The picture shows a bee quite

unlike the present species, but because we shall, of course, never be sure of its

identity, any suggestion would be mere guess-work. Panzer’s later interpretation

(1804: 34—35) of Schaeffer’s picture of “Apis sexta” (1766, tab. 22 fig. 14) must

remain equally uncertain, as it shows a much greater resemblance to a male of

M. a. albovaria Erichson or, even more so, to M. luctuosa (Scopoli). In the same

work, Panzer also compared the picture with the description of Nomada?

duodecimmaculata Rossi, which he rightly thought to be a different species (see

under that species and M. luctuosa).

I believe that the characters used for a new group definition, summarized in the

opening paragraph of the key on p. 145, are such as to justify the removal of the

albifrons series from other Eurasian members of the genus. In addition, several

more features proper to the taxa of this particular group could be added to those

already known and employed in the same key. The discovery of new characters

not shared by most of their congeners has led to the conviction that no useful

purpose would be served to comment upon earlier attempts to clarify the relation

between the present species (i.e. armata olim) and non-related taxa such as grandis,

luctuosa and several others compared with it, whose essential structural characters

remained unstudied by authors like Pérez (1883) and Gribodo (1893).

LIEFTINCK: Palaearctic Melecta

SEE

LES

Figs. 19—25. M. albifrons albifrons; 19, left antennal segments 3—5, frontal view (¢ lectotype Apis punc-

tata, England); 20, fragment of right fore wing (same specimen); 21, dorsal view of apex of tergite 7,

showing slightly aberrant emargination, sculpture and hair omitted (same specimen); 22, right lateral

and dorsal view of partly exposed pygidial plate (@ syntype Apis punctata, England); 23, right hind tarsal

claws, lateroventral view (Netherlands); 24, ventral view of tergite 7 (same specimen); 25, ventral view

of apex of sternite 6 (same specimen). — Figs. 26—29. M. albifrons nigra (3 lectotype M. fasciculata,

Genoa, Italy); 26, dorsal view of apex tergite 7; 27, ventral view of apex sternite 6; 28—29, veniral view

of sternites 7 and 8, with apices of both more enlarged (lower fragments in figs.)

195

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Biology. — The earliest and most concise account of the biology and host-

parasite relationship of M. albifrons is the one summarized by Friese in his classical

work “Bienen Europa's” (Apidae europaeae), 1895: 154. It was based on an

investigation of the cells of Anthophora fulvitarsis Brullé (= personata Illiger, olim),

a large species, which in those days (1888) nested in loess (loss) banks and formed

large colonies in the vicinity of Strasbourg (Alsace). This big species was

parasitized by M. armata var. gigantea Friese, which is, indeed, only an oversized

local variety of typically brown-haired albifrons, adapted to its equally large-sized

host, both probably restricted to some particularly suitable places, inasmuch as

normally sized individuals of albifrons, possibly committed to smaller species of

Anthophora, were also found in the suburbs of that town.

Distribution. — The marginal populations occurring within the area occupied

by the nominate subspecies albifrons, are discussed under albovaria and nigra, two

fairly distinct geographical races which, I believe, can be maintained in spite of the

poorly defined limits of their distribution area. Numerous intergradations are

known to occur in the borderline zones, radiating almost in any direction. The

northern and eastern limits of the range of a. albifrons, which is apparently centred

in Middle Europe, are very insufficiently known, and it is not possible either to

precisely define those of the other subspecies. This is why no maps can yet be

provided to indicate the territories occupied by each of them.

Melecta albifrons albovaria Erichson

(figs. 1, 7, 31—37, 72, pl. 1 figs. 1—2)

Melecta albovaria Erichson, 1840, in Wagner, Reisen Algier 3: 192 (¢ Algier). Stat. nov. — Lieftinck,

1958, Comment. Biol. Helsingfors 18 (5): 22 (note on ¢ type, Algeria).

Melecta punctata: Lepeletier, 1841, Hist. nat. Ins. Hym. 2: 441—442 (pars, 9 & “toute la France”). —

Lucas, 1849, Explor. sci. Algérie 3 Hym.: 221 (Algérie; various locs., no descr.; partim?).

Melecta armata & var. grandis: Pérez, 1883, Actes Soc. linn. Bordeauz 37 (ser. 4 t. 7): 302—307 (pars

Q 3; compar. notes).

Melecta pseudoarmata Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 183, fig. 37

a-c, i,k (Z struct., “® g Caucase: Lagodechi”’). Syn. nov.

Melecta armata Var. Mediterranea Gribodo, 1893, Bull. Soc. ent. Ital. 25: 413—414 (pars! © ¢ “Francia

meridionale (Montpellier) e dall’ Algeria-Algeri?’’). Syn. nov.

Melecta armata var. grandis: Friese, 1895, Bienen Europa’s 1: 159 (key, pars; ‘“Sideuropa, Nordafrika”).

Melecta luctuosa var. albovaria: Friese, 1895, Bienen Europa’s 1: 163 (pars! “Südeuropa’’).

Melecta armata: Dusmet y Alonso, 1905, Bol. Real Soc. esp. Hist. Nat.: 152—153 (pars? Spain, locs.).

Melecta luctuosa Stammform 9: Alfken, 1914, Mém. Soc. ent. Belg. 22: 236 (9 Algerien).

Melecta Novellai Dusmet y Alonso, 1915, Mem. Real Soc. Esp. Hist. Nat. 8 (7a): 330—331 (9 ¢ Marra-

quesh, Morocco). Syn. nov.

Melecta luctuosa var. minima Friese, 1925, Konowia 4: 28 (¢ Algerien). Syn. nov.

Type and syntypic material. — Algeria: | ¢ (diss., figs. 34—35), labelled “Bone

Wagnler]”’ (written, white), ‘‘a/bovaria Er.* 226” (same writing), “Type” (print on

red). Lectotype M. albovaria Er. by present selection (MNB). — Portugal: I 9,

labelled “Portugal (Lusit.)” (written in light green) “726” (print), “Type” (print on

red), “Albovaria N. Erichs.* Wagner, Algier/Lusit.” (all written on one label,

Erichson’s handwriting), ‘‘a/bovaria Erichs.” (written). Paralectotype (syntype)

M. albovaria Erichs. (MNB). — USSR: 1 & (diss, left antenna missing, genit. glued

LIEFTINCK: Palaearctic Melecta 197

on card), labelled ‘“Caucasus” (print), “Type” (old print on red), “Pseudoarmata”’

(Radoszkowski’s handwriting), “‘Melecta pseudoarmata Radoszk. Type Dr.

Enderlein”. Lectotype M. pseudoarmata Rad., selected by Enderlein and here

confirmed (MNB). — Tunisia: 2 ©, labelled: “Tunisi, Belv[edere] 12.111.1882, G.

& L. Doria”, and ‘‘M. armata 3 Q Panz. var. mediterranea Grib. D. Gribodo”

(both in Gribodo’s writing on blue). Lectotype and paralectotype M. armata var.

mediterranea Gribodo by present selection (MCG). — Morocco: 1 &, 9, both

labelled ‘““Marruecos Marraquesh, IV.1907, Escalera” (print), “Melecta Novellai

Dusm. GQ” (Dusmet’s writing), “de la collección Escalera” (do.), “Typo”

(written on white). Lectotype G and paralectotype 9 M. novellai Dusmet, by

present selection (IEM).

The above synonymies are partly new and require a brief explanation.

M. a.albovaria. — Since the earlier discussed male from Bone was meant to

represent the “holotype” (Lieftinck, loc. cit., 1958), that specimen should be

declared more explicitly lectotype of the present taxon. Male and female, from

Algeria and Portugal respectively, are both in the Berlin museum (MNB).

M. pseudoarmata (Caucasus). — This proves to be a medium-sized, entirely

white-haired specimen of albovaria, as was to be expected, judging from

Radoszkowski’s sketches of the male terminalia.

M. a. var. mediterranea (Tunisia). — Correctly placed as a variety of “armata”.

The two females in the Genoa museum, named by Gribodo himself, conform in

size to large individuals from other North African localities. Though only

Montpellier and Algeria are given as localities, Gribodo’s remarks preclude any

possibility of confusion with his meridionalis (= italica Radoszk.) and/or

leucorhyncha, two other new names introduced previously to denote varieties of

“Juctuosa” (sensu Gribodo): ‘‘L’esame però della scultura, e della forma della

piastra epipigiale, delle antenne, oltre alla struttura ed armatura delle gambe li

fanno immediatamente distinguere” (loc. cit: 414, the spacings are mine). The

male from Montpellier is evidently one of the bees collected by Lichtenstein and

identified with M. grandis by J. Pérez.

M. luctuosa “Stammform” (Algeria). — It will be explained sub M. italica and

leucorhyncha that luctuosa was misinterpreted by Alfken: by so doing he introduced

a superfluous new varietal name for one (or both) of the two taxa just mentioned.

In point of fact not one of Alfken’s Algerian specimens still available for

comparison proved to be conspecific with luctuosa !

M. novellai (Morocco). — As with the type(s) of albovaria, my selection of the

male lectotype seems warranted. The female syntype is undoubtedly conspecific.

Both agree perfectly with the original description, as far as it goes.

M. luctuosa var. minima (Algeria). — The type male, labelled as such in the

Berlin museum collection, is a dwarfish example of a/bovaria and has nothing to do

with /uctuosa. The specimen is one of the smallest known, measuring only 7.3 mm

for the body. The author merely says of it: “in allen plastischen Merkmalen mit

M. luctuosa übereinstimmend (Analsegment, Metatarsus), aber Segment | schwarz

behaart, 2—5 seitlich mit weisshaarigem Bindenfleck: Antenne dunn, normal.

Länge 7% mm. Breite 2% mm. ¢ von Algerien.”

198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Further material (roundabout Mediterranean basin, W to E). — Morocco: 1 &, Maroc, Oulmes,

iii.1961, Meinander (MH); 1 ©, Maroc, Midelt, 21.v.1947, J. de Beaumont (CB) and 1 9, Midelt,

18.v.1965, W. Linsenmaier (CL); 1 &, Ifrane, 13.v.1944, K. M. Guichard (BM); 9 & (1 diss.), Azrou,

1250— 1400 m, 20—24.v.1966 and 27.v.1968, and Dayet Aaoua nr. Imouzzèr, 1350 m, 26—29.v.1968, all

M. A. Lieftinck (ML); 1 9, Casablanca, 3—11.iii.1919, R. Benoist (MP); 1 ©, Maroc, 622.19 (MG). —

Algeria: 1 &, N.Afrika, Constantine, 17.v.1929, H. Bauer (ZSM); 1 &, blue disk, Constantine, coll. J.

Pérez (MP); 1 9, Sétif, M. grandis, det. Friese & M. funeraria, det. Radoszkowski (MNB); 2 9, Sétif,

vi.1891, Handlirsch, M. armata var. grandis Lep., det. Friese 1893 (MNW); 1 ©, Alger, coll. de Saussure

(MG); 1 &, Oran 1910, Schmiedeknecht (SMF); 1 &, Alger (MCG); 1 g 1 9, Algerien, Les Tamarins,

10.v.1954, W. Linsenmaier (CL); 1 g, Algir, Oran 1890 (MBUD); 1 & (very small ex.), Oran,

Lepeletier, sub M. notata Klug (MT); 1 9, Sa. Cruz, Oran, J. Bequaert, M. albovaria det. Alfken 1933

(MNB); 3 ©, Oran, coll. Ernest André 1914 & J. Pérez (MP); 1 &, 276/3/M. armata var. grandis, Sidi-

bel-Abbes, coll. Drescher, ded.v.1936 (MKB); 1 4, 123 (square label), coll. Pérez 1915, M. armata det.?

(MP); 1 g, Alger, Husseyn-Dey, J. Bequaert, M. albovaria det. Alfken 1933 (MNB); 2 g, Bone, coll. J.

Pérez (MP); 1 g, Bône, coll. O. Sichel 1867, Mars 1864 (MP; 1 &, “Algérie, L. Dufour, coll. O. Sichel

1867, M. grandis Lep. Alg., det. par L. Dufour” (old written labels in L. Dufour’s writing (MP); 1 & 1

2, (B green disk), Ain Draham & Biskra, coll. J. Pérez (MP); 2 9, Biskra, 16.11.1897 & 20.iii.1894

(BM); 1 g, Sidi-bel-Abbès, ex coll. Pittioni, sub M. albovaria (BM); 1 ©, Ain Sefra 1931, R. Arlé,

M. armata var. grandis det.? (MP); 1 9, Sétif, C. Saussure det.: M. grandis Lep., coll. O. Sichel 1867

(MP); several 49, Algier, Mascara, Cros, 1 © “ex larva, 8 avrile, Cave Prade”, and 1 & “ex larva, 1

avrile 1907” (sic) (MNB); 1 9, 59 (green disk), M. grandis Bg. 3757, leg. & det. O. Sichel 1867 (MP); 1

©, Laghouat, Sahara algér., coll. O. Sichel 1867 (MP); 1 & 1 9, S. Algérie, Laghouat, iii—iv.1929, Dr.

R. Meyer, ex coll. Pittioni (BM); 1 ¢ 19, S. Algérie, Laghouat, iii—iv.1929, R. Meyer, M. albovaria

det. Alfken (MNB); 2 ©, Algeria, Laghouat, iv.1911, M. armata Panz., det.? (BM). — Tunisia: 1 9,

Tunesia, Tozeur, iii.1978, K. Guichard (BM); 1 ¢, S. Tunesia, Gabes, 1.iv.1925, J. Omer Cooper (BM);

2 ©, Gafsa, Birò (MBUD); 1 4, Tunisi dint.IV.1882, G. & L. Doria (MCG); 2 g (1 diss., fig. 36 & pl. 1

fig. 1), Tunis, coll. J. Perez (MP); 1 &, Médinine, coll. J. Vachal 1911 (MP); 1 ©, blue disk, Tunis, coll.

J. Pérez (MP); 3 9, Tunisie, Gafsa, A. Weis 1904 (MP); 2 9, Tunisie, Tozeur, C. Dumont 1921 (MP). —

Libya: 1 g 1 9, Tripolis, M. armata v. albovaria = grandis det. Friese 1896 (MNB); 1 g 1 9,

Tripolitania, Zliten, 11.iii.1952, Garian Hills, 31.iii.1952, K. M. Guichard (BM); 2 g, Cyrenaica,

Cyrene, 1800 ft., 26.iii.1954, K. M. Guichard (BM); large series ¢ 9, Tripoli, 30.xii.1954—4.iii.1955, K.

M. Guichard (BM, ML). — Egypt: I & (diss., fig. 37), Aegyptus, Cairo (MBUD); 1 & 1 9, labelled

“Egypt Waltl caesia Ill.’ and ‘‘caesia Waltl-Egypt” (MT); 2 &, Egypt, Cairo (ML); 1 &, Egypt, Cairo,

M. armata var. mediterranea (red label, det.?) (ZSM); 2 ©, Wan (?) 1912, Kulzer, one with Melecta spec.,

det. Blüthgen 1947 (ZSM, ML); 1 g, Le Caire, coll. J. Pérez 1915 (MP); 2 g (diss., fig. 37), Egypten,

Schmiedeknecht (Schmiedeknecht’s writing) coll. A. Weis, and 1 ©, Cairo, coll. A. Weis, M. armata

var. albovaria Er., det. Friese 1900 (SMF); 1 ©, Egypt, Ismailia, 8.iii.1965, K. V. Krombein, “hovering in

front of mud wall” (USNM); 2 ¢ 2 9, Aegypten, A. Andres, Wadi Rachid, 1.iii.1914, and Esbeth el

Nakleh (1 & 1 9, MNB), ii.1914, all M. albovaria Er., det. Alfken (SMF); 1 g, Unter Aegypten, Ad.

Andres, M. armata det. Friese 1911 (SMF); 4 ©, Egypte, Naville (MG); 1 g_ 2 9, Fayun,

Schmiedeknecht 1900, M. luctuosa 9, det. Kohl (NMW); 1 ©, Egitto, Gebel Asfar, 6.ii.1936, coll. A.

Mochi 8073 (MA); 1 © (pl, 1 fig. 2), Aegyptus, Cairo (print) (ML, ex MBUD); 1 9, Egypten, R.

Malaise (NRS); I 3, Egypten, Schmiedeknecht, coll. A. Weis (SMF); 1 &, Aegypt, Cairo, M. armata

var. mediterranea Gribodo (red, Gribodo’s writing?) (ZSM); | g, Egypt, Fayed, ii.1943, H. Priesner

(CP); 1 3 3 9, Egypte, Kerdasa, iii.1927, R. Mabrouk, El Maro, ii.1932, M. Aly, and Nawa, i.1913, leg.

Neguib (MNB); 1 3 2 9, Cairo, iii.1925, Min. Agric.(g), & with three labels: grandis armata, armata &

aegyptiaca, all det. Friese (MNB) 2 g, Egypt, Cairo, sub var. mediterranea Grib., ex coll. & det. Pittioni

(BM); | g, Cairo, sub. plurinotata F.-W. (sic) (BM). — Israel-Jordan: 1 ¢ (headless) 1 ©, Israel, C.

Jordan valley, Deganya A, 27.ii.1939 & 9.iii.1942, nos. 2890 & 6333, Y. Palmoni (AID); 1 ¢ 4 Q, Israel,

Jericho, Hisham Palace, 8.iii.1975, K. M. Guichard (BM); 1 &, Palestine, Haneger, Mt. Mishmaz, no

date, H. Bytinski-Salz (CBS); 1 3, Jerusalem, 12.ii.1942, H. Bytinski-Salz; 2 g 5 9, Palestine, Jericho,

23.11.1941 and 3—9.iv.1943, M. albovaria Er., det.?, M. armata grandis Lep. (1 39), det. G.

Mavromoustakis, and M. luctuosa var. (13 9), det. G. Mavromoustakis (CBS); 1 9, Palestine, Daphne

Oaks, 13.v.1940, H. Bytinski-Salz (CBS); 1 ©, id., Desanlah, 18.iii.1941, same coll. (CBS); 1 9 (diss.,

LIEFTINCK: Palaearctic Melecta 199

very small), Mt. Hermon, 1600 m, 9.vi.1975, M. Kaplan, and 1 9, same loc., 1900 m, 27.vii.1971, H.

Bytinski-Salz (CBS); 2 9, Israel, Yeroham Retama, 12.iii.1963, J. Wahrman (CBS, ML); 1 & 1 9,

Jerusalem, 12.11.1942, 9 with “24/Anthop. 9”, g with M. armata albovaria Er. = grandis Lep., det.

Mavromoustakis (CBS); 1 4, Israel, Bir-Rekhme, 15.iv.1953, 1 g, Gilgal, 1.iii.1973, M. Kaplan, | 4,

Ein Gadi, 19.iii.1975, A. Freidberg, and 1 9, Israel, Negev, Ze’elim 300 m, 18.iv.1967, leg. Kugler (all

CBS). — Lebanon: 1 Z, N. Libanon, Cedern bei Becharre, 1900 m, 3—6.vi.1931, Zerny (NMW). —

Syria: 1 9, Syria 1899, coll. A. Weis, M. ashabadensis det. Friese 1900 (SMF); 1 & 1, Syria, viii. 1899

(MNB); 1 9, N. Syrien, Sendschirli, 500 m, v. Lüschau (MNB); 2 g (diss.), Syria, Becker (NRS, ML); 1

Q, Syrie, coll. Ernest André 1914, var. grandis Lep., det.? (MP). — Cyprus: 1 9, Led[erer?] Cypern

1854, M. armata, det. Friese (NMW); 1 ©, Cyprus, Limassol, Yermasoyia, 17.iii.1979, L.-A. Janzon

(NRS). — Iran: 2 g (both diss.), Iran, 40 mi. S of Hamadan, 13.v.1960, R. v. d. Bosch (USNM, ML); 1

9, yellow disk16/40, Perse, Aucher 16—40 (MP); 1 &, Persien, Elburs, 800, S. Kuramahad, iv (MNB).

— USSR: | 2, Transcauk. (Armenia), Helenendorf 1886, M. armata, det. Kohl (NMW); 1 4, Caspian

Sea, Enseli, leg. (?) Mocquerys (USNM). — Turkey (Asia minor): 1 9, Kleinasien, Smyrna, Loew, with

“pseudoarmata”, det.? (MNB); several 3 9, Kleinasien, Sewdiköib nr. Smyrna, iv.1917, La Baume

(MNB); I &, Asia min., Tokat, M. armata var. mediterranea Grib., det.? (ZSM); 1 3, Asia min.

septentr., Alem Dagh, 600 m, 26—30.vi.1964 and 1 Z, Asia min. merid., Bulghar Dagh, 1200 m,

6—15.vi.1964, leg. Demelt (FAG); 1 g, Türkei, Bulga Haden, 12.vi.1965, leg. Rene (CMS); 1 9,

Turkiye, Konya, Sultan Dag, Cankurtaran Koy, 1900—2000 m, 19—20.vi.1967, leg. Reinig (SMF); 2 9,

Brussa 1882, M. armata var. grandis Lep. 9, det. Friese 1893 (MBUD); 1 9, Adana, Cilicia, 12.v.1911,

coll. J. Vachal (MP); 1 9, Turkey, above Hasanoglan nr. Ankara, 1500 m, 29.vi.1962, Guichard &

Harvey (BM); 1 ©, Turkey, Bebeh, iv.1960, M. Burr (BM); 1 4, Turkey, Rize, at sea level, 22.iv.1959,

K. M. Guichard (BM); 1 9, Araxesthal, Reitter 1890, M. armata var. grandis, det. Friese 1893 (NMW);

1 ©, Türkei, Urgiip, medio vi.1960, leg. Schläfle (CL); 1 4, Constantinople, coll. J. de Gaulle (MP). 6

3, 4 9, Türkei, Ankara, 3.vi.1972, Erzurum, Ispir, 17.vi.1973, Birecik/Urfa & Halfeti/Urfa,

16—17.iv.1976, and Östl. Türkei, Sirnak/Siirt, 3.vi.1977, all K. Warncke (CKW, ML); 1 3 2 9, Türkei,

Icel: Sertavul, 1400 m, 9.vi (3), Urfa: Halfeti, 29.v (9) & Nevsehir: Urgüp, 6.vi.1978 (9), all Max.

Schwarz (CMS). — Bulgaria: 2 ©, Arkutino, Black Sea coast, vi—vii.1970, K. Bleyl (CFP); 1 g, N.O.

Bulgarien, Pisanec, v.1955 (MNB). — Romania: 1 3, Mehadia, Mann 1859, M. luctuosa, det. Pohl

(NMW). — Greece: | 9, Morea, Cumani, Brenske (print) and 1 9, same loc., M. armata var. grandis

Lep., det. Friese 1893 (MBUD); 1 3, Greece, Athens, Theseion, 15.iv.1968, J. P. van Lith (CVL); 1 9,

Griechenland, Chalkis, Euböa, 12—16.v.1956, Fr. Borchmann (MKB); 1 9, Graecia, 189, coll. A.

Weis, M. ashabadensis det. Friese 1904 (SMF); 1 9, Graecia, Xiloxastron, 8.v.1962, W. Linsenmaier

(CL); 1 & (diss.), Peloponnesus, Chelmos, 2100 m, 1.vi.1962, M. Schwarz (CMS); 1 9, Kalamata,

14.v.1964, M. Schwarz (CMS); 2 9, Ellas, Athene, Mt. Imitos, 1.v.1963, S. Daan & V. van Laar (MA); 1

3 (diss.), Graecia, Delphi, 11.iv.1963 and 1 g, Agrinion, 17.iv.1963, Kl. Warncke (CKW); 1 g, Greece,

Salomis I., 6.iii.1932, E. E. Green (BM); series ¢9, Graecia, Olympia, Mykene & Lamia, iii.v. 1964,

Delphi, iv. 1966; Sparta, 9—13.iv.1969; all W. & E. Grünwaldt (CWG); Sikyon, Kastania & Trypi, iv-

v.1973, W. Grosz, coll. Grünwaldt (CWG); 1 &, Mt. Parnes, 600 m, 17.iv.1977, K. M. Guichard (BM); 1

4 9, Delphi, 11.iv.1963 (together with M. fulgida spec. nov.!), K. Warncke (CKW); 2 9, Chalkis,

Euboea, iv.1926, Holtz (MNB). Cyclades: 1 9, Kykladen, Pholegandros, v.1934, Werner & Wettstein

(NMW); 1 2 (diss.), Kiklades, Mikonis I., Lino, 3 km SE of town, 13.iv.1974, and Tigani, 5 km NE of

Ano Mero, 18.iv.1974 (3), A. C. & W. N. Ellis (MA); series 9, Mykonos I., iv.1927, and Ikaria I.

iv.1934 (MNB); 1 ©, Cykladen, Keos [Khios], v. Oertzen (MNB); Samos I., 1 ©, Samos, Kokkari,

25.iv.1977, H. Teunissen (CT); Kos I., 1 9, Kos, Asclepieion, 300 m, 30.iv.1971, M. A. Lieftinck (ML);

Rodos I.: 1 g, Rhodus, Hedenb. 1904 (NRS); 3 3, Rhodos, Fileremos & Archangelos, 29—30.iii.1977,

at Anchusa hybrida, A. Nilsson (DEU); 1 g, Rodi, 17.iv.1928, M. armata, det. Hedicke (IEB); 5 4,

Rodini, 22.iv.1970, H. Teunissen (CT, ML); 1 3 19 (g diss.), Rodos, Lindos, 26.iii & 17.iv.1970, A. C.

& W. N. Ellis (MA); 1 2, Rhodos, Fileremos, 23.iv.1976, H. Teunissen (CT); 1 g, Rhodos, Profitis

Ilias, 8.v.1975, H. Malicky (CG). — Crete: 1 4, Crete, nr. Kanea, D. M. A. Bate (BM); 1 ©, Kreta,

Omalos Ebene, 1000 m, vi.1942, Kl. Zimmermann (MNB); | 9, Ost-Kreta, Ssiwa, Ep. Pirgiotissa,

17.v.1925, A. Schulz, M. armata v. albovaria, det. Alfken (MNB); 2 ¢ 2 9, Creta, Canea and

Heracleion, iii.1906, Biro (MBUD, ML); 1 & 4 9, Crete, Chania, 15.v.1972 & 1.v.1974 (3 ©), and

Omaios, 1250 m, 13.v.1972 (1 g 1 9), K. M. Guichard (BM); 1 4, Kreta, Tybaki, iv.1914 and 1 9,

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Knossos, coll. F. Werner, M. armata var. albovaria, det. Alfken (NMW); 2 9, Kreta, Knossos,

13.v.1968, J. Gusenleitner (CG); 1 9, Kreta, Knossos, 13.v.1963, K. Kusdas (CMS); 1 9, Kreta,

Kaminaki, 900 m, 12.v.1971, Dr. Malicky, coll. K. Kusdas (CG); 2 & 3 9, Crete, Arkhanes, 28.iv.1972,

K. M. Guichard (BM); 1 9, Kreta, Lassithi, 850 m, 11.vi.1976, K. Warncke (CKW); 1 9, Kreta,

Neapolis, v—vi.1904, Rebel (NMW). — Ionian Is: 1 9. Ionian Is., S. Saunders (BM); 1 g 1 ©, Corfu,

Paganetti 1903 (MNB). — Yugoslavia: | g 2 9, Makedonien, Babuna Schlucht, 25,iv.1976, M. Kraus

(thoracic pubescence brownish: vers albifrons!) (CKW); 2 8, Nea, Agatoupolis, 27.iv.1976, on Thymus,

M. Kraus (CKW); 2 &, Dalmatia, coll. Simony and Mann, M. armata var. grandis, det. Friese 1893

(NMW); 1 ©, Dalmatia, C. Marzio Arbe, 4.vi.1914, M. luctuosa, det. Maid! (NMW); 1 9, Dalmatia

(MBUD); 1 3, Fiume [Rijeka] Mann 853, M. armata var. grandis, det. Friese 1893 (NMW); 2 ¢ (1

diss.), Dalmatien, one with M. armata det. Friese 1896 (MNB). — Italy: 2 9, Trieste, 8.v.1896, Ducke,

with two labels: armata & p. punctata, both det. Alfken (MNB).

N.B. — Some (not all!) of the last mentioned bees from Yugoslavia and Trieste,

across the Italian border, are moderate-sized and possess the usual regularly

arranged white abdominal spots of albovaria. The pubescence on head and thorax

is, however, not pure white but palest brownish grey, thus approaching more

northern individuals of typical albifrons. Further westward into the northern

provinces of Italy and the peninsula, these intermediates become much darker and

are true to the melanistic subspecies nigra Spinola (see under that taxon).

Except in a few places at the Côte d'Azur in the extreme SE corner of France,

purely white haired bees corresponding with typical albovaria, are well known also

from some mid and west European countries, but have mainly a southerly

distribution. In regard to body size, these individuals are as variable as

nominotypical albifrons of northern occurrence though possessing, as a rule, larger

and more regularly placed abdominal spots. Material examined from the above

countries, arranged more or less from E to W, can be listed from the following

localities.

Switzerland: 1 9, Wallis, Sitten, 19.vi.1914, Th. Steck (NMB). — France (N to S): 2 9, St. Cyrs s.

Mer (S. et M.?), 14.vi.1953 (CVZ); 1 &, Argentlat?] (Corrèze), 20.v.1887, coll. J. Vachal 1911, M.

luctuosa 9, det.? (MP); 1 &, Dieulefit (Drôme), 22.vi. 1960, J. v. d. Vecht (ML); 1 & (diss.),

Montmeyran (Drôme), 15.iv.1979, J. Teunissen (CT); several 3 9, Carpentras (Vaucluse), v.1952 &

v.1953, P. M. F. Verhoeff, and 4 9, same loc., 30.v.1959, W. Linsenmaier (CL); series ¢ (diss.) 9,

Digne (Bses Alpes), ult v.1957, M. A. Lieftinck (ML); 1 9, Montauroux (Var), 30.vi.1960, J. v. d. Vecht

(ML); 1 &, Fayence (Var), 19—23.iv.1973, G. Barendrecht (MA); 1 &, Callian (Var), 21.v.1963, W.

Linsenmaier (CL); 1 3, Beau Vallon, 12.iv.1967, O. W. Richards (BM); 2 9, Gall./Sichel, M. armata

var., det.? (NRS); 1 &, Montpellier (Her.), D. Lichtenstein 216/11, M. grandis Lep., det. J. Pérez

(MCG) and 1 9, same loc., Lichtenstein (MNB); series 3 9 ( fig. 7), same loc., Jardin botanique,

13.iv.1961, M. A. Lieftinck, “together with Anthophora acervorum L.” (ML); 1 9, Vallon de Connuls,

iv.1893 and 1 9, Nimes (Gard) (BM); 1 9, Toulon (MNB); 2 9, Pont du Gard (Gard), v.1952, exped.

H. Engel (MA); 1 ©, St. Jean de Luz (B* Pyrén.), Le Lac, 2.v.1946, coll. V. Muspratt (BM). — Spain:

1 9, N Spain, Huesca, Boltona (Aragon), 17.v.1953, I. H. H. Yarrow (BM); 1 9, NW Spain, Pontevedra

above Vigo (Léon), 200 m, 3.iv.1964, K. M. Guichard (BM); 1 &, Ponferrada (Léon), 25.iv.1955, I. H.

H. Yarrow (BM); 1 9, N Spain, Santander, Santillana, 4.v.1958, excurs. ML (ML); 1 9, Adradas (prov.

Soria, Castilia), 9.vi.1961, H,, excurs. ML (ML); 1 &, Burgos (Castilia), 1.v.1955, I. H. H. Yarrow

(BM); 2 3, Hispania, Albaracin (Catalon), vi.1953, leg. Fabian (CMS); 1 g, violet disk, Barcelone,

coll. J. Perez 1915 (ex MP, ML); 2 g, Canet de Mar (Catalon), 25.iii.1960 & 5.iv.1962, Feo. Vergés

(CMS); 3 & 3 9, Barcelona, Schmiedeknecht, coll. A. Weis (SMF, ML); 1 9, Gerona (Catalon),

Blanes, 5.iv.1956, and 2 &, Barcelona, Vich & Montisquiu, 18.iv.1956, I. H. H. Yarrow (BM); 1 g

(diss.), Sierra Morena, Santa Elena (Jaen), 4—8.iv.1926, H. Lindberg (MH); 1 9, Ribas (Catalon),

Busmet, coll. J. Pérez 1915 (MP); I 9, Spanien, prov. Taragona, Flix, F. Haas 1914—17, M. albovaria

LIEFTINCK: Palaearctic Melecta 201

Er., det. Alfken 1925 (SMF); 1 9, E Spain, Tarragona, 7 km E. 28.v.1960, excurs. ML (ML); 1 9, W

Spain, Abadia, 700 m, 5.v.1960, excurs. ML (ML); 3 9, Caceres (Estremadura), Tornavacas, 1200 m,

V83, and Jerte, 900 m, 22.vi.1961, at Echium, excurs. ML (ML); 1 &, Monchique, 7.iii.1906/Madrid. G.

Schramm (MP); 1 9, prov. de Madrid, Meco, 610 m, J. Alvarez (CCS); 1 9, Aranjuez (Madrid),

25.v.1959, W. Linsenmaier (CL); 3 4, Espagne, Uclés (MCG); 1 &, Waltl, Spani 1830, M. armata var.

grandis Lep., det. H. Friese 1893 (NMW); I &, Granada, Staudinger (BM); 1 4, Valencia, Betera,

Quilis, M. albovaria, det. Alfken (MNB); 1 3, Hisp., Godelleta Valencia, Peres Torres, coll. Peres, M.

Fig. 30. M. albifrons nigra, sternites 7 and 8 (3 Rome, Italy). — Figs. 31—33, M. albifrons albovaria; 31,

dorsal (upper) and ventral view of tergite 7 (& Egypt); 32, partial ventral and dorsal view of genital cap-

sule (same specimen, bristles partly omitted); 33, lateral aspect of gonostylus showing dorsobasal and

ventrobasal processes (same specimen). Scale lines 1 mm (fig. 30) and 2 mm (fig. 32)

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

luctuosa Scop., det.? (CCS); 2 9, Hispania, Valencia, Cafiada, Giner Mari (CCS, ML); 3 9, Andalusia,

Sevilla, 14.v.1972, W. Grosz (CWG, ML); series g, Elche (Alicante), v.1933, Hering (MNB); 2 g,

Südspanien, Granada, 30.iii.1959, W. Schlafle (CL); 1 g, Algeciras, 25 iv, coll. O. Vogt (MA); 2 g,

Andalusia, Algeciras, 1—10.v.1925, and Tarifa, 13.v.1925, Zerny (NMW); 1 g, Espana, Cadiz, M.

López Banus (CCS); 1 &, Porto Sta Maria nr. Cadiz, 17.iii.1906, coll. Schramm, coll. J. de Gaulle 1919

(MP). — Portugal: 1 9, Portugal, Wattison coll. (BM); 1 g, Lisboa, 3.iv.1955, N. F. d’Andrade (ML);

1g 1 9, Rezende, 14.iv.1950 (together with 1 & M. luctuosa Scop.!), N. F. d'Andrade (ML); 2 9,

Portugal, Algarve, Praja da Rocha, v.1934, H. Main (BM).

Varicoloured specimens intermediate between typical albifrons and albovaria,

with head and thorax pubescence distinctly brownish and with conspicuous pure

white abdominal spots, are occasionally met with in mixed populations of the

latter, comprising all intergradations. These were found in the following localities:

Austria: 1 3, almost all white, Ob.-Österreich, Traunsteingeb., 7.vi.1951, R. Loberbauer (CMS). —

France: 4 large-sized ¢ (diss. fig. 71), Antibes sur Colza (Alpes-Mar.), 24.iv.1969, J. Riom, head &

thorax definitely brown, abdominal spots small, white (INRA); | large 9, vers albifrons typ., with small

white-spotted tergites 2—3, Bordeaux (Gironde), 28.iii.1974, H. Hamann (CMS). — Spain: 2 9, Soria,

St. Maria de Huerta (prov. Soria, Castilia), 13.vii (sic) 1969, P. M. F. Verhoeff, thorax pubescence

mixed with brown (CV); 3 3, Ronda (Catalon) and Canet de Mar nr. Barcelona, thorax pubescence

whitish fading to grey-brown (CVS); 6 &, small-sized (10—11.5 mm!), Pozuelo de Alarcon nr. Madrid,

R. Ruano, 3.iv.1959, one with distinctly grey-brown thoracic pubescence, abdominal spots of large size

(CCS, ML); large series 4 9, body measuring 9.5—17.5 mm, all profusely spotted but partly with head

and thorax pubescence definitely brown, env. Valladolid (Castilia), nr. Fuensaldana, Los Alamos &

Villabafiez, v-vii and xi (sic) 1974, at nest entrances of Anthophora acervorum (L.), E. Asensio de la Sierra

(CAV, ML); 2 ¢ 1 9, various sizes (see above), Toro (prov. Zamora), ca. 30 km from Zamora,

25.iv.1974 & i-ii.1975, in nests and at nest entrances of Anthophora acervorum (L.), crinipes Smith (ii.

1975) and fulvitarsis Brullé, E. Asensio de la Sierra (CAV, ML).

Interesting information concerning the host relations of the present taxon in

southern France was found (sub armata) in the manuscript catalogue of J. Pérez’

bee collection in the Paris museum, made available to me now already 20 years

ago. Corresponding remarks are contained in the author’s more elaborate account

of Mediterranean Melecta (Pérez, 1883: 302—307). At present we know for certain

that several white-spotted ““near-alike” species may occur together all over the

Mediterranean region inclusive of the southern provinces of France. Therefore it

must remain uncertain whether all forms discussed by Pérez were really albovaria.

Yet I think the author’s observations are of some historical interest and worth

quotation in his own language:—

“No. 156 — Melecta armata Lep. primitivement marqué punctata puis rayé.

Périgueux, Tarbes, Bordeaux, très commune, en avril et encore en mai sur un

grand nombre de fleurs, en particulier celles du chou, dans les jardins comme dans

les champs. Parasite des Anthophora parietina |= plagiata (Illiger)] et personata (=

fulvitarsis Brullé); aussi la voit-on fréquemment sur les talus où nichent ces

apiaires. Grandes variétés de taille et de vestiture. Les sujets qui éclosent des

cellules de personata sont plus grands, plus fourrés de poils; ceux qui sont parasites

de la parietina sont plus petits, leur tête notamment est peu volumineuse; leurs

poils sont moins longs et moins épais. Les taches des segments 3, 4, 5 peuvent

disparaitre complètement; d’autres fois elles sont très larges particulièrement chez

quelques 2. Le corselet est généralement chez les ¢ couvert de poils fauves en

LIEFTINCK: Palaearctic Melecta 203

dessus, sauf sur l’écusson et le post-écusson qui les ont noirs, ainsi que le voisinage

de l’insertion des ailes. Chez la 9 cette variation est assez rare; généralement le

noir gagne davantage sur le corselet et ordinairement on distingue quelques

houppes de poils noirs dont 2 plus distinctes en avant. Face entierement noire chez

une 9. — 9, premiers jours de juin, talus habités par les Anthoph. personata,

parietina, aestivalis, crinipes, volant en ce moment.”

From the above notes on the colour design, it is quite clear that, apart from

typical albifrons and albovaria, specimens of nigra Spinola, from maritime districts,

were also at the writer’s disposal; further details are unfortunately not known.

During recent years an investigation of the host-parasite relationships was

started with more precision by the Spanish hymenopterologist Dr. Enrique

Asensio, in Castilia and Zamora (north-central Spain). During a period of five

years (1971—1975), the nesting sites of three Anthophora species were regularly

inspected, viz. those of acervorum (F.), crinipes F. Smith and fulvitarsis Brulle.

These bees are among the first members to emerge in the spring (mid April) and

overwinter as adults. They formed mixed colonies, the nesting sites being

preferably in tali of deep and little frequented roads in three or four farming areas

just outside Valladolid or at some distance to the W of that town. The contents of

many nests were excavated, the adults reared and all stages of development

preserved for further study. All species were associated with the cleptoparasitic

M. a. albovaria, i.e. the only Melecta observed in the nesting area. The big host bee

A. fulvitarsis was first noticed in May 1971 visiting the racemes of vine in company

with albovaria at a farm near town. Already in July 1974 adults of albovaria were

present in A. acervorum cells near Toro, some 30 km E of Zamora, and also in

November of the same year at a nesting site N of Valladolid. During April 1974 and

again by the end of January 1975, while digging out the contents of A. crinipes

nests, also near Toro, a number of surprisingly small adult males of albovaria were

unearthed resting in cells (or burrows?) of this equally diminutive Anthophora. Nest

entrances of A. fulvitarsis and acervorum were close by, and Dr. Asensio noticed

that sometimes females of these two species used the same entrance hole, the

parasitic intruder being usually also present.

The interesting point here is the striking difference in stature and size exhibited

by individuals of a single species of Melecta within a limited area. Nests in small

mixed colonies of at least three Anthophora species were all robbed by albovaria

alone. As these host bees are not at all closely interrelated, differing markedly inter

se in structure and dimensions, they will probably have their own foraging

capacity, each provisioning its cells with a specific quantity of pollen and nectar.

Therefore the outsized individuals of albovaria may well have emerged from the

well-stored cells of A. fulvitarsis, while the smallest would have reached maturity

by consuming the less copious food supply available in the cells of the little

A. crinipes. Two other species of Melecta, viz. italica Radoszk. and luctuosa (Scop.)

were also found in Spain by Dr. Asensio. Perfectly fresh, small-sized males of

italica were collected in May 1972 and April 1973, in a different area near

Valladolid, flying in company with A. cf. dispar Lep.; females of luctuosa were

caught only at Salamanca.

204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

This predominantly white subspecies of albifrons is one of the most widely

distributed and undoubtedly the commonest Melecta of the entire Palaearctic

region. Both sexes of the three subspecies are sufficiently characterized in the key

to the males. Once again it must be emphasized that several local populations of all

forms are rather puzzling with regard to the colour of the pubescent pattern and

extent of white abdominal spots: these aberrant individuals are briefly discussed

under the subspecies concerned.

M. a. albovaria occurs discontinuously throughout southern Europe, i.e. with the

exception of practically the whole of Italy, the Tyrrhenian islands, and Malta, in

which area it is replaced by the next subspecies, a. nigra. Its further range

comprises North Africa north of the Sahara and eastward as far as the Caspian Sea

and into Iran.

Melecta albifrons nigra Spinola

(figs. 26—30, 73)

Melecta nigra Spinola, 1806, Ins. ligur. spec. nov. 1: 44, tab. I, fig. XIV (Habitat in Liguria, haud infre-

quens). — Friese, 1895, Bienen Europa’s: 159—160 (not seen!). — Alfken, 1937, Boll. Ist. Ent. Bo-

logna 9: 111 (9 & Ronzano: 9 g Bozen; ¢ Calabrien: Antonomina; notes: “Wirt vermutlich Antho-

phora crinipes F. Smith”). — Lieftinck, 1958, Comment. Biol. Helsingfors 18.5: 30 (note on type).

Melecta fasciculata Spinola, 1806, Ins. ligur. spec. nov. 1: 45, tab. 1, fig. XV (Habitat propè Genuam, ra-

rior). — Friese, 1895, Bienen Europa’s: 160 (not seen!). — Lieftinck, 1958, Comment. Biol. Helsing-

fors 18.5: 20—22 (synon. notes), pl. 1 fig. 11 (3 struct., lectotype M. fasciculata Spin., Genova, = ni-

gra Spin.).

Crocisa atra Jurine, 1807, Nouv. méth. Hym. & Dipt. 1: 241, Taf. 12 Gen. 34 (9 sine patria (= Europe!).

— Frey-Gessner et al., 1882, Mitt. Schweiz. Ent. Ges. 6: 387—397 (remarks on Jurine’s type). —

Lieftinck, 1958, Nova Guinea, new ser. 9: 24 (list, rect. = Melecta). Syn. nov.

Melecta punctata var. Spinola, 1809, Ins. ligur. (edit. Francofurti): 153 (diagn. variations).

Melecta bipunctata Lepeletier, 1841, Hist. nat. Ins. Hym. 2: 446 (9 Italie, Toscane). Syn. nov.

Melecta aterrima Lepeletier, 1841, Hist. nat. Ins. Hym. 2: 447 (Q Ile de Noirmoutiers, Dejean). Syn. nov.

Melecta calabrina Radoszkowski, 1876, Horae Soc. Ent. Ross. 12: 95—96 (3 — not 9! — Calabria). Ra-

doszkowski, 1893, Bull. Soc. Imp. Nat. Moscou n.s. 7 (2—3): 183, fig. 38 a—c, i (Z genit.). Syn. nov.

Melecta armata cum subspec., Pérez, 1883, Actes Soc. Linn. Bordeauz 37 (ser. 4, vol. 7!: 302—307 (pars:

compar. notes).

Melecta luctuosa Scop. var. nigra Spin., Friese, 1893, Bienenfauna Deutschl. u. Ungarn: 37 (Bozen). —

Pérez, 1890, Actes Soc. Linn. Bordeaux 44: 36 (cat. France, sine locs.).

Melecta armata Panz. var. aterrima Lep., Frey-Gessner, 1907: 144 (Locarno & Geneva, note).

Type and syntypic material. — Italy: 1 © (not ¢!), over drawer label ““Melecta

notata var. J/tota nigra/Liguria” (white). Lectotype M. nigra Spin. by present

designation (MT). — 1 & (diss., figs. 26—29), over drawer label ‘‘Melecta

fasciculata m. Ins. ligust. @... notata var.? Gênes” (white). Lectotype

M. fasciculata Spin. by earlier designation (MT). — 1 4, labelled “Italia Genua

Spinola S.” (modern writing), “Type” (print on red), and old black-rimmed label

“M. fasciculata Spin. mas S.pr. punctatae/Genua Spinola” (probably Spinola’s

writing). Syntype M. fasciculata Spin. (MNB). — N. Italy or S. Switzerland: 1 9,

originally mounted upside down at strong pin; pictured in Jurine, loc. cit., with

white written label “rufipes x”, sine loc. Holotype Crocisa atra Jurine (MG). —

Italy: 1 & (diss.) labelled ““Calabr.” (white, Radoszkowski’s hand), “Type” (print,

LIEFTINCK: Palaearctic Melecta 205

red), ‘‘calabrina” (pencil), “Melecta calabrina Radoszk. Type” (written, Dr.

Enderlein det.). Holotype M. calabrina Radoszk. (MNB). — Other syntypes. Italy:

1 & 4 2 (syntypes nigra and fasciculata): 1 3 1 9, g with pin-label M. punctata?

var. fasciculata Spin. Ped. 39; 1 9, over drawer label M. notata Kl. 39...

punctata Lep. Gênes; | 9, with pin label M. punctata? var. F. Lep. 9 Torino; 1 9

over same drawer label as lectotype M. nigra (all MT).

Further material. — Italy: Liguria & Piemonte: 17 & 18 2 in coll. Fea, Magretti and Gribodo,

from env. of Genoa, Borzoli, 10—20.iv.1909 and iv.1883, Orte Botanico, Caneva 1883, C° Amata,

Paderno Dugnano, spring 1895— 1908; Piemonte, Genola, v.1885 and Coazze, leg. Gribodo (all MCG);

1 ©, Liguria, Mortola-Inferiore, Giardino Hanbury, 28.iv.1975, H. Wiering (MA). Liguria: 1 & 1 9,

Genua, Lavagna, 26.v.1970, B. J. Lempke (MA); 1 4, Italia, Spezia, 10.iv.1906, Dr. Uzel (NMW); 1 9,

Bordighera, iv.1961, W. Grünwaldt (CVS). Lombardia: 1 ©, Monza, R. Parco, III.1932, D.

Prestifilippe (IEP). Alto-Adige-Trentino: 1 & 1 ©, Italia, Sanertal/728, © with M. testaceipes, det.?

(MNB); 11 3 1 9, iv-v, various years, | © M. armata aterrima Lep., det. Friese (MZB); series ¢ 9, Süd-

tirol & Bozen, variously labelled M. nigra + fasciculata, aterrima, det.? (MNB); 1 9, Bozen, Tirol,

25.iv.1898, Anchusa, leg. Friese, M. nigra det. Alfken (MNB); 1 g, Bozen, 2.iv.1897, M. armata var.

bipunctata Lep., et. Friese (MCG); 3 ¢ 2 ©, Bozen 1886, F. Kohl, M. armata var. bipunctata, det. Friese

1893 (NMW); large series 9, Südtirol, Bozen, at Saponaria ocymoides, Stöcklein leg. (ZSM); series 14 ¢

9 ©, Bozen, various dates, A. Weis (SMF) & 1 g, id., M. armata var. bipunctata Lep., det. Friese 1893

(MBUD); 2 2, S. Tirol, Weidbruck, 15—16.v.1960, H. Priesner (CMS) Emilia-Romagna: 1 &,

Modena, Casinalbo, iv.1943, G. Fiori (MA); 1 g 1 9, Bologna, Ronzano, M. nigra and fasciculata, both

det. Alfken (MNB); series 24 ¢ 19 9, env. of Bologna, Ronzano, Gaibolo, iv-v, various years, leg.

Grandi et al, M. nigra M. Spin. fasciculata M. Spin., det. Alfken 1937 (1 ©), M. nigra Spin., det. Alfken

1937 (4 ©), M. fasciculata Spin., det. Alfken 1937 (21 3), M. armata, det. Hedicke (1 g), armata f.

bipunctata Lep., det. Hedicke 1921 (2 3), and armata f. nigra Lep., det. Hedicke (3 9) (all IEB), and 6

9, Bologna, Gaibolo, v.1953, leg. Grandi (MA); 1 ©, Bologna, Prj. Fiori (MCG); 1 9 (totally black!),

Castelvetro, 18.viii. (sic) 1885, coll. Baldini (MT) and | &, same loc., 7.x. (sic) 1885, same coll. (MT).

Toscana: 1 g, Florence, coll. O. Sichel 1867 (MP); 1 9, Fiesole nr. Firenze, 8.v.1925, at Salvia, and 1

©, Siena, 16.v.1925, M. A. Lieftinck (ML); 1 9, Florence, Picioli, coll. Tournier (MG); 1 & 2 9,

Castiglioncello (Livorno), 3—11.v.1958, G. Barendrecht (MA); 11 49, Pisa, iv-v.1953 (CTP). Marche:

23 1 9, Macerata, 17—30.iv.1939, leg. G. Soika (CVZ). Abruzzi: 1 9, Cerchio, Aquila (MCG); 4 ¢ 1

©, E-coast, Pescara, 21.iv.1963, W. Grünwaldt (CKW). Lazio: 1 &, Marino, iv.1933, and 1 9,

Caffarella, iv.1935, (ZSM); 1 g (diss, fig. 73), Roma, Aquatraversa, 13.iv.1949, M. Comba (CMC); 5 &

6 9, env. of Roma, Bosco Sacao, Subiaco (S. Sco. Castica), Marino, Acilia, Priverno, Formia, Capo

Circeo, and Sabaudia, iii-v.1932—1945, leg. Castellani et al. (all INER). Campania: | 9, Neap., “nigra

et aterrima Lep.” and “punctata Grib. nigra Lep.” (old writing) (MNB); 1 9, Naple, coll. O. Sichel 1867,

M. nigra (unknown writing, MP); 1 9, Neapel, M. testaceipes, det. Friese (MNB); 1 &, Roy de Naples,

O. Costa, 1.53 (MP); 17 ¢ 9, Portici, various spring data (IEP); 2 & 2 9, Pisciotta, 6—10.v.1936,

M. armata aterrima & a. nigra, det. Alfken (IEP); 1 9 (totally black), Monteforte, Intrino, 12.iv.1918

(IEP); 1 3, Areve, 2.v.1930 and 1 g, S. Vito de Norm, v.1905 (IEP); 1 3, Neapel, Coll. Rhd./28985,

bipunctata (old writing) and 1 9, Neapel/728, M. arm. var. aterrima, det. Friese 1896 (MNB); 1 9,

Portici (MCG). Puglia: 2 3, Monte Argentario & Monte Gargano, 12—14.iv.1975, H. Teunissen (CT,

ML); 1 9, Manfredonia 1904, Paganetti(NMW); | 3, Barletta, v.1893 (MT); 1 g, Apulien, Spongano,

18.iv—3.v.1933, v. Loudon leg. (MNB); 1 9, Apulien, same loc., 27.ii—16.iii.1933, same coll.

M. luctuosa, det.? (ML). Basilicata: 33 & 11 ©, Rionero, iv—vi.1890—93, coll. Baldini (MT); I g,

Monticchio, 15.iv.1890 (MT); 8 g 1 ©, Vulture, all vii. (sic) 1891 (MT); 1 9, Rionero, 1.v.1891,

M. armata, det.? (INER); 1 9, over old drawer label “M. nigra h. in Italia Scop.” (ex Scopoli?) (MP). 2

g, sine loc., armata, det. Saunders (MBUD). Calabria: 1 ¢ 1 9, Calabria, Antonimina, Paganetti,

M. nigra, det. Alfken (MNB). — Corsica: 2 g, Corsica, Klinckow (NRS); 2 g, Corse, Mann 1855,

armata, det. Kohl (very large ex.), and armata var. bipunctata Lep., det. Friese 1893 (smaller ex.)

(NMW). — Sardinia: 1 9, Sardinia (print, MBUD); 2 g, Sardinia, Oristano, Kraus leg. (AMNH); 1 9,

“Sardin. Kutit.” (?), testaceipes, det.? (MNB). — Sicilia: 1 9, Sicile (lilac disk), coll. J. Pérez 1915

(MP); 1 g, Taormina, V. narbon. mm 20, 20.iv.1937, Kuntzen leg. (MNB); 1 9, Sicilia orient.,

206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Taormina, 200 m, dint.22.iv.1950, Hartig & Grshm. leg. (INER); 1 4, Sicilia, Taormina, 23.iv.1976, J.

A. W. Lucas; 2 4, Sicilia 1885, M. armata, det. Mocsary (MBUD); 1 9, Sicilia, M. Cuccio, 19.v.1953,

A. Fiori (MA); 1 g, Sicilia (MCG); 1 4, Sicilia, Taormina, 22—30.iv.1921, Zerny, M. armata, det.

Maidl (NMW); 2 9, Sicilia, Palermo, 10—28.v.1921, Zerny, M. armata var. nigra Lep., det. Maidl

(NMW); 2 ¢ 2 9, Sicilia, 1858 Mann, M. luctuosa det. Pohl (1 g), armata var. grandis Lep., det. Friese

1893 (1 3), and armata var. nigra Lep., det. Friese 1893 (2 9) (NMW); 2 g, Palermo, 30.iii.1965, M.

Schwarz (CMS); 2 9, Sicilia, prov. Palermo, Bosco di Ficuzza, 16.v.1965 and S. Martino d.Sc.,

1—12.vi.1954, J. Klimesch (CMS); 2 9, above Trapani, Sicily, 200 m, 16.iv.1965, K. M. Guichard

(BM); 1 g (diss.) 1 9, Syracus, iv.1898, $ with punctata var. albovaria, det.? and 9 with M. luctuosa var.

ruthenica, det. Friese (MNB); 2 &, Sicilia, Monte Gargano, 12—14.iv.1975, H. Teunissen (CT, ML).

Synonymy. — Here follow a few remarks on a number of types synonymized

with the present subspecies.

The holotype of Crocisa atra Jurine possibly originates from northwestern Italy,

or even from south Switzerland, and does not differ from certain examples

occurring in the former country. The mandibles are plainly exposed and of the

shape characteristic for this and other species of the genus. The specimen lacks

two legs (fore and hind), which for the rest are wholly black. The abdomen carries

distinct greyish spots, one each side, on tergites 2—4.

Of the five species named by Lepeletier, the localities of two obscurely coloured

bees, nigripennis and testaceipes, are unknown and since the types are no more in

existence, their names should be rejected (see species incertae sedis, at the end of

this work). M. nigra Lep., — the name given to a melanistic female from Genoa —

is homonymous with Spinola’s bee and undoubtedly belongs to the same

subspecies. The type of a fourth species, bipunctata, a female reportedly from

Tuscany, could not be recovered in any collection, but since the brief diagnosis

applies perfectly to specimens of nigra collected by myself and others in the same

province (Toscana), should also be considered synonymous with the present

subspecies. Lastly, the type of aterrima Lep., described from Ile de Noirmoutier(s),

at the Atlantic coast of Bretagne, has also become lost but could not have been

anything else but a melanistic individual of what is here regarded as nigra Spinola

(see below).

The type male of M. calabrina Radoszkowski is a true member of the albifrons

group, its structure agreeing with south Italian specimens of nigra. The mid tibiae

are predominantly white, the hinder pair only bear a small basal spot. Tergite 1 of

abdomen black with greyish hair tuft on either side, those on 2 being larger, while

the sides of 3—4 are each marked with a single much smaller white spot placed

more inward. The genital organs were dissected out and glued on a card by its

describer.

Specimens like this should not be confounded with other black-coloured

Melectae occurring in Italy and elsewhere in the Mediterranean, like italica and

leucorhyncha taormina, which are closely similar superficially. Though often

occurring together with the latter in one locality, nigra can be immediately

distinguished from these by the characteristic shape of the tarsal claws, its wing

venation, and the greater complexity of the male copulatory apparatus.

Variability. — In the past the black-haired Melecta occurring in the

Mediterranean region and further eastward have given rise to much confusion,

LIEFTINCK: Palaearctic Melecta 207

Figs. 34—37. M. albifrons albovaria; 34, ventral view of tergite 7 (3 lectotype Algeria); 35, sternites 7

and 8 (same specimen); 36, sternite 8 (4 Tunisia); 37, apices of sternite 8 (two 3 from Egypt, showing

variation). Scale line 1 mm (figs. 35—36)

taxonomically as well as, inevitably, in respect of their nomenclature. Feelings of

apprehension regarding the proper names for these bees were plainly revealed by

Friese (1893, 1896) and Alfken (1937), who assigned no less that seven names to

the colour variations of the present subspecies of albifrons. This nigra has its

distribution centre in Italy and the Tyrrhenian islands. The striking variation in the

extent of grey and pure white hair covering the body and legs of Italian

populations, is best demonstrated by a large series of 58 ex. (44 & and 14 Q) in

coll. Baldini (MT, females labelled var. aterrima by Zavattari), collected in the

provinces of Emilia and Basilicata, and by 31 ex. (15 3 and 16 ©) from Lombardia

and the environs of Genoa, in coll. Gribodo and Magretti (MCG). Though not all

of these are still at hand, they were studied by me in 1959 and formed the basis of

the key characters. All males have white faces, spotted legs, and at least much grey

hair upon the mesonotum, but the white spots on the tergites 3—4 are present only

in about 50 per cent of the total, in all others they are restricted to the first two.

Females are considerably darker, either entirely black, or marked only with small

white abdominal spots.

It will be noted that specimens from Sicily, here all listed under albifrons nigra,

208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

are not at all homogeneous either. Yet both sexes differ from each other only in

the distribution of white and black hair. g: White plus variants differ from

mainland populations by having a complete row of spots on tergites 2—5, the one

on 2 being largest and comma-shaped. 9: Head, thorax and legs completely black,

save for a vestigial spot at base of hind tibia, the abdomen with complete row, as in

gd, except that all spots are a little smaller (from Taormina, Palermo and

Syracuse). — g': Minus variants only have the two basal tergites white-spotted, the

mark on 2 being small, hence are similar to many mainland examples of that sex

(from Mts. Gargano and Argentario); 9 not available.

Melanistic females inseparable from typical nigra occurring in Italy, have been

examined from the following countries (see also under albovaria).

France: | ©, Alpes marit., Castella sur Menton, 1.xii.1954—1.v.1955, J. Briedé (MA); and from

more northern and coastal localities in France: 2 9, Ouessant (Finistére), 31.v—3.vi.1960, G.

Kruseman et al. (MA); 1 9, Bar. s. Seine, M. aterrima Lep./aterrima (print), coll. Gribodo (MCG); 1 9,

Passy, Reg. Paris, coll. O. Sichel 1867, with old separate labels “Pass.” and “M. aterrima”, in Sichel’s

hand (MP); 1 9, St. Nazaire, 25.v.1911, coll. J. de Gaulle 1919, ‘M. luctuosa var. noire inédite” (MP); 1

Q, Loire Atl., Nantes, Dominique, coll. Tournier (MG); 1 9, Nantes, Coll. J. Perez 1915 (MP); 1 9,

Bretagne, Roscoff, vi.1957, J. Pasteels (ML); 1 9, dark blue disk, coll. J. Pérez 1915 (MP); 1 9, Loire

inf. (print), coll. J. Pérez 1915 (MP); 1 9, Gironde, Vivonne, près Bordeaux, coll. J. Pérez 1915 (MP).

The Atlantic populations found at the French coast and those listed occurring

more scatteredly in the northern parts of that country, are strikingly similar to

some specimens from the Channel islands and southern England. This is the form

named M. aterrima Lep., described from Ile de Noirmoutiers, an island off the

French westcoast. In the Brit. Mus. (Nat. Hist.) and the University Museum at

Oxford (OUM), I have seen equally dark and occasionally almost entirely black

specimens from various counties in Southern England. These are not now before

me but I noted the following localities: Colchester (Essex), Oxford, New Forest

(Hampshire), Wimborne (24.v.1953), Sandown (Isle of Wight), Bude (Cornwall),

and Guernsey I. (Channel Is.). A male from Chouet, Guernsey, 18—23.v.1960,

taken by Mr. I. H. H. Yarrow (BM), bears the collector’s label “indistinguishable

from Italian nigra”, a statement with which I concur. On the other hand, I have

before me a male from Jersey I. (also of the Channel Is.), 8.v—6.vi.1965, J. Briedé

(MA), which I am unable to distinguish from typical albifrons Forster! The same

applies to males from near the Atlantic coast in western France and the Channel

Islands; in them the thorax pubescence is distinctly brown while the abdominal

spots are relatively of small size, e.g. a male from Finisterre, Ile de Batz,

11.vi.1958, J. H. van Bree (MA).

Apart from the Atlantic forms so closely approaching nominotypical nigra,

some remarkable varieties also make their appearance toward the east and south,

in the marginal parts of its distributional area. Notably two puzzling females

should still be mentioned, one from Poland, the other from eastern Turkey. Both

are indistinguishable from some white-spotted Sicilian examples of the same sex;

yet, on account of their unusual origin and the absence of males, I am referring

them to a. nigra with some misgivings. Poland: 1 9 (diss.), Tal von Otuzy [ =

Otusch or Otush, 25 km WSW of Posen, teste Dr. Fischer in litt.], 22.iv.1923, W.

LIEFTINCK: Palaearctic Melecta 209

Wuczeticz leg. (NMW); and Turkey (Asia minor): 1 © (diss.), labelled Taurien,

Magaratsch (print) [Kurdistan, Armenian Taurus, Marasch?], M. luctuosa, det.

Kokujew (CO).

Pronounced melanism is a well known phemomenon amongst other melectini

occurring in Italy and islands round the Tyrrhenian Sea (see also M. italica,

leucorhyncha, and members of the allied genus Eupavlovskia). I see no profit in

guessing at the factors effecting the localized and discontinuous distribution of

a. nigra and its outlying forms. However, the facts speak in favour of climatic

influences responsible for the general appearance of these obscurely coloured

bees, — at least in so far as the analogous Atlantic populations are concerned. To

sum up, I still believe that the melanistic nigra can be most conveniently

maintained as a fairly defined and recognizable geographical subspecies centred in

Italy and the big Tyrrhenian islands.

2. Group of Melecta duodecimmaculata (Rossi)

It will be understood from our illustrations that the species group here treated,

so well characterized by the presence of paired lateral abdominal spots, genuinely

only forms a “subgroup” of M. albifrons (Forster). The morphology of this alliance

considered as a whole, is quite homogeneous, all members having the same ‘facies’

and show great uniformity of structure. This similarity of characters, expressed

also in the wing venation, is undeniable and demonstrates their intimate relation.

On the other hand, there is unlimited variety in the pubescent colour pattern,

which is very confusing. Whereas albifrons only breaks up in a number of fairly

defined geographical subspecies, each showing a more or less characteristic

design, the diversity within the present cluster is more complicated and less easily

understood. Apart from the extent and scheme of coloured markings, its members

exhibit slight but apparently constant differences in the armature of the legs, wing

colour, nature of pubescence generally, and length of hair. Yet the valuation of

these mixed characters must remain subjective and is difficult to reconcile with the

distribution of the various components of the group.

In the key for the males not much value has been assigned to the form of the

apical sternites and appendages of the gonocoxite. Of these structures, many more

than those figured were dissected out and compared, — without much advantage,

however. Differences in proportion and outline of the sternal plates sometimes

appeared to be incidental and of little significance. For that matter, a comparison

of my sketches reveals the same inconstancy throughout the albifrons group (and

some other species in the genus as well!), thus emphasizing their dubious value as a

means of distinction in certain cases.

As will appear from the key characters of the subgroup in question, it comprises

only three recognizable taxa deserving full specific rank, viz., duodecimmaculata

Rossi, chinensis Ckll., and excelsa spec. nov. Of the previously proposed names,

only jakovlewi Radoszk. is here re-established, to denote a subspecies of the

earliest described taxon of the group, duodecimmaculata.

210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Melecta duodecimmaculata duodecimmaculata (Rossi) stat. nov.

(figs. 38—42, 51—56, pl. 1 fig. 3)

Nomada ? 12-maculata Rossi, 1790, Fauna Etrusca 2: 110—111, no. *931 (Italy; sex not stated).

Nomada duodecimmaculata: Panzer, 1804, Syst. Nomencl. Schaffers Abb. regensb. Ins. 1: 35 (compar.

note with A. punctata F.). — Diniz: (with question mark), 1960, Mem. Mus. Zool. Univ. Coimbra,

266: 34 (list: Portugal).

Melecta plurinotata Brullé, 1832, Expéd. sci. Morée, Zool. 3: 343, pl. 48 fig. 13 (9 Morée). — Lepeletier,

1841, Hist. nat. Ins. Hym. 2: 442-443 (Q Bagdad; 9 Sicile; 2 Oran). — Lucas, 1849, Explor. sci. Al-

gérie 3 Hym.: 211 (“Oran, vers la fin de l’automne”; other locs. sec. Lepeletier). — Dours, 1874, Cat.

synon. Hym. France, Mém. Soc. Linn. Nord 3: 205 (France). — Morawitz, 1880, Bull. Acad. Imp.

Sci. St. Pétersb. 26: 492 (SE Mongolia; Samarkand). — Pérez, 1883, Act. Soc. Linn. Bordeaux 37

(ser. 4 t. 7): 307—308 (9 diagn., France ?). — Friese, 1893, Bienenfauna Deutschl. u. Ungarn, Ber-

lin: 61 (Mehadia, Hungaria). — Gribodo, 1893, Bull. Soc. ent. Ital. 25: 407—408 (9 ¢ Spain: Catalo-

nia and Andalusia (first description of 3, after Spanish examples!); Greece; Italy: Sicilia; and (par-

tim?) as far as Persia and Turkestan). — Friese, 1895, Bienen Europa’s 1: 156—157 (GQ key),

160—161 (2 Korfu; 9 Mehadia & Baleares). — Dusmet y Alonso, 1905, Bol. Real Soc. esp. Hist.

nat.: 152 (key), 154 (Spain, locs.). — Alfken, 1914, Mém. Soc. ent. Belg. 22: 235(9 Algeria, locs.). —

Strand, 1915, Archiv f. Naturgesch. 81A: 166 (3 9 Creta, name only). — Schmiedeknecht, 1930,

Hym. Nord u. Mitteleuropas: 830 (9 ¢ Südeuropa bis Südungarn). — Alfken, 1931, Konowia 10:

164—165 (pars! 9 3 key, with supposed M. 14-punctata Fisch.-Waldheim, © g sine patria!; Alfken,

1940, Sitzber. Ges. naturf. Freunde Berlin: 243 (syn. note, see citation below). — Garcias Font,

1953, Bol. Soc. Hist. Nat. Baleares 1: 10 (Mallorca). — Móczár, 1957, Fauna Hung. 19 Hym. 3: 31

(note: since 1886 not found in Hungary). —Iuga, 1958, Acad. Rep. Pop. Rom., Ins. 9 Hym. An-

thoph.: 208, 212—213 (Mehadia: Hungary). Syn. nov.

Melecta quatuordecim-punctata Fischer de Waldheim, 1843, Rev. et Mag. Zool. 13: 3—4(¢ “ad Ural flu-

vium superiorem’’). Syn. nov.

Melecta baeri: Alfken, 1935, Apidae in Wiss. Ergebn. Niederl. Exped. Karakorum, etc., Zool. I:

253—254 (9 Yarkand, notes).

Melecta söderbomi Alfken, 1936, Arkiv f. Zool. 27A: 21 (4 S.W. Mongolei, Lanchow, 28.iv.1928). Syn.

nov.

Type material. — China: 1 & (diss), Central China (prov. Kansu), labelled

“Sven Hedins Exp. Ctr. Asien/S.W. Mongol. Sòderbom” (print), ‘“Lanchow (Lan-

chou) 25.7.28 (9)” (pencil writing), ““Melecta söderbomi m. J. D. Alfken det. 1935”

(Alfken’s writing), “Typus” and “(5)” (print on red). Holotype (NRS).

Further material. — Europe. Portugal:1 g, Portugal (MNB). — Spain: 3 9, over white drawer label

“Melecta plurinotata Brul. & Lep. D. Ghiliani Espagne” (MT); 2 3, 288/1 M. plurinotata Br. Spagna, D.

Lichtenstein, det. Gribodo (MCG); 2 9, Hispania (OUM, MNB); 1 &, Barcelona (Cataluna),

Schmiedeknecht (SMF); 1 9, Elche (Valencia), 10-30.iii.1883, Friese (MNB); 1 9, Alicante (Valencia),

Callosa de Ansarra, 2.iv.1956, I.H.H. Yarrow (BM); 1 & (diss, figs. 41—42), Spanien, Lorca (Murcia),

24.v.1965, W. Linsenmaier (CL); 1 3, Granada (Andalusia), Schmiedeknecht, coll. A. Weis (SMF); 1

©, Andalusien, Pto. Sta. Maria, Hering coll. iv.1933 (MNB); 2 9, Gibraltar (OUM). — Balearic Is.,

Ibiza:1 4, Ibiza, ‘da Schmiedekn. plurinotata/lviza/Parente n. Anthophora hispanica” (unknown hand)

(MCG); 1 &, Ibiza, S. Augustin, Giner leg. (ZSM); 2 9, Ibiza, 20.iv.1883, Friese (MBUD) and id.

2.iv.1883, Friese (MNB). — Mallorca:1 & 1 9, Mallorca, 2.iv.1883 & 1 9, id., 2.v.1883, Friese coll. A.

Weis (MCG, SMF) & (MNB); 5 ©, Mallorca, Palma, 1895 (CCS & ML). — Italia: (Nto S): 1319,

Dintori di Roma (Lazio), Acilia, 12—14.iv.1932, M. 12-maculata Rossi (sic!), det. H. Hedicke (INER); |

1) This specimen unfortunately gives no information about its origin or collector’s indications, the ver-

tical pin-label only suggests that it might well be Brullé’s type specimen from “Morée” (Greece)!

LIEFTINCK: Palaearctic Melecta 211

©, Marino (Lazio), 1.v.1934, M. armata, det. Stocklein 1955 (ZSM); 1 &, Lazio, Roma, leg. Giordani

Soika (CGS); 1 g, Lazio, Civitavecchia, 24.iii.1935, O. Castellani (ML); 2 3, Foggia (Puglia), iv.1905

(IEP); 1 g, Brindisi (Puglia), Schmiedeknecht (MNB); 1 9, Calabria, Antonimina 1905, Paganetti

(MNB). — Yugoslavia: I g, Spalato (Split), Gasperini, M. plurinotata, det. Friese (MNB). — Greece:

1 ©, Graecia (MCG); 1 9, with old vertical pin-label “plurinotata Br.” (MP)'); 1 g 1 9, Korfu

(Kerkira), 20—30.v.1973, K. Vegter (coll. K. Vegter); 3 9, Steni, Euboea & Chalkis, Euböa, iv.1926.

Holtz (MNB); 2 g (1 diss., pl. 1, fig 1 & fig. 3), Athen, J. Sahlberg (MH & ML); 1 g, Athen, 7.iv.1927

(MNB); 1 & 2 9, Tinos, Erber (ML); 1 9, Naxos (Cyclades), Krüpper (MNB). — Crete: 1 3, 20/2

Creta, v.O., ex coll. Vogt (MA). — Turkey (Asia minor): 1 ©, Smyrna (Ismir), ex coll. Vogt (MA); 1 9,

Kleinasien, Sewdiköib, Smyrna, 3—4.17, La Baume (MNB). — Romania:; 1 9, SW Romania,

Mehadia 1886, Friese (MBUD). — Malta: 1 © ,with small white label ‘Malta’ over drawer label “M.

plurinotata Brul. & Lep.” (MT). — Cyprus:; 1 g 1 9, Limassol, 5.ii.1931, G. Mavromoustakis (CK); 1

d, Cyprus, Amathus, 25.11.1966, same coll. (FAG); 1 g, Limassol (ZSM) & 1 g, same loc., 23.11.1927,

same coll., coll. P. Magretti (MCG); 1 & 1 9, Limassol, same coll. (NRS) and series 4 9, Limassol,

same coll. (MNB); series ¢ 9, Limassol, various dates, ii—iii, all G. Mavromoustakis, ex coll. Alfken

(MNB); 14 1 9, Is. Cipro, Limassol, iii. 1932, Mavromoustakis (IEP). — North Africa. Algeria: 1 g 1

Q, Prov. d’Alger, Chellala, Jardin Romanetti, 1893, Vauloger, 9 with 138—97 (MP); 1 9, same loc.,

1895, M. plurinotata, det. Friese (SMF); 1 g 1 ©, Algeria, Mascara, Sidi Daho, J. Béquaert (MNB);

small series ¢9, Alger (OUM); 1 9, Sétif, coll. de Saussure (MG). — Libya: 1 9, Tripolitania, Sidi

Mesri, iii.1940, G. M. Martelli, M. plurinotata, det. D. Guiglia (BM); 1 9, Tripolis (MNB); 1 9,

Cyrenaika, Mars el Brega, ii.1942, Kirchberg leg. (MNB); 1 g, Tripolitania, Tripoli, 4.iii.1954, K. M.

Guichard (BM). — Egypt: 1 9, Egypt, Mansouriah, ii.1926, Min. Agric., R. Mabrouk coll. (MNB). —

Israel: 1 ©, Kappernaum, 11.iii.1935, coll. Hecht (CBS); 2 9, Jerusalem, Mt. Scopus — L., 22.ii.1946,

at Lavendula and Rosmarinus (CBS); 1 9, Palestina, Audja, v.1945, Bodenheimer leg., at Asphodelus

(MNB). — USSR: 1 9, S.O. Kazakhstan, Kurtagai, 7.vii.1959, leg. Scopin (MBUD); 1 ©, Turkestan,

Adjan (MNB); 1 g, Djarkent, Turkestan (MNB); 1 &, Tadzhikistan, Geb. Alai Pamir, 1890, M. 14-

punctata F.-W., det. Friese (MNB); 1 9, Turkestan 1912, M. 14-punctata F.-W., det.? (MNB); 1 ¢ (diss.,

figs. 53—55), Siberia, Krasnojarsk (Jenisei R.), (93° long. E, 56° lat. N), 56/Trybom 223/61 (red), and 1

©, same loc. and collector, 13/G (NRS). — USSR-China: 1 9, Jarkand (W Chinese Turkestan, So-

che), 1300 m, 8—27.iv.1930, J. A. Sillem leg., Ned. Karakorum Exped., Melecta baeri Rad., det. J. D.

Alfken 1933 (MA); 2 ©, “Jarkend Raquette” (print), and 2 ¢ 8 9, “Kaschgar (W Chinese Turkestan,

K’ashih), Raquette” (print) (NRS, ML); 1 & (diss., figs. 51—52) 10 9, (Chinese?) Turkestan, Fluss

Usek, v.1909, ex coll. C. & O. Vogt, acq. 1960 (MA,ML). — China: 1 9, Nord Pékin, Jehol, A. David

1865 (MP).

N.B. — In the last category of “Further material’’, a number of specimens are

listed originating from the USSR and frontier states of China (Tadzhikistan,

Kirgizyia and Sinkiang, in the interior of central Asia). With the exception of the

Kurtagai female, the pair from Krasnojarsk and the female from Jehol, they form a

homogeneous series of small bees measuring only 10—12 mm, thus equalling M.

chinensis in size. All are, however, characterized by shorter pubescence, slender

legs, sharply defined small, subcircular abdominal spots placed in a regular row,

much black at the thoracic sides, and absence of a white tuft between the scutellar

spines. Structurally, the male is rather similar to chinensis Ckll. (figs. 57—59), but

differs, among others characters, in the armature of the hind legs. These bees are

only provisionally classified under the present name and labelled by me “M.

duodecimmaculata subsp.?’’. At the moment this form is hardly worth naming, but

with much more material may ultimately prove to represent some kind of

“ecological subspecies”’ confined to higher altitudes.

This is evidently the species hitherto known in the literature as M. plurinotata

Brullé, originally described from Greece. It was probably Hedicke who first

212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

consulted Rossi’s work and, realising that the author’s specimen of “Nomada”

from Italy should have been a Melecta, acknowledged the validity of the specific

name given to it. Hedicke also examined Italian examples (collected in 1932 near

Rome, see above) and named them correctly, but, as far as I know, did not publish

about his findings. The necessity of a nomenclatural change was also fore-

Figs. 38—42. M. duodecimmaculata duodecimmaculata; 38, sternites 7 and 8 (4 Athens, Greece); 39, par-

tial ventral and dorsal view of genital capsule (same specimen, bristles partly omitted); 40, lateral as-

pect of gonostylus showing dorsobasal and ventrobasal processes (same specimen). — Figs. 43—44.

M. excelsa (3 holotype, Jalalabad, Afghanistan; 43, sternites 7 and 8; 44, partial ventral and dorsal view

of genital capsule (bristles partly omitted); g.a. = gonocoxal angle

LIEFTINCK: Palaearctic Melecta 213

shadowed by Alfken (1940), who wrote: “Es ist erstaunlich, was Rossi alles als

Nomada angesehen hat. In der Fauna etrusca.... hat er sogar die Melecta plurinotata

als Nomada 12-maculata beschrieben”. Yet Alfken did not mention the last name in

his later publications. With a view to verify his statement, I asked the opinion of

Herrn Max. Schwarz at Ansfelden, the well known specialist in Nomada, who had

ascertained already that this Italian bee could not have been a Nomada. The

following passages, referring to the scutellum and abdomen, are taken from

Rossi’s “Fauna etrusca” (part 2): “Scutellum porrectum, dentibus duobus

instructum, qui non apparent nisi pilis abrasis, sed nec postice productum, nec

emarginatum’’. And further: “Secundo & tertio segmento punctis utrinque

duobus; reliquis utrinque unico albis, adeo ut puncta abdominis alba, omnino

duodecim.” (loc. cit.: 111). These words are sufficient to prove the correctness of

Alfken’s note, so that plurinotata unfortunately must be dropped as a synonym of

duodecimmaculata (Rossi).

Next to the preceding, we have to consider plurinotata and quatuordecimpunctata

Fischer-Waldheim. While the first relates, of course, to populations of Rossi’s bee,

which is distributed all over the Mediterranean basin (see above), the last-

mentioned name applies to a small-sized individual from the Ural mountains. A

locality nearest to this is Kurtagai in NE Kazakhstan, whence I have examined

another small specimen (12 mm) indistinguishable from the western nominotype.

It will be seen that further toward the south we meet with a more profusely white-

haired form, jakovlewi. The whereabouts of Fischer de Waldheim’s type are

unknown to me, but that the species belongs here is almost beyond doubt, as

witness the scraps taken from the original diagnosis: “*.... scutello acute bispinoso...

segmente (2—4) maculis utrinque duabus niveis. Anus subbidentatus’’, — which

clearly point to a male of the present species, whose name antedates all others.

Consequently, it seems best to follow the opinion expressed by Pérez in Friese,

1895: 162), who placed it in the synonymy of plurinotata, i.e. of the nominate

species duodecimmaculata.

The last synonyms included in the above list are “baeri”, employed by Alfken

(1935) for a mis-identified individual of duodecimmaculata, and söderbomi Alfken,

which came from Lanchow in SW Mongolia, described by him one year later.

Remarkably enough, though possibly as a result of its remote habitat, Alfken

failed to compare his far eastern species with the present one and merely placed it

“in die Gruppe der M. armata Pz.”. As a matter of fact, the type of söderbomi, also

before me, is hardly distinguishable from a male collected at Krasnojarsk, in

Siberia. These two specimens are surprisingly similar to normal nominotypical

examples from the Mediterranean region, even with regard to the pubescent

pattern, as well expressed — though not stated — in terms of Alfken’s description!

The following additional description is based on specimens from the type

localities of both duodecimmaculata and plurinotata, hence on populations

occurring in the Mediterranean region and adjacent countries, i.e., approximately

between 47° and 33° lat. N, and 10°W—S0° long E.

Black and white pattern of head and thorax differing in both sexes.

Male. Black are: most bristles upon labrum, sides of clypeus, vertex medially

214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

and temples, lateral tuft below wings and — interrupted by white patch — a

narrow zone on lateroventral portion of mesepisternum; also the scutellum and

parascutella (except occasional white tufts on either side of the short spines and,

more rarely, a few recurved white hairs midway between the latter); and lastly,

middle of propodeum and metapleural sclerites. Coxae and trochanters with some

longish white hairs; femora all black, tomentum on closely punctate hind femur

posteriorly extremely short, dense and comb-like, the central carina almost acute,

extending along distal half to four-seventh; white external mid tibial pad elongate-

oval, incomplete (black) on both ends, the posterior fringe conspicuously white;

basal half or less of hind tibia white externally (except black knees), as are most of

the tarsal segments. Abdominal spots composed of longish, suberect silvery hairs;

with outer row of large spots on tergites 2 and 3, rarely continued rearward also

upon 4—5, most conspicuous on 2, and with inner row on 2—6, the spot on 2

smallest, circular, and occasionally vestigial. Disk of sternites clothed with black

hairs, 3—5 moreover with long and dense subapical brush of partly white raised

hairs. Tergal plate 7 closely longitudinally striatopunctate lacking interspaces,

usually with low middorsal ridge; distal border yellow-brown; recurved ventral

border with pair of narrow ridges close to apical margin. Genital capsule of large

size, 1.7—2.3 mm long.

Female. Differs as follows: White hairs covering clypeus more distinctly raised,

not markedly fanned. Thoracic pattern alternatingly black and white: mesonotum

white only as far back as base of tegulae, with pair of compact deep black spots

placed on each side upon middorsum, immediately behind these the dorsum being

also black, but with pair of white dots placed obliquely to the inside of tegulae;

Figs. 45—50. M. duodecimmaculata jakovlewi; 45, external view of left hind basitarsus (¢ Bairam ali,

Turkmenia); 46, abdominal pattern, right dorsolateral view (same specimen); 47, dorsal view of apex

tergite 7 (other specimen, same loc.); 48, ventral view of sternite 7 (same specimen); 49, partial dorsal

view of genital capsule (same specimen); 50, 9 abdominal pattern, right dorsolateral view (same

locality)

LIEFTINCK: Palaearctic Melecta 215

posterior parts black with conspicuous tufts of white only on each side of both

scutellum and propodeum, and (more rarely) a few recurved white hairs also

between scutellar spines; sides and ventral parts mostly black, except an isolated,

almost circular, white dot upon middle of mesepisternum. Abdominal markings

much as in male though more compact: with outer row of large spots on 2 and 3

(rarely wanting on 2) and inner row on 2—5 (rarely on 3—5 only); suberect hairs

and gradular fringes on sternites conspicuous but more evenly distributed and

invariably black. Outer faces of mid and hind tibiae more densely hairy,

concealing most of the surface.

The nominate subspecies and most characteristic “form” of the whole series, is

a large, conspicuously marked and densely hairy bee, occurring from Iberia

eastward into Turkey, but its range extends all over southern Europe in the same

direction as far as Mongolia and N China. Measured specimens before me from

North Africa are quite similar though generally somewhat smaller, the average

sizes being about 12 and 15 mm, respectively. As stated above, our series from

Central Asia comprise still smaller individuals agreeing with chinensis in size, but

the latter differ markedly in the coloured maculations and longer pubescence. I

am convinced that all except chinensis and excelsa belong to a single, polytypic

species.

Melecta duodecimmaculata jakovlewi Radoszkowski stat. nov.

(figs. 45—50)

Melecta Jakovlewii Radoszkowski, 1877, Horae Soc. Ent. Ross. 12 (1876): 333—334 (¢ “des environs

d’Astrakhan’’).

Melecta Jakowlevi: Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 183, fig. 39

a—c,i(g genit type).

Melecta plurinotata: Morawitz, 1895, Horae Soc. Ent. Ross. 29: 38—39, descr. notes, 9 & “Dus-olum,

Turkmenia, von Pomeranzew gesammelt”.

?Melecta Baerii (nec Radoszk.): Kohl & Handlirsch, 1889, Sitzber. Zool.-bot. Ges. Wien, Verh. 39:

273—274 (9 Aschabad, descr.).

Type material. — USSR: 1 ¢ (diss., genit. glued on card), with red-gold label in

poor writing “astrak(an)’”, “Type” (print on red), and ‘M. Jakovlewii Radoszk.

Type det. Dr. Enderlein”. Holotype (MNB).

Further material. — USSR: 1 ©, Turcmenia/Aschabad (Ashkhabad) Leder/Reitter, M. plurinotata,

det. Kohl (NMW); 1 & (diss., figs. 45—49), Turkmenia, with written label, transl. from Russian

“Transcaspian region, Bairam-ali, N. Simonov” and “coll. Kokuev” (print) (ZIL); 1 9 (fig. 50), with

two written labels, transl. from Russian ‘12.v.1960 Bairam-ali rayon, Zovkhoz”, “Bairam-Ali

Turkm(enistan), T. Atdaev”, and on reverse side “at wall of old fortress’, and ‘‘M. plurinotata Brullé,

det. Ponomareva 1960” (ZIL); 4 9, with printed labels in Russian ‘“Turkmenia, Kara-kala (Aral Lake?),

Siumy, Petrishcheva, VII.1931 (3 9) & 9—10.IX.1931 (1 9) (ZIL, ML). — Iraq: 2 9, Mesopotamia,

both with old vertical pin-label ‘‘Bagdad/orientalis’ and ‘‘plurinotata Br.” (MP); 1 & (diss.), Iraq,

Baghdad, 20.ii.1955, Fathi, M. plurinotata, det. Mavromoustakis, and 1 ©, Iraq, Abu Ghraib,

18.iii.1950, A. I. Derwesh, same identification (CKW).

The type of jakovlewi came from Astrakhan, the others from the SE side of the

Caspian Sea. They agree in most respects with our specimens from Baghdad

(Iraq), which for that reason are also referred here. Unfortunately, the internal

216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Figs. 51—56. M. duodecimmaculata duodecimmaculata; 51, ventral view of tergite 7 (¢ Fluss Usek, Tur-

kestan); 52, sternites 7 and 8 (same specimen); 53, frontal view of labrum (¢ Krasnojarsk, Siberia); 54,

dorsal view of apex of tergite 7 (same specimen); 55, sternites 7 and 8 (same specimen); 56, apex of right

gonobasis, dorsal view (same specimen). — Figs. 57—59. M. chinensis (3 Shanghai, China); 57, sternites

7 and 8; 58, partial ventral and dorsal view of genital capsule; 59, external view of right gonostylus,

showing dorsobasal and ventrobasal processes, bristles partly omitted

sexual organs of the type were cut into bits by Radoszkowski and are useless for

comparison, as are his sketches of the genital capsule.

The arguments in favour of restoring the name jakovlewi are based on the greater

density, predominance and purity of all white markings, most luxuriantly

developed in our small series from Turkmenia (figs. 45—50). In them the patches

on tergite 2 are not definitely interrupted, the outermost portion being the largest

and irregular in shape, emitting a subtriangular extension pointing basad and

nearly meeting the hind angle of a strongly arched collar on tergite 1. The spots on

3 are also unusually enlarged, the outer subcircular, the inner transverse and at

least twice as large again; spots 4—6 (&) or 4—5 (9) single, diminishing in size, all

except the last broader than the interspaces. Sternal plates 3—5 of male adorned

LIEFTINCK: Palaearctic Melecta 217

with white transverse bands tapering inward and narrowly interrupted medially,

composed of straight decumbent hairs projecting somewhat beyond hind margin.

M. jakovlewi had been synonymized already with plurinotata by Friese and

contemporary writers, but is here re-instated as a subspecies of duodecimmaculata,

the new name for the latter. The type measures about 15 mm, thus corresponding

with the average-size of the nominotype but, along with a number of closely similar

individuals, differs from examples inhabiting the Mediterranean region. The

vestiture on all parts of the body (including the white abdominal spots) is shorter,

composed of decumbent instead of suberect hairs, thus giving the insects a less

“woolly” appearance than those of more western occurrence. A striking feature

affects the enlarged innermost spot of the lateral patches on abdominal tergite 2.

These spots reach their maximum extent in our series from Bairam-ali and Kara-

kala, where they are completely fused together (figs. 45— 50). In this respect these

bees correspond very closely with the one briefly characterized by Morawitz, who

wrote: “… Auf dem 2-ten und 3-ten Hinterleibsringe sind 2 neben einander

stehende weisse Filzmakeln jederseits vorhanden, von denen die auf dem zweiten

häufig zu einer fast nierenförmig gestalteten zusammenfliessen” (loc. cit 38).

I am not quite sure about the identity of M. baerii sensu Kohl & Handlirsch,

from Aschabad; another specimen from the same locality listed above and labelled

plurinotata by Kohl, surely belongs to jakovlewi, as the spots on tergite 2 are

broadly coalescent. The type of jakovlewi, and the very similar Iraqi specimens as

well, differ from the above only in that the same spots, though much larger than in

typical duodecimmaculata, are not completely united. A character perhaps of

minor concern and therefore not entered into the key, affects the antennae. These

are, in both sexes, a little longer and more slender than in the nominotype,

segments 4—12 (13) being slightly longer than broad and of equal length, whereas

in the great majority of the better known subspecies the 4th is perfectly square and

just a little shorter than the next flagellar segments. I believe that these features,

taken together with those summarized in the key, justify its separation as a fairly

recognizable subspecies of duodecimmaculata. The abdominal pattern of a

Turkmenian pair and some structural details of the male are shown in figs. 45—49.

The southernmost range of jakovlewi in West Asia remains to be established. It is

worth noticing that the — very few — examples which I have been able to examine

from more northern countries, viz. Kurtagai (S Ural) and Krasnojarsk in the east,

differ in no way from the typical subspecies.

Melecta excelsa spec. nov.

(figs. 43—44, pl. 1 fig. 4)

Type and paratypic material. — Afghanistan: 1 & (holotype, mounted and

diss., figs. 43—44 & pl. I fig. 4), labelled “O. Afghanistan, Prov. Nengrahar,

Jalalabad (east of Kabul, nr. Kyber Pass), I—III.1965, legt. D. Povolny” (print); 4

d 4 Q (paratypes), “O. Afghanistan, Prov. Nengrahar, D. Povolny et coll”, and

same area, ‘(85) Nemla, 450—500 m, 29.3.1967” (4 & 3 2) and “(13) Nemla,

18.2.1966, Povolny & Tenora” (1 9) (MMB, ML).

A large species, evidently belonging to the duodecimmaculata group and closely

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

related to that species. There is nothing of real importance to improve on the

characterization in the key. The sexes are very much alike and can be at once

distinguished from the above mentioned taxon by the short-haired body and legs,

the more regularly arranged and sharply outlined white abdominal spots, and the

broad, contrastingly tinted smoky brown wing border. As in nearly all regional

Melecta, antennal rhinaria are present on 3—13, quite distinctly impressed,

becoming gradually smaller distad, the one on 3 being largest, more or less

sausage-shaped, the remainder subtriangular, except the one on 13, which is in the

form of a circular pit.

Wing expanse of photographed holotype male, 33 mm.

The habitat of M. excelsa is of great interest, since very few Melecta have so far

become known from Afghanistan. The holotype, though perhaps collected at a

much higher altitude than the females, does not in any way differ from the latter,

except for the sexual characters.

Melecta chinensis Cockerell

(figs. 57—59)

Melecta chinensis Cockerell, 1931, Amer. Mus. Novit. 466: 6 (43 Zi-ka-wei, E. China (= Shanghai), Piel).

Melecta 14 punctata: Alfken, 1931, Konowia 10: 164—165 (key 3 2, with M. plurinotata Brullé; pars!

Shanghai & Soochow only).

Material. — E China: | & (diss, figs. 57—59, topotypical M. chinensis Ckll.!), Shanghai, 4.iv. 1926,

coll. E. Suenson, M. /4-punctata F.-W., det. Alfken (ZSM); 1 £, Shanghai, i.v. 1891, with old written

label, “M. plurinotata similiter affin”. (MNB); 5 ©, Shanghai u. Umgeb., Dr. Eidmann, one with 2 labels

“M. 14-punctata F.W., det. Alfken”, and “nicht /4-punctata”, det.” (MNB); 2 &, with “Shanghai” and

“China”, respectively (MNB); 4 ©, Szetschwan, Omisen, exped. Stôtzner, Hedicke (MNB); 1 &,

Chekiang (Tsekiang), Tien-tai-shan, 30.iv.1935, H. Höne (MNB); 2 & 4 ©, Chekiang, Hangtchéou

(Hangchou), A. Pichon 1925 (2 & 2 ©), and same loc., no further data (2 9) (MP, ML); 2 &, Hanchow

1934/China, one with “chinensis”, det.?, the other with ‘tex coll. TC Ma, M. chinensis Ckll.”, det.?

(MNB, ML); 1 ©, Chine, Nanking (Nanching), J. de Joannis 1908 (MP); 16 & 2 Q, Prov. Fukien,

Kuatun, 2300 m, 27°40’ lat. N, 117°40' long E, 6—29.iv.1938, J. Klapperich (NMW, ML).

Male (additional), — The sternal plates are clothed all over with not very long,

suberect, mostly pale hairs and somewhat longer bristles, 3—5 moreover with a

subapical brush of almost depressed, mostly silvery hairs forming poorly defined

bands on either side of the median line; hind margin of sternite 5 shallowly

emarginate. The shape of the hidden plates 7—8 gives no clue as to the inter-

relation of this and allied members of the group. Curiously enough, sternite 8, with

its prominent marginal convexities, approaches most closely the form it has in

excelsa, which is altogether different in other respects (see also under

duodecimmaculata).

Our specimens agree with the original description, as far as it goes. Cockerell

compares his male of chinensis with that of ‘‘p/urinotata’’ (from Cyprus), stating

that it could be regarded as a subspecies of it. The combined characters

enumerated in the key seem to justify its position as a full species.

Without mentioning any locality in particular for his plurinotata, Alfken (1931)

gave a number of male and female characters in tabular form to separate this

species from the allied quatuordecimpunctata, the type of which he had, of course,

LIEFTINCK: Palaearctic Melecta 219

not examined. The characters employed refer to the colour and pattern of the

body pubescence, and the spinosity of the hind tibia and basitarsus of the female.

It is clear that plurinotata in this table conforms with the earlier described

duodecimmaculata, while the characters given for the other species from Shanghai

(called /4 punctata by Alfken), are obviously those of chinensis, as here understood.

N.B. — The type locality, Zi-ka-wei, is in the SW part of Shanghai, the

headquarters, or stronghold, of the Roman Catholic mission in east and central

China. It is also the site of one of the oldest observatories, libraries, and museums,

all founded and maintained by the mission. The museum was moved later into a

downtown area and renamed Musée Heude (pers. comm. of Prof. T. C. Maa).

II. REMAINING PALAEARCTIC SPECIES AND SUBSPECIES

Melecta fulgida spec. nov.

(figs. 60—70, 74, pl. 1 figs. 5—6, map 1, p. 223)

Type material. — Bulgaria: 11 4 6 9, SW Bulgaria, Sandanski steppe,

Ortakenei, 25—31.v.1967, at Anchusa and Stachys, together with Anthophora

crinipes F. Smith, M. Kocourek leg., some with “ashabadensis Rad., det.

Kocourek. Holotype Z and one paratype Z (one diss. pl. 1 fig. 5 & figs. 60—64,

67—69), and two paratopotypic 9, same loc. and dates, in coll. Kocourek; also

several topotypes of either sex (CK, ML).

Further material (from SW Europe south-eastward roundabout the Mediterranean). — France: 1 9,

Fréjus (Var), 28.iv—4.v.1976, H. Wiering (MA). — Yugoslavia: 1 4, Macedonia, Ohrid,

29.v—10.vi.1972, H. Teunissen (ML); 1 9, Macedonia, Gradsko, v.1916, Müllenhoff (MNB). —

Bulgaria: | 3, “Orta-kenei (M. tr. Balcani), D. De Magistris” (topotypical!) (IEP). — Greece: 1 Z

(diss., fig. 66), ‘“Parnass’’. (Mt. Parnes?) Vogt coll. (MA); 1 2 (fig. 65) 2 2 (fig. 70), Graecia, Patras,

4.v.1962, W. Linsenmaier (CL, ML); 1 9 (fig. 74), Graecia, Meteora, 26.v.1963, W. Schläfle (CL); 1 9,

Greece, Euboia, Eretria, 26.iv.1968, J. P. van Lith (CVL); 6 3, Graecia, Delphi, 11.iv.1963, together

with M. a. albovaria, K. Warncke (CKW, ML); | 2 (diss.), Greece, Ancient Olympia, 19.iv.1968, on

Asphodelus, (CVL); 3 3, Graecia, Lamia & Florina, 2.v & 5.vi.1964, W. & E. Grünwaldt; 2 3 (diss.),

Nemea & Korinth, 5—28.iv.1969, same coll. (CWG); 2 4 (diss.), Graecia, Amphissa & Kiaton,

17—29.iv.1973, W. Grosz, coll. Grünwaldt (ML); 1 9, Greece, Samos,22.iv.1962, H. Bytinski-Salz

(CBS); 1 2 (diss.), Graecia, Lesbos, Lepetimnos, 28.v.1975, H. Maticky (CG). — Rodos: 2 4, Profitis

Ilias, 800 m, 20.iv.1970, H. Teunissen, and 3.v.1971, M. A. Lieftinck (ML); 5 3, Rodos, Attaviros &

Profitis Ilias, 24—28.iv.1976, H. Teunissen (CT, ML); 1 9, Rodos, Epta Piges, 50 m, 20.iv.1971, M. A.

Lieftinck (ML); 2 9, Rodos, Lindos, 10.iv.1970, A. C. & W. N. Ellis (MA). — Kos I: 1 £ (diss.),

Asklepieion, 300 m, 1.v.1971, M. A. Lieftinck (ML). — Turkey (Asia minor): | 9, Asia min. 1890, coll.

Magretti, M. ashabadensis, det. Friese 1897 (MCG); | © (diss.), Amasia, Mann 1860, M. ashabadensis,

det. Kohl (NMW); 1 ¢ (diss.), Asia min., Erdschias, (= Erciyas) 12.v., Penther leg., ashabadensis, det.

Kohl (NMW); 1 ©, Asia min., Angora , iii.29, Náday 1911 (MBUD); 1 ©, Turkey, Ankara, 15.v.1961

(CVZ); 2 4 (diss.), Turkiye, prov. Ankara, Cubuk Baraj, 17—19.iv.1963, J. Leinfest (MA); 1 3, Turkey,

Amasya, 1600 ft., 22.v.1959, K. M. Guichard (BM); 1 4 (diss.), Asia minor, Amasya, 800 m, v—vi.1967,

H. & U. Aspöck (CMS); 1 ©, Trabzon Area, Turkey, 19.iv.1959, K. M. Guichard (BM); 1 ¢ (diss.),

Asia minor, Beysehir, 4—6.vi.1964, J.Gusenleitner (CG); 1 3 1 9, Turkey, Konya, ult.v.1965, J.

Gusenleitner (CG); 4 3 (diss.), Türkei, Ürgüp, medio vi.1960, W. Schläfle (CL, ML); 3 2 4 9, Türkei,

Ankara, 3.vi.1972 (3 ¢ 3 9), together with M. a. albovaria, and Ephesus, 7.iv.1972 (9), all K. Warncke

(CKW, ML); 1 4, Türkei, Göreme, 15.v.1972, on Rosa spec., K. Kusdas, M. Kraus (CKW); 1 3,

Dijarbekir, 6—7.vii.1937 (MBUD); I 2 1 ©, Brousse, coll. J. Pérez 1915 (MP); 1 3, Brussa, 1894, 1015/

220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

112 (MBUD); 1 & 1 9 (diss.), Asia min., Angora, 29.iii (©) and Seräj-Köj, 8.vi, leg. Náday 1911, M.

ashabadensis Rad., det. Friese 1893 (MBUD). — USSR: 1 g, Transkauk/Helenendorf/1886, M.

ashabadensis, det. Friese (MNB); 2 g 2 ©, Transkauk., Helenendorf 1886, M. ashabadensis, det. Kohl

(NMW, ML); 1 9, Armenia, Monastero Chérard, 13.vii.1963, A. Giordani Soika (ML). — Iran: 1 g

(diss.) 1 ©, 16/40 (on yellow disk), Perse, Aucher 16.40 (MP). — Syria: 1 ¢ (diss.), Syria, GO (lilac

label), 531/8 (black-rimmed square), 4 (red) (MBUD); 1 © (diss.), Akbes, C.D. 1891, coll. J. Pérez

(MP); 2 & (diss.), Syria, Leder, 854, M. ashabadensis, det. Friese 1893 (NMW); 1 3, Syria 1899, A.

Weis, M. ashabadensis, det. Friese 1893 (SMF). — Lebanon: | & (diss., pl. 1 fig. 6), Syria, Beyruth, ex

Staudinger (ML); 1 9, Syria, Gödl, 856, M. ashabadensis, det. Friese 1893 (NMW). — Israel: 1 g,

Palestine, Jerusalem, Mar Saba, 15 km SE, 15.iii.1922, P. A. Buxton, M. armata Panz., det. B. Uvarov

(BM); 1 9, Palestine, Jerusalem, 13.iii.1923 (BM); 2 ©, Jerusalem, M. Pic 1928 (MP); 1 & (diss.),

Jerusalem, J. Sahlberg 113 (MH); 1 ©, Jerusalem, M. ashabadensis Rad.? “vide long 2nd joint on the

flagellum”, det. Saunders, E. Saunders coll. 1910—266 (BM); 1 3, Haifa, 26.11.1977, A. Freidberg

(CBS); 2 3 1 g (diss.), Jerusalem, 15—30.iii, 1—15.iv.1939 & 3.iii.1940, all H. Bytinski, M. rugosa Drs.,

det. Mavromoustakis (CBS, ML); 1 ©, Jerusalem-Jericho road, 31.iii.1969, H.Bytinski-Salz (CBS); 2

2, Kabara Hills, Zarga Br., 10.iv.1946, from Anthophora nests (CBS); 1 ©, Israel, Aqua Bella, Hebrew

Univ., 22.111.1954, J. Wahrman (CBS); 4 © (diss.), Israel, Upper Galilea, Rosh Pinna, 600 m, 29.iii. 1951

& 1952, Verechson (CBS); 2 & 1 9, Jericho, -200 m, Wadi Kelt, 6—27.iii.1975 (2 &), Hisman Palace,

8.111.1975 (©), K. M. Guichard (BM); 1 9, Israel (?), Kfar Jecheyghal (?), spring, T. Kurtzmig (?), rugosa

Drs. var. © (unknown writing) (CBS); 1 & 1 9, Palestine, Dan, 20 & 26.iii.1941, H. Bytinski-Salz

(CBS); 1 ©, Jerusalem, 1—15.v.1939, H. Bytinski-Salz, M. rugosa Dours var., det. Mavromoustakis

(CBS); 1 9, Centr. Jordan Valley, Deganya A, 28.iii.1962, B 228, Y.Palmoni (AID); 1 g, Bethlehem,

Melecta sp., det. Blüthgen (ZSM); 1 3, Palestine, Mt. of Olives, 9.iii.1918, Major E. E. Austen (BM); 1

9, Palestine, Nazareth, 1800 ft, 16.iii.1920, P. J. Barraud, Crocisa luctuosa, det. Uvarov (BM); 1 9,

Benjaminah, 6.iii.1924, Asphodelus. Hedicke (MNB). — Jordan: 1 9, Trans Jordan, Nuwaigis near

Amman, 23.11.1922, H. St. J. B. Philby (BM); 2 & (diss.), Jordan, AP Baliat, 17.v.1974, and via H.

Elmosa, Calihut, 2.iv.1977 (ML); 1 &, Jerash, 30.iii.1979, K. M. Guichard (BM).

The following descriptions are based on both sexes of the typical series from

Bulgaria (pl. 1, fig. 5).

Male. — Labrum (fig. 60) in frontal view broadest at or slightly before middle,

surface concave, shining basally and upon the low tubercles, with few punctures of

different sizes, for the rest closely rugosely punctate lacking interspaces, a low

smooth median carina usually present on distal half; anterior border not upturned,

almost straight. Maxillary palpi 5-segmented, relatively short, the second longest.

Mandibles sparsely, finely striato-punctate, surface rather shiny. Clypeus little

convex, surface along anterior border glossy, with few large punctures, gradually

more finely and very closely punctate posteriorly; rest of head dull, closely

coarsely punctate, the interspaces smaller than one puncture width, except very

narrow, almost impunctate area (much smaller than in species like albifrons)

immediately beside each lateral ocellus. Antenna (fig. 61), thin and slender,

segment 3 longer than scape, more than twice as long as its width at apex and in

frontal view twice as long as 4, which is slightly longer than broad, as are also the

next flagellar segments, rhinaria distinct though small and feebly impressed, the

one on 3 elongate, the succeeding ones circular, gradually fading away on 6—12.

Thorax segments without peculiarities, all parts closely punctate even on

mesonotal disk, where the rather large punctures are about equal in size to the

somewhat shining interspaces. Basal half of tegulae finely punctate, the rest

dullish, smooth. Scutellar tubercles straight, slightly divergent, directed obliquely

upward and backward, shorter than surrounding pubescence. Legs thin and

LIEFTINCK: Palaearctic Melecta 221

Figs. 60—66. M. fulgida; 60, frontal view of labrum (4 holotype Sandanski, Bulgaria); 61, frontal view

of left antenna (same specimen); 62, segments 3—4 more enlarged (same specimen); 63, external view

of right hind tibio-basitarsus (same specimen); 64, oblique dorsal view of right hind tarsal claw (same

specimen); 65, partial view of right fore wing (¢ Patras, Greece); 66, dorsal view of tergite 7 (3° Par-

nass, Greece)

slender; all femora finely closely punctate on a shiny surface. Mid tibia straight in

profile, moderately swollen and broadened as far distad as slightly before the

truncated apex, which, besides carrying some short marginal bristles, is produced

posteriorly into a thick, bluntly triangular spine; outer face very closely punctate

basally (similar to fore tibia), but apical portion polished, with much larger and

fewer punctures, some of which bear strong bristles. Hind tibia slender, subequal

in length to femur, distal three-seventh to one-half of outer face uneven but

conspicuously polished and brilliantly shining, coarsely sparsely punctate, each

depression carrying a thick, suberect spine-like seta; apex obliquely truncated but

its border not excavated, produced posteriorly in a robust triangular tooth (fig. 63).

Basitarsi thin and slender, those of hind legs laterally compressed, gradually a little

broadened from base to apex, the latter terminating in an acute external tooth; the

same segment slightly but distinctly outcurved in posterior view, surface smooth

and rather superficially punctate, punctures smaller than interspaces. Hind tibial

spurs unequally long, both almost straight, the inner spur being not undulated.

Inner ramus of fore tarsal claw slender and a little shorter, flatter and broader than

outer, those of mid and hind claws only about half as long and twice as broad as

outer, distinctly more expanded than this and also more strongly curved and

pointed (fig. 64).

Wing venation brown, membrane invariably obscured, smoky brown, with

229 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

subhyaline areas occupying only most of the basal portion of fore wing, streaks in

first two submarginal cells, and an irregular spot just outside the third; third

submarginal strongly elbowed distally, almost as much as in albifrons and

immediate allies, though a little shorter than these (fig. 65).

Abdomen deep glossy black, apical sternal segments distinctly lighter; all

tergites finely superficially and not at all closely punctate, punctures successively

more widely spaced posteriorly on all segments, closest basally but all much

smaller than interspaces, postgradular (posterior marginal) areas rather broadly

impunctate, those of tergite 1 narrowly, of 2—5 and 6 very finely tessellate,

punctures at base of 6 more crowded together but still smaller than interspaces.

Sides of exposed portion of tergite 7 strongly converging, dorsal surface hollowed

out and shining. with widely spaced punctures, apex narrow, posterior margin with

distinct crescentic emargination and rounded edges, border dark brown; apex in

ventral view without indication of tubercles or ridges. Sternites similar, but much

less shiny and more densely punctate than tergites, especially 4 and 5—6, the

whole surface of these bearing punctures. Sternite 6 lacking a distinct median

impressed area, hind border evenly rounded; 7 and 8 of characteristic shape, basal

arms of 7 relatively broad, almost straight, disk more or less square, in the form of

a thin flattened plate, apex not emarginate, almost bare, lacking bristles or setae;

apex of sternite 8 projecting, ending in a thin bilobed plate (fig. 67). Genital

capsule as in fig. 68.

Pubescence generally thin and fluffy, consisting of finely branched black and

white hair nowhere concealing the surface, except at sides of tergites 1—5 (9) or

1—4 (9) and partly on outer faces of tibiae. Labrum, mandibles outwardly, lower

part of paraclypeal area, upper part of genae, and a line bordering inner orbits,

with long raised black bristles; hair on genal area finely branched. Clypeus for the

most part bare, rest of head above clothed not very densely with white as far back

as about halfway between base of antennae and median ocellus, anterior

pubescence partly depressed, posteriorly raised and much longer; genal and

occipital areas with mixture of back bristles and silky white hair. Antennal scape

with few black bristles roundabout and a fringe of long, mostly white, hairs

anteriorly. Thoracic vestiture thin and diffuse, alternatingly black and white:

mesonotum anteriorly with fairly broad white collar extending back as far as a

little beyond anterior border of tegulae, this collar either entire or subinterrupted,

some black hairs usually being interspersed on either side of the middle; an

indistinctly triangular spot of white and better defined white tufts also in front of

tegulae, the latter occasionally forming a whole with the collar; behind this the

dorsum is black, but more inward behind tegulae are a second pair of whitish tufts,

just in front of the parascutella; entire scutellar area and posterior thoracic

sclerites black-haired save for large white metapleural tufts on either side behind

the wing bases; hair at sides of thorax long and raised, mainly black but in the

centre of mesopleuron an ill-defined tuft of white; ventral parts of thorax black.

Legs sparsely clothed with mainly black hairs, which are long and partly white on

coxae and femora, much shorter but mixed with long black bristles on tibiae and

tarsi; all femora posteriorly with long raised black hairs, those on distal parts being

223

LIEFTINCK: Palaearctic Melecta

wy 00S

saroads D199J2W aWOS JO Sp10991 AyEdoT ‘| dew

07 Ol

VOVILdADAV #

VIINILNY *

val DINI ®

224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

even longer than the diameter of femur; tibiae less hairy, but at least outer faces of

mid and hind tibiae with an elongate patch of much denser silky-white appressed

tomentum, these spots nearer base than apex and ill-defined, not broadened or

pad-like on mid tibiae, which also lack a posterior fringe of long hairs; traces of

white frequently also on outer faces of fore tibiae. Abdominal tergites 1—5 each

with pair of pure white postero-lateral hair spots, hairs on 1 long, tufty and raised,

the innermost exceeding hind margin of tergite, those on 2—5 regular, compact,

transverse, subrectangular and depressed, spots on 2—3 subequal in size, about

twice as broad as deep or slightly broader, but separated by a distance four to

three times their own breadth, those on 4—5 smaller, about half as broad as the

black interspaces, on 5 only half the size of preceding spots; tergites 6—7

invariably unmarked. Dorsal hair otherwise short and scanty, all setae suberect

and arising from the punctures, but much longer black bristles are present at base

and sides of tergite 1, alongside and at graduli of 2—6, and also on disk (with

additional marginal fringes) of 7. Sternites 1—2 clothed sparsely with longish black

setae, which on 3—6 become shorter, depressed and more numerous, arising from

the punctures, but do not quite conceal the surface.

Wing expanse (¢ paratype, Bulgaria, pl. 1 fig. 5), 26 mm.

Female. — Strikingly similar to male in body sculpture and pubescent pattern,

but apart from the more obvious sexual characters, the following differences can

be noted.

Antennae a trifle more slender but practically of the same length by having all

segments relatively longer, especially 3 and 4; rhinaria wanting. Inner ramus of all

tarsal claws about half as long as outer but distinctly broader and more flattened

than this, much as in male; apex rather abruptly hooked and acutely pointed.

Third submarginal cell, though varying in length, averages a little shorter, with its

apex less strongly elbowed than in male, several individuals, however, showing

hardly any sexual difference in this respect. Legs including tibiae and basitarsi of

the same slender form and consistency, polished areas identical; hind basitarsus

similarly and but feebly outbent; white pubescent patch at outer face of mid tibia a

little less dense and silky, several short, black, suberect spinulose setae remaining

clearly visible between the decumbent hairs. Posterior fringes at mid and hind

femora shorter than diameter of femur. Abdomen shaped as in male, the raised

black and white pubescent spots on dorsum and sides of thorax of the same length

and equally diffuse; compact lateral tergal spots also similar but restricted to 1—4.

Pygidial plate when fully exposed (fig. 70) almost twice as long as its breadth at

extreme base, sides rectilinear or a little incurved before halfway length, then

straight-lined and less converging, margins upturned before apex; surface flat,

shining, but very finely tessellate except at apex, with low but distinct median ridge

that broadens at tip; at either side of the pygidial plate the tergite bears a dense

fringe of straigth stiff bristles, the longest of these surpassing the rounded apex of

the plate. Sternites much as in male, but posterior marginal areas of I—5 markedly

broader medially and more abruptly becoming impunctate.

The sexes are equally variable in size. Measurements (type series), length

LIEFTINCK: Palaearctic Melecta 225

Figs. 67—70. M. fulgida; 67, sternites 7 and 8, scale line 1 mm (¢ holotype, Sandanski, Bulgaria); 68,

partial ventral and dorsal views of genital capsule (same specimen); 69, external view of right gonosty-

lus, slightly flattened (same specimen); 70, dorsal view of pygidial plate, bristles and hairs omitted (9

Patras, Greece)

9.5—10.8 mm, fore wing 8.5—9.0 mm; length (remaining specimens), 8.5—15.0

mm.

Variation. — In respect of body size and pubescent pattern, the typical

individuals — all from one locality — form a homogeneous series, though not all of

them are quite alike as to the extent of black and white spots. Yet fulgida, like most

other melectines, is evidently a very variable species throughout its known range.

True, very dark specimens and profusely white-marked bees, whether or not

varying in size, need not have been collected at the same time and in exactly the

same locality, but populations of these extremes frequently occur in close

proximity in the same area. In our material specimens agreeing closely with the

type series are from the following localities: — a male from Ohrid in Yugoslavia;

both sexes from Delphi, ‘“‘Parnass’” and Samos in Greece, as well as others from

226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Helenendorf, Konya and Beysehir in Turkey. Our series collected in Greece

(Patras, Olympia and Euboea), Lebanon (Beirut) and Palestine (Jericho,

Bethlehem and Dag) include males as well as females varying much in size (body

10.0—13.5 mm), the largest among them at the same time being more robustly

built. They are remarkable by having the light thoracic pubescence much more

extensive by fusion of the white tufts, which together form a very broad collar,

whereas in some of them the white abdominal spots are so small as to be reduced

to mere points on tergites 3—4, the spot on 5 being occasionally absent in the

male. On the other hand, all individuals from Rodos and Kos, off the westcoast of

Turkey, are very white and also superior in size, measuring 12.0—15.0 mm in

length. They exhibit a pattern indistinguishable at first sight from large-spotted

examples of M. albifrons albovaria and tuberculata spec. nov., two species with

which it was associated in more than one locality. ')

Careful examination of all salient characters of fulgida has revealed no structural

differences between representative specimens from nearly all countries whence

the species has become known. It has apparently no near allies, although remote

(yet undeniable) affinity is suggested with the albifrons group, approaching the

latter somewhat in the form and sculpture of the hind legs (including the tarsal

claws), the shape of the third submarginal cell, and the female also in the

somewhat similar form of the pygidial plate. However, it differs widely in the

antennal structure and male genitalia, being in fact easily distinguished from all

other Eurasian species by a combination of characters, as follows: (1) antennal

segment 3 fully twice as long as 4; male rhinaria incomplete distally, not deeply

impressed; (2) apices of hind tibia and basitarsus produced posteriorly into a

triangular process; (3) exterior surface of apical portion of mid and hind tibiae

uneven, though polished and brilliantly shining, bearing scattered setiferous

punctures, the thick setae triangular and spine-like, with few long dark bristle-like

hairs interspersed; (4) fore wing membrane strongly enfumed save basally, disk

with subhyaline spots; (5) male tergite 7 narrowly tapering, apex distinctly excised;

(6) male sternite 7 of peculiar shape, ending in a thin subrectangular plate; (7)

female pygidial plate slender, gradually narrowed with almost straight sides; (8)

pubescent spots on dorsum and sides of thorax ill-limited, made up of long and fine

raised hairs.

An additional feature not expressly stated in the description is, that the white

tufts behind the wing base on each side of the thoracic dorsum, if at all present, are

1) On Rodos and other islands of the eastern Mediterranean, many Anthophoridae were accumulated

over the years along with their melectine parasites and dealt out among several museum collections.

As to Melecta, collecting was done indiscriminately by various entomologists including myself. At

first the composite nature of these captures was not clearly understood, but with the discovery of

better specific characters it became evident that several closely similar taxa may occur simultane-

ously in any given locality. In fact it was established only recently that M. fulgida in Rodos, besides

keeping company with the two species mentioned above, may be attended also by two (or even

three) other congeneric species, all of equal semblance. Thus there is definite proof that during the

spring in this one small island at least five species of Melecta may occur together in any suitable loca-

lity.

LIEFTINCK: Palaearctic Melecta 227

isolated, never continuous across middle of scutellum, which always remains

black.

With this species the opportunity is given to spend a few words on the armature

of the mandibles in Eurasian Melecta. Following the example set by Michener et

al. (1978), I have figured one of the mandibles of some regional species. As was to

be expected, they are similar to those of Thyreus (loc. cit., fig. 15), almost

invariably bidentate in fresh specimens, i.e. armed with a single, flattened

submedian (pre-apical) tubercle, either subtriangular or more or less molar-

shaped. Like the apical tooth, the organ is liable to wear off rapidly with age, an

abraded mandible being shown in fig. 72. In the present species, however, the

inner tooth is apparently obliterated or altogether absent, even in fresh individuals

(fig. 74).

Melecta luctuosa (Scopoli)

(figs. 2, 4, 8, 79 —96)

Selected references:

Apis luctuosa Scopoli, 1770, Annus hist. nat. Lipsiae, 4: 13 (Hungary). ?‘‘Apis sexta’ Schaeffer, 1766,

Icones Insectorum.... etc. Ratisbon. 1(1), tab. 22 fig. 14 (insect, sex?), “Apis sexta. Sechste Biene”.

? Apis albifrons: Rossi, 1790, Fauna Etrusca 2: 111 (compar. note with Nomada ? 12-maculata).

? Apis punctata: Panzer, 1804, Syst. Nomenclat. Schäffers Abbild. regensb. Ins. 1: 34—35 (comments on

Schaeffer’s picture of “Apis sexta’’).

Melecta punctata: Lepeletier, 1841, Hist. Nat. Ins. Hym. 2: 441—442(¢ 9 partim! ‘Toute la France”).

? Melecta fasciculata Fischer de Waldheim, 1843, Rev. et Mag. de Zool. 13: 3—4 (? sex, ““Hab. ad Ural

fluvium superiorem’’).

Melecta luctuosa: Pérez, 1883, Actes Soc. linn. Bordeaux 37 (ser. 4 t. 7): 305—307 (inclus. key 49, 9

partim, not £?). — Radoszkowski, 1886, Horae Soc. ent. Ross. 20: 18, pl. 3 fig. 15 (3 genit. caps.,

Askhabad). — Pérez, 1890, Cat. Mellif. Sud-ouest, Actes Soc. linn. Bordeaux 44: 166, pars? (Saint-

Sever, Bordeaux, Tarbes, Toulouse). — Friese, 1893, Bienenfauna Deutschl. u. Ungarn: 37 & 61

(Mecklenburg; Thiringen; Elsasz. Budapest; Fiume; Triest). — ? Radoszkowski, 1893, Bull. Soc.

Imp. Nat. Moscou, new ser. 7 (2—3): 178—180, fig. 30 a—c, i (3 genit., loc.?). — Gribodo, 1893,

Bull. Soc. Ent. Ital. 25: 408—412 (pars! exclus. vars. meridionalis Grib. & leucorhyncha Grib.). —

Friese, 1895, Bienen Europa’s 1: 162—164 (pars, exclus. syn. & vars.). — Saunders, 1896, Hym. acul.

Brit. Is, etc. London: 345 (key), 345—346, pl. 45 fig. 4, 9, 10 (locs., “associates with Anthophora retu-

sa”). — Dusmet y Alonso, 1905, Bol. Real Soc. esp. Hist. Nat: 152—153 (pars) (Spain locs.). —

Frey-Gessner, 1907, Fauna ins. helv. Hym. Apidae 2: 143—145 (pars, Switserland, locs., ecol. notes).

— Alfken, 1914, Mém. Soc. ent. Belg. 22: 235 (pars! exclus. var. acutivalvis Alfk. & ““Stammform

9”). — Dusmet y Alonso, 1915, Mem. Real Soc. esp. Hist. Nat. 8 (7a): 330 (pars ? Morocco). — Bi-

schoff, 1927, Biol. Hym. Berlin: 399 (hosts). — Schmiedeknecht, 1930, Hym. Nord u. Mitteleuropas:

830. — Giordani Soika, 1936, Boll. Soc. Ent. Ital. 68: 47—48, figs. 1—2 (early larval stage). —

Stoeckhert, 1954, Fauna Apoid. Germ., Abh. Bayer. Akad. Wiss. N. F. 65: 64 & 74—79 (full bibli-

ogr. Pal. Apoidea). —Möczär, 1957, Fauna Hung. 19 Hym. 3 Apidae: 32—33 (key 9 g, pars, Hunga-

ry). Möczär, 1958, Rovart.Köslem 11 Cat. Hym. 14: 414 (distrib., ethol. Hungary). — Iuga, 1958,

Fauna Rep. Pop. Rom. 9(3) Hym. Apid. Anthoph.: 208—209 (pars, exclus. syn., Romania). — Lief-

tinck, 1958, Comment. Biol. Helsingfors 18.5: 23 & 26 (key ¢Q), fig. 8, 15, 30—34 & 37, pl. 1 fig.

9—10(¢9Q struct., Hungary & Holland). — Osychniuk, 1959, Bull. Acad. Nauk Ukraine SSR, Kiev:

80 (pars!) (¢ © Ukraine, flower records). Osychniuk, 1960, Entom. Obosz. Akad. Nauk USSR Mos-

cow 39: 391 (tab., flower records, Ukraine). — Leclercq, 1965, Bull. Inst. agron. Stat. Rech. Gem-

bloux 33: 114 (Belgian locs. prior to 1950). — Lieftinck, 1972, Tijdschr. Ent. 115: 256 (hosts), 264. —

Leclercq, 1971, Atlas provis. Ins. Belg. Gembloux, carte 548 (Belgian locs. prior to 1950). — Kullen-

berg, 1973, Zoon (Uppsala) Suppl. 1, pl. 2 fig. 6 (good col. phot. of live 9, Sweden). — Erlandsson,

1976, Entomologen 5: 20 (Swedish locs., hosts).

228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

9 Melecta luctuosa: Radoszkowski, 1886, Horae Soc. Ent. Ross. 20: 18, pl. 3 fig. 15a—c & i, g genit. (g

Transcaspia).

9 Melecta Eversmanni Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 180 (@ only,

not fig. 3la—c & i, g genit.!), “Orenbourg, Astrakan (Ryn-Peski), Tachkend”. See under that spe-

cies.

Melecta Eczmiadzini Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 181, fig.

33a—c & i(¢ genit.), 9 incert. (3 Eczmiadzine). Syn. nov.

Type material. — USSR: 1 ¢ (diss, fig. 81—82, right ant. missing), Caucasus,

“Caucas/Portz” (print on pale blue), ‘“Eczmiadzin” (Radoszkowski’s writing),

“Type” (print on orange), ‘“Fass.” (unknown handwriting), ““Melecta eczmiadzini

Rad. Type, rev. Dr. Enderlein”. Holotype M. eczmiadzini Radoszk. (MNB).

M. luctuosa is one of the most widely distributed Eurasian members of the genus,

ranging approximately from lat. 60° to 40° N., the centre of its area lying between

lat. 40° and 50° N., although the species may occur as far east as long. 90° in west

Figs. 71—80. Dorsolateral (external) view of mandibles of Eurasian Melecta; 71, M. albifrons albifrons

(3 9 Antibes, S France); 72, M. albifrons albovaria (worn ©, Montagne Noir, S France); 73, M. albifrons

nigra (3 Lazio, Italy); 74, M. fulgida (fresh 3, Meteora, Greece); 75, M. transcaspica (9 Konya, Tur-

key); 76, M. amanda (9 Krakow); 77, M. guichardi (9 Israel); 78, M. italica (2 St. Guilhem, S France);

79 , M. luctuosa (3 Peña Ubina, Spain); 80, M. luctuosa (3 Iran). All except fig. 72 taken from fresh spe-

cimens, drawn on the same scale; bristles omitted

LIEFTINCK: Palaearctic Melecta 229

Central Asia. With the exception of data included in publications dealing with the

bees of northwestern Europe, /uctuosa has been frequently confounded in the

literature and collections with similarly looking species occurring in more

southern and eastern parts of Eurasia, i.e. in countries with a milder, sub-tropical

climate. Males of /uctuosa, though bearing a close prima facie resemblance with

several other congeners, are always easily recognized, viz. (1), by the absence of

impressed olfactory organs on the ventral face of the antennal flagellar segments,

and (2), by the unmodified and normally pubescent outer faces of the mid tibiae

(see specific key). Females, on the other hand, are much more difficult to identify

with certainty and often indistinguishable — even structurally — from one or more

congeners inhabiting the same area, with which they often mix, but whose males

are on all occasions easily recognizable. Identification of females originating from

countries in the temperate zone of northwestern and parts of central Europe,

where no Melecta other than albifrons and luctuosa have so far been found, offered

no problem as both are easily held apart. These countries are: Scandinavia, Great

Britain, the Benelux, West and East Germany, Poland, Czechoslovakia, Austria,

and also the greater part of France (to about lat. 46° N.). To save space, drawer- or

pin-labels of museum specimens from these parts of its range (all based on

specimens examined by myself) are given in condensed form and simply replaced

by locality lists, arranged alphabetically for each country, as was done for the well

known nominotypical M. albifrons (Forster). In some cases certain territorial

Occurrences are mentioned separately. Countries are arranged somewhat

arbitrarily, from north to south (east).

As indicated above, recognition of luctuosa females collected outside the

northwestern territory of Europe often proved impossible, except of course under

special circumstances, e.g. when both sexes were captured simultaneously in some

isolated or remote locality. I think that the most reliable impression of the

distributional pattern of luctuosa can be obtained by enumerating all available data

of fully authenticated males (whether or not accompanied by females), and by

copying all labels in full. In that way comparisons can be made with the specified

data available on the labels of other species. In all instances where only solitary

females had come in hand, it has been expressly stated, indicating that they are

regarded conspecific. Other localities are left without the prefixed sex symbol,

which means that the proper identity of these bees is a little doubtful. The status of

a considerable number of soiled or worn females — mostly from the southern

Mediterranean and Near Eastern countries — had to be left undecided; these

specimens were simply omitted and left unidentified.

Further material. — Sweden: 2 & 1 9, Ostro Gothia (Östergötland), Aby, 30.v & 6.vi, leg. Haglund

(NRS), and 1 g 2 9, “V.G./Schh” & ““GL/Bhn-P.Wg” (NRS). Also known from the islands Gotland

and Oland (S. Erlandsson, 1976 & in litt.). Apparently rare. In Fabricius’ collection (MC) is a typical 9

(sine patria) with a label “Fals-Samml unter punctata” (recent handwriting), which means that the

insect had originally been placed under M. punctata F. (now a. albifrons Forster) (pers. comm. by B.

Petersen, Copenhagen). — Denmark: no specimens examined. Recorded by Jorgensen (in 1921) from

three localities on Jylland (Jutland), and two others on Jylland and Sjaelland (Zealand), communicated

by Dr. Borge Petersen. — Netherlands: Formerly (prior to 1950) widely though sparingly distributed,

except in the northern provinces (about 10 more records), nowadays a scarce species. Only eight

authentic localities from 1950 onward in the central and southeastern provinces. — Belgium: Also

230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

more widely spread before 1950 (21 locs., all in central provinces); since then apparently nowhere

turned up again. — Great Britain: 1 & 2 ©, Angleterre, coll. O. Sichel 1867, with old label “3 M.

luctuosa Scop. & Angl. F. Sm(ith)” and © with “Baly, M. luctuosa Q” (MP). Restricted to the southern

counties of England. In the British Museum (Nat. Hist.) are few specimens from Berkshire,

Buckingham, Essex, Gloucestershire, Surrey and Sussex. — West Germany: Atzwang; Bamberg;

Bensheim a. B.; Bonn (fig. 94); Ebelsbach nr. Bamberg; Frankfurt & Schwanheim nr. Frankfurt a/M;

Furth; Hannover; Kaiserstuhl; Karlstadt (Spessart); Koln; Lorelei (Rheingau); Lüneburgerheide;

Marburg/L.; Mühlhausen; Starnberg (Ober-b.); Strassbourg; Soden nr. Aschaffenburg); Tübingen;

Vilshofen (Nordb.); Waldorf/Gulda; Weilburg (Nassau); Weissenfels (Elsasz). — East Germany: gd 9,

Altenburg; Berlin; Dolgelin & Falkenberg (Mark Brandenburg); Frankfort a/Oder; Greifswald; Haabe

& Jena (Thüringen); Hainburg (Hundsheimerberg/Donau); Kyffhäuser; Naumburg a/Saale; Mittl.

Odergegend; Saaletal; Schwerin (Mecklenburg). — France: Nyons (Drôme) being the northernmost

locality of congeneric species having a more southern or eastern distribution, we may safely assume the

46th degree of latitude to form a boundary line for the three or four other species resembling /uctuosa in

certain respects. Examples from French localities to the south of this line, enumerated by Leclercq

(1965), would seem to require re-investigation. — Northern France: Finisterre, Fouesnant (Bretagne)

and Vitré (I. et-V.); Paris and Fontainebleau (S. et M.); Villers Allerand and Bazancourt (Marne);

Viarmes (?), v.1902, L. Chopard (MP). — Southern France: 4 9: St. Sever (Landes), coll. J. Pérez

1915 (MP); Argent (Corrèze), 25.v (MP); Tarbes (Htes Pyrén.), lilac disk, coll. J. Pérez 1915 (MP, ML);

Mont Canigou & Vernet-les-Bains (Pyren. or.), A. Weis 1908 (MP). — © only: Carpentras (Vaucluse),

v.1952, P. M. F. Verhoeff (ML); Callian (Var), 11.vi.1931, Th. Steck (NMB); Fréjus & Valescure (Var),

28.v.1971, K. M. Guichard (BM); Valescure, 15.iv.1913 & Hyères, iii.1898 (BM); Croix-Vaimer (Var),

v.1952, P. M. F. Verhoeff (ML); Vallouise (Htes Alpes), vi.1934 & 18.vi.1939, P. M. F. Verhoeff (ML);

B.-Alpes, 26.vi.1948, P. M. F. Verhoeff (ML); Montpellier, D. Lichtenstein, M. punctata Latr., det.?

coll. Gribodo (MCG). — Switzerland: 39; Basel; Valais, E. Favre (CB); Genève, 222/61 (red),

Sauss(ure) (NRS). — © only: Belp; Bern; Biel; St. Blaise; Lausanne; Locarno; Lugano; Sierre: (diss.,

fig. 87); Sion; Alp. Sussillon; Useigne. — Portugal: 29, Porto (Oporto), and Porto (MUC); &

Coimbra, O. Hospital (MUC); Rezende, 15.iv.1950, N. F. d’Andrade (4 only, ML). 9: Cardigos; Favo;

Figuera da Foz, 20.vi.1967, J. van der Vecht (ML). — Spain: £ 9, Bilbao (Basque), coll. J. Pérez 1915

(MP); g, Burgos, Sta. Maria del Invierno, iv-v.1964, I. H. H. Yarrow (BM); 2 ¢ (both diss.),

Navarredonda de Gredos (Avila E.), 2.vi.1979, H. Teunissen (CT, ML); 9, Barcelona, 11.iv.1924, M.

Martens, M. luctuosa, det. Alfken (SMF); ©, Albarracin (Aragon), vi.1953 (CS); ©, Albaracin,

H.Wagener (MNB); 4, Biescas (Huesca), 13.v.1953, I. H. H. Yarrow (BM); 9, Espagne (Ma.), Puerto

El Paular, 9.vii.1950, F. Schmid (CB); 2 ©, Batuecas nr. Salamanca (Léon), 13—15.v.1976, at nesting

site of Anthophora dispar Lep.(?), E. Asensio (CAV, ML); 9, Sierra Guadalupe (Caceres), v.1904, G.

Schramm (MP); 2 & (fig. 79), Espagne, Pefia Ubina (Valencia), 2.vii.1963, R. Desmier de Chenon

(INRA); Alicante (Valencia), Callosa de Ensarria, 2.iv.1956, I. H. H. Yarrow (BM); E. Spain (Valencia),

6 km NE Blagaz, 2.v.1960, exped. Leiden Mus. (ML); g, Espagne, “Hisp. 69”, O. Sichel 1867 (MP); ©,

Spagna, coll. Magretti (MCG); 2 9, S. Spain, Ronda (Malaga), 1000 m, 1—S.vi.1974, K. M. Guichard

(BM); 9, Andalusien (MNB); series 4, Algeciras (Cadiz) (NM W). — Balearic Is., Ibiza: No males!

©, Balearen, Ibiza, ult v.1956, M. luctuosa, leg. & det. Rebmann 1961 (SMF). — Mallorca: No males!

©, Mallorca, coll. Gribodo (MCG); Palma, 4.iv.1958 and Mallorca, without further data, A. Compte

Sart (ML); Porte Golom, 18.v.1956, F. Keiser (NMB); 9, Mallorca, 14.iv.1883, Friese (MNB). —

Morocco: A few doubtful © from the High Atlas are probably /. leucorhyncha Gribodo! (see under that

species). — Algeria: No males! 9 Algérie, Mascara, 28.iv.1908, Dr. A. Cros (MP); Algerien, Batna

and Lambesa, vi.1891, leg. Handlirsch (NM W). — Poland: No males! © only: prov. Pinczöw, vii.1955;

prov. Zabkowice, Muszkowice & Stolek, v.vi; E. Poland, Sandomierz a/Wisla (Weichsel), vi.1953; all

leg. W. J. Pulawski (CP, ML). — Czechoslovakia: ¢ 9; Dobrany (Dobrzan, S of Plzen) (Bohemia),

1.vi.1930 (ML); Jinonice (Bohemia), v.1953, Pádr (MUC); Bohemia, Chodau, v.1977, R. von Stein

(BM); Böhmen, 16.v.1904 (MNB); series 49 (diss., fig. 93), Brno (Moravia), Räjecek, v.1937, A.

Hoffer (MMB, ML); Ceje (Moravia), vi.1940, A. Hoffer, and Velke Pavlovice, leg. Stricha (ML);

Kobyli (Moravia), 21.v.1965, M. Kocourek (CK); Hostivor, 20.v.1953, leg. Pàdr (CK); Hvanice &

Hodanice (Moravia), 10—14.v.1942, M. Kocourek (CK); Vyskov (Moravia), 14—15.v.1965, M.

Kocourek (CK); Dobsina (Slovenia), vi.1954, Pàdr (MUC); Gbelce (Slovenia), 10.vi.1956; Jinonice,

1.v.1953, leg. Pádr (MUC). — Austria: 49, many locs., a.o. Bad Hall; Bisamberg; Bucklige Welt;

LIEFTINCK: Palaearctic Melecta 231

Dornbach (N.O.); Eisleben; Gottlesbrunn (N.O.); Innsbruck; Leopoldsberg; Linz; Mannersdorf

(Leithagebirge); Maschegg; Neusiedl; Oberweiden (N.Ò.); Plesching (Ob.O.), 5.v.1952, M. Schwarz

(CMS); St. Pauls (Tirol); Villach; Weidbruck (S. Tirol); Wien; Winden (Burgenland). — Hungary: a

series of 42 3 5 © with printed labels ‘“Särszentmihäly Bird 1923 v.1927” and “Dabas 1855, leg.

Metelka”, all 5 9 with identification labels M. luctuosa, det. M. Móczar & Mocsáry et Moczar

(MBUD, ML); 49, Simontornya, 8.v.1938, Pillich (NMB). — 9; Thebner Kogel; Budapest; Hungaria

centr.; Neusiedlersee, E-side (NMW). —Italy: ¢ 9: Cortona (Toscana), 20.v.1975, H. Teunissen (CT,

ML); M. Cimius (Lazio), 600 m, 18.iv.1949, M. Comba (ML); M. Sibillini (Marche), 2000 m, Monte

Sibilla, 26.vi.1930, G. Binaghi (IEP). — 9: Corfino di Garfagnana estata (Piemonte), 1954, Wiering

(MA); Aosta, Val Pelline, 22.v.1957, Bischoff (MNB) and Torino (Piemonte), Gianelli, coll. P. Magretti

(MCG) and Torino, 27.v.1918 (CKW); 2 ©, Genova (Liguria), under white drawer label “M. notata

Klug &Q (both are 9!) — punctata Lep., Gênes”, together with © M. a. nigra Spinola! (MT); Varazze

(Liguria), v.1917, F. Invrea (MCG); 1 9 (diss.), Bordighera (Imperia), 1.iv.1961, W. Grünwaldt (ex

CVS, ML); Appiano (?), Egat, Monticolo, v.1932, “lehmiger, bewachsener Abhang” (ML); Oltr’ Adige

(Trentino), Bolzano, Montioggi, 11.vi.1930 (ML); Bozen, v-vi, A. Weis (SMF) & same loc. (NMW);

Cortina d'Ampezzo, Mann leg. (NMW); Ponte Ticino (Lombardia), 13—28.v.1934, D. Prestifilippe

(IEP); Udine (Venezia), Val Rio del Lago Predil, 950 m, viii.1958, Br. Theowald (ML); Lido Venezia,

Mura, 10.v.1930, M. armata, det. Biegeleben (IEP); Bologna (Emilia), r.Pr.Fiori, M. Gibbio, 11.v.1893

(MCG); Lagopesole (Basilicata), 1.vii.1895, M. armata Pz., det.? (MT); Aspromonte (Calabria),

Paganetti (MBUD) and “Calabria. Aspromon Paganetti 1905” (NMW); 3 © (diss., much worn),

Aspromonte, 1350 m, 1.vii.1973, H. Bytinski-Salz (CBS). — Sicily: 49, Sicily, Zafferana, 800 m,

25.iv.1965, K. M. Guichard (BM); © (diss), Vittoria, 170 m, 3.vi.1908, G. Mantero, coll. Gribodo

(MCG); 3 9 (diss.), Mte Etna vers. occ. Pineta, 1700 m, 8.vi.1949, Hartig leg. (INER, ML); 9, Sicilia,

Südhang Aetna, vi-vii.1949, H. Hamann (CMS). — Yugoslavia: g Dobrovnik (Slovenia), 1.vi.1930

(NMW); Istria and Istrien, coll. Graeffe (NMW); Brioni Is. (NMW); Split (Spalato) (NMW). 9

Dalmatia & Arba in Dalmatia (NMW); Croatie (Hrvatska), coll. E. André 1914 (MP); prov. Kosovo,

Kosovska Mitrovica, Akad. Balkan Exped. (MBUD); Katlanovska Banja (Macedonia), 6.vi.1965, W.

Vervoort, and between Ohrid & Resen, 19.vi.1965, C. van Heyningen (ML). — Romania: 4, Mehadia,

Mann 1859, M. luctuosa, det. Friese “Type” (NMW); Tultscha (E Romania, Tulcea). — Bulgaria: g 9:

loc. illegible, 1.vi.1952 & v.1955, M. luctuosa, det. M. Kocourek (CK); Arkutino (Black Sea coast),

19.vi—4.vii.1970, K.Bleyl (CFP).

— 9: N. Zagora, 21.vi.1963, S. Niedl (CK); SW Bulgaria, Sandanski steppe, 26—31.v.1967, M.

Kocourek (CK). — Albania: 29, Albania Exped., Kula Ljums, Kruma & Hodzha nr. Prizren (NMW).

— Greece: No males! Females almost certainly belonging to /uctuosa are labelled as from the following

localities:Graecia, Ikaria, leg. Werner (NMW); Corfu, Athen and Delphi (NMW); Attiki, Neo

Peramos, 30.iv.1932, Dr. Fodor (MBUD); Peloponnesus, Vytina W of Tripolis, 22.v.1962, W.

Linsenmaier (CL); Olympia, 21.iv.1964, W. Grünwaldt, and Trypi, 19.v.1973, W. Grosz (CWG);

Karadagh, 30.v.1924, auf Euphorbia, and Umgeb. Tokluk (near Turkish frontier), 21.v.1924 (NMW). —

Turkey: ¢9: Turkey, Erzurum, 31.v.1972, H. Özbek, at Anchusa (CKM, ML); Turkey, Madensehir/

Konya, 20.vi.1973, K.Warncke (CKW). — 9: Erzurum, 18.vi.1965, 9.viii.1966, 28.v.1970 & 15.vii.1971,

H. Ozbek (CKW, ML) and Erzurum, Ispir, 17.vi.1973, K. Warncke (CKW); Ankara, 25.iii & v.1934, A.

Seitz (SMF) and Ankara, 3.vi.1972, K. Warncke (CKW); Türkei, Giglikara, 1860 m, N. N.

Zedernwalde, 30.v.1966, H. Felten, M. luctuosa, det. D. S. Peters (SMF); Ciftahan, and Posanti nr.

Ulukishla (Taurus), v.1955, Seidenstucker (CKW, ML); Taurus, Akseli, 1300 m, 13.iv.1974, K.

Warncke (CKW); Turkey, Halfeti/Urfa, Insesu/Kayseri, Ergek/Van, and Agri, N. Süphan Dagi, iv-

vi.1976—77, all K. Warncke (CKW). — USSR: £ (4 ex., one diss., figs. 88—90), Uralsk, Bartl (print),

one with “M. luctuosa aggr. sp. thyridia variable”, det. D. B. Bakker (NMW); & 9, Ukraine, Kievskaja

dist., Christinovka, distr. Umanj. gub. Kiev, 12.v.1901, I. Zhicharev leg., M. luctuosa, det. Osychniuk

(CO); 2 3 1 ©, Chersonskaja dist, Novaja Tjaginka and vicinity of Cherson, 14 & 19.v.1954, M.

luctuosa leg. & det. A. Z. Osychniuk (CO, ML); 1 £, Ross. mer., Uschakoff (MH); 2 4, Russia mer.,

with two partly illegible labels in Russian ‘“‘Woro Zaljogwitsj, Galitscha Gora, Kutschark, 15.v &

22.v.1938, A. Golidche” (2) (MNB); 9, Caucasus, Araxesthal, Leder-Reitter, M. luctuosa var., det.

Friese (MNW); 1 ©, Irkutsk, 22.iv—12.v.1904 (MNB); 1 g 1 ©, Irkutsk, 14—27.v.1913, O. Hesse

(MNB); & (diss., ex alcohol), Jelan, Region du Baikal et env. d’Irkutsk, Paul Labbé 1902 (MP); ¢ 9,

“Dschungarey, Post 1 1878” (Dzhungarskiye, SE Kazakhstan) (NMW); 1 ©, Siber (ia), M. luctuosa,

232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Figs. 81—82. M. luctuosa; 81, ventral view of tergite 7 (holotype 8 M. eczmiadzini, Caucasus); 82, ster-

nites 7 and 8 (same specimen). — Figs. 83—94. M. luctuosa; 83, oblique dorsal view of right hind tarsal

claws (g © Hulshorst, Netherlands); 84, ventral view of tergite 7 (4 same locality); 85, sternites 7 and 8

(same specimen); 86, right gonostylus, oblique extero-lateral view, with separate dorsobasal process of

left gonostylus (same specimen); 87, dorsal view of pygidial plate (9 Sierre, Switzerland); 88, ventral

view of tergite 7 (4 Ural); 89, sternites 7 and 8 (same specimen); 90, genital capsule, ventral and dorsal

halves (same specimen); 91, ventral view of tergite 7 (¢ Iran); 92, sternites 7 and 8 (same specimen); 93,

exterior view of left gonostylus (4 Moravia); 94, dorsal view of pygidial plate (@ Bonn, W Germany)

LIEFTINCK: Palaearctic Melecta 233

det.? (MBUD); 9, Transkauk (asia), Helenendorf 1880, M. luctuosa, det. Kohl (NMW); 4 9,

Samarkand, Aman Kutan, 1—2.vi.1919, J. Niedl (CK); Usbekistan, Samarkand, 12.vi.1957 (MNB); 2 3,

Ost-Turkestan, Narin, E. A. Böttcher (MNB). — Iran: 2 & (diss., figs. 80, 91—92), Iran, Karadj Lake/

Keredj river/, 2000 m, ca. 30 km W Teheran, 24.v.1972, H. Bytinski-Salz (CBS, ML).

Synonymy. — Concerning the questionable synonymy cited above, any

comment would seem too speculative to be of any use. M. fasciculata Fischer-

Waldheim is preoccupied by fasciculata Spinola (= albifrons nigra Spinola),

eversmanni is most likely a synonym (see species incertae sedis), while eczmiadzini

and /uctuosa are undoubtedly also conspecific.

Distribution. — It will be seen, that several references cited above under

luctuosa are affixed ““pars’”’, which means that more than one species may be

involved. Therefore, all published information on this species based on material

originating from the more southern and eastern parts of its range, should be

considered with caution. For instance, Radoszkowski (1893), after briefly

describing the pubescent pattern, mentions the following localities taken from

specimens in his own collection: “Astrakhan (Ryn-Peski), Odessa, Ukraine,

Siberie, France, Italie, Kissingen”. It must, of course, remain uncertain how many

— and if any — of the author’s “‘/uctuosa’’ are really that species! As to the

material examined by myself, and reviewing all reliable data, it must be admitted

that, as long as no authentic males have become known, the occurrence of /uctuosa

remains to be established in the following countries (arranged alphabetically): —

Balearic Is., Corsica, Crete, Cyprus, Egeian Is., Iraq, Israel, Jordan, Lebanon,

Malta I., whole North Africa (incl. Egypt!), Rodos I., Sardinia. In many of these

countries and islands the species is likely to turn up sooner or later; but, by the

absence of males, many solitary females supposed to be luctuosa, mainly from

Asiatic Turkey, had to be left unidentified.

Note on the occurrence of M. luctuosa in the Netherlands.

In former days, from 1938 to 1942 (and once again in 1947), the late biologist F.

C. Mijnssen regularly observed /uctuosa in May and June inspecting the nesting

site of Anthophora retusa (L.), on the S-slope of a railway cutting near the station at

Baarn (prov. of Utrecht). Though no definite evidence of parasitization could be

acquired, the Melecta always kept company with the retusa bees, which

undoubtedly acted as host of /uctuosa in this particular place. While collecting

together at the same locality on May 15, 1938, we observed the two species jointly

gathering nectar at flowers of Lamium album, Lithospermum officinale, and Nepeta

cataria.

Melecta rutenica Radoszkowski

(figs. 97—98, map 1, p. 223)

Melecta luctuosa var. Rutenica Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3):

179— 181, fig. 30a—c, i(g genit.) (Q & “Cechocinek, Pologne” [err.!], see below).

Type material. — USSR: 1 4 (diss. & fig. by Radoszkowski), labelled “Nickon:

p. Stani:” (2) (poor writing, unknown hand), “Rutenica” (Radoszkowski’s

234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

handwriting), “Type” (print on red), “Melecta luctuosa var. rutenica Rad. Type”,

rev. Dr. Enderlein (MNB). Type designation by Enderlein confirmed and here

selected lectotype.

Further material. — USSR: 1 9, “Distr. Kiev, Kirill-ravine, 24.v.1917, V. Alexand.” (transl. from

written label), ‘M. luctuosa v. rutenica Rad. ©”, det. A. Z. Osychniuk (CO). — Turkey (Asia minor): 1

3 (diss., figs. 97—98), “Mann Brussa 10” (written), “167” (ditto), ““Melecta luctuosa var. 3, det. Friese

1893” (NMW).

The identity and habitat of this species are of some historical interest,

warranting the following observations.

Radoszkowski’s diagnosis of the two sexes runs as follows:

“A Cechocinek, en Pologne, il y a une varieté que je nomme: Var. Rutenica.

Poils de la tête et du thorax complètement noirs, excepté la touffe blanche du

chaperon. Le male de cette variete (il a été pris avec sa femelle a la même place)

est garni de poils gris sur le prothorax [i.e., anterior portion of mesonotum] et de

chaque côté du métathorax. Dans ma collection, je possède aussi un exemplaire

[sex not stated!] de cette localité avec le chaperon et les pieds sans taches

blanches”. Follows a description of the Z genitalia of which sketches are also

Figs. 95—96. M. luctuosa (Q Hungary); 95, antennal segments 3—4; 96, left scutellar tubercle, intero-

dorsal view. — Figs. 97—98. M. rutenica (3 Brussa, Turkey); 97, ventral view of tergite 7; 98, sternites

7—8. — Figs. 99—102. M. brevipila(3 holotype and 9 paratype, Almasy, Turkestan); 99, dorsal view of

3 tergite 7; 100, sternites 7 and 8 (same specimen); 101, external view of left gonostylus (same speci-

men); 102, dorsal and left lateral view of 9 pygidial plate (paratopotype)

LIEFTINCK: Palaearctic Melecta 235

given. As these organs had already been figured earlier by the same author (1886)

for a supposed luctuosa from Ashkhabad, the drawings accompanying Radosz-

kowski’s 1893 description are clearly those of this rutenica. These figures are not

alike, but too schematic to be of any use.

Concerning the habitat of this taxon, Dr. W. J. Pulawski, of the Zoological

Institute, Wroclaw, kindly gave me important information, now several years ago.

He wrote to me as follows: “The historical term Ruthenia is more or less the same

as the present day Ukraine. The most western part of it is the Lvov district, which

since 1939 is part of the Soviet Union. Therefore, M. ruthenica Rad. certainly does

not belong to the Polish fauna. It is more probable that this bee was described

from specimens from the Kiev area or even the Black Sea coast. I have no idea

about the type location.”

Male. — As the Brussa specimen agrees in every respect with the lectotype,

found in the Berlin museum and identified by Radoszkowski, we are now

somewhat better informed about the occurrence of rutenica. With luctuosa and

brevipila it is the only species of which the male can be recognised by the absence

of well developed impressed antennal sensoria. It can be distinguished from the

former and most other Melecta by the reduction of white marks, small size, fairly

long black pile, and relatively strong antennae. The genital capsule of the type has

been cut to pieces, but the sternal plates are still intact and do not differ from

those here figured for the second specimen (fig. 98). Seeing how variable these

structures are in its near ally, /uctuosa, the differences shown are almost negligible

and would point to their close relationship.

Female. — The dark specimen from the Ukraine (Kiev area), is the only one

that can be assigned to rutenica with reasonable certainty. Of course, more

material of both sexes, preferably from one and the same locality, is needed to

confirm the correctness of the present sex association. Here follow a few more

characters of the specimen in hand, in addition to those given in the key.

Antennal segments 3—12 only little longer than broad, 3 less than twice as long

as its breadth at apex and less than one and one-third longer than next segments,

the intermediate joints almost square. Punctures on disk of mesonotum partly

smaller than rather shiny interspaces; median mesonotal and parapsidal lines fine,

not impressed. Scutellar tubercles moderate, parallel, conical, raised obliquely

upward and backward, shorter than surrounding pubescence. Third submarginal

cell in fore wing distinctly higher than long (24: 20), its distal side moderately

angled. Inner ramus of mid and hind tarsal claw about one-third length of outer.

Abdominal tergites markedly shiny, setiferous punctures smaller than interspaces.

Little could be found to distinguish rutenica from our series of leucorhyncha

taormina, i.e., the subspecies commonly distributed all over Italy and some of the

islands in the Mediterranean. It is held distinct from that taxon on account of the

non-developed antennal rhinaria and longer hair.

Melecta brevipila spec. nov.

(figs. 99—102, pl. 2 fig. 7)

Type material. — USSR: 1 3 (diss., figs. 99— 102, pl. 2 fig. 7), with two printed

236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

labels “Vallis Kabak” and “Turkestan, Almasy 1906”. Holotype (MBUD).

Further material. — USSR: 2 & 1 9 (diss., figs.), all with same labels as holotype (MBUD, ML). —

Turkey (Asia minor): 1 & (diss., much worn, wings frayed), Asia minor, Umgebung Konya, 4.vi.1964,

H. H. F. Hamann (CMS).

The rather comprehensive diagnosis of this remarkable new species, found in

the descriptive key to the males, is given on purpose for comparison with that of

luctuosa, mainly because they are almost unique by the absence of antennal

rhinaria. For that reason alone the two species run out together in the same

paragraph of the key. There are, however, no further points of agreement between

them, and they do not seem to be at all closely related.

Wing expanse of male holotype (pl. 2 fig. 7), 27 mm.

Here follow a few more characters not mentioned in the key.

Male. — Labrum subrectangular, distinctly though slightly longer than broad,

surface at first convex, then markedly hollowed out, the anterior border finally

again upturned; side angles rounded but apex distinctly projecting, bearing a

subacute median denticle, the latter in two paratypes prolonged basad for a short

distance, forming an indistinct median ridge; surface deeply, irregularly punctate.

Maxillary palpus slender, 6-segmented, the last joint very short. Silvery white

patch upon clypeus squarish, the long hairs straight, lying flush upon surface, tips

not or scarcely exceeding anterior border; sides black. No distinct smooth areas in

front of median and just beside lateral ocelli, surface dullish; distance separating

ocelli little shorter than their own diameter. Head and thorax closely, almost

contiguously punctate; median mesonotal line not impressed. Wings subhyaline,

but fore wing membrane gradually somewhat infuscated beyond cells; third

submarginal cell shaped much as in many luctuosa, with distal side rather strongly

angled. Subbasal white spots at outer faces of mid and hind tibiae and at sides of

tergites 2—5 characteristically small and isolated.

Female (unique). — An aged specimen, with much of the body pubescence

rubbed off and with fore wing borders revelled out. Probably conspecific, agreeing

with the male in many respects and bearing identical printed locality labels.

Chiefly characterized by predominantly black head and thorax (including sides

of the latter), dullish abdominal tergites, caused by relatively more closely set

setiferous punctures than in similarly looking species of equal size. Flagellar

segments of antenna also more markedly longer than broad than usual. Third

submarginal cell much higher than long, the distal sides of second and third cells

moderately angled but more nearly parallel than in most other species of the same

sex. Length 11.3 mm approx. Pygidial plate, fig. 102.

Melecta sibirica Radoszkowski

(figs. 103—111)

Melecta sibirica Radoszkowski, 1890, Horae Soc. Ent. Ross. 25 (1—2): 246—247 (9 Irkutsk, feu Ec-

kert).

? Melecta sibirica Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 184(9 Irkutsk).

LIEFTINCK: Palaearctic Melecta 2817

Figs. 103—111. M. sibirica; 103, posterior view of antenna (3 Ramit, Tadzhikistan); 104, dorsal view of

tergite 7 (same specimen); 105, sternites 7 and 8 (same specimen); 106, apices of sternite 8 (upper) and 7

(lower) (second specimen); 107, external view of left gonostylus (same specimen); 108, partial dorsal

view of gonocoxal angle (+), with dorsobasal process (same specimen); 109, apex of sternite 7 (left) and

whole sternite 8 (right) (4 Akfasch/Tashkent) and 110, the same of second specimen from same locali-

ty; 111, external view of left gonostylus (3 Akfasch/Tashkent). — Fig. 112, M. eversmanni, dorsal view

of pygidial plate (@ lectotype, Orenbourg). — Fig. 113, M. ashabadensis, dorsal view of pygidial plate

(Q lectotype, Ashabad). — Figs. 114—117, M. turkestanica (3 lectotype, Tashkent); 114, ventral view

of tergite 7; 115, sternites 7 and 8; 116, ventral view of right half genital capsule; 117, external view of

left gonostylus

238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

“Type” (print on orange), “sibirica” (pencil writing, Radoszkowski’s hand),

‘“Melecta sibirica Radosz. Type, rev. Dr. Enderlein” (MNB). Holotype designation

by Enderlein, confirmed.

Further material. — USSR: NW Tadzhikistan: 2 & (both diss., figs. 103—108, one with frayed

wings), with written labels ““Tadcikistan, Ramit [NE of Dushanbe], 1.vi.1966, Niedl’’ (CK), and “Tadz

[Fadrobod?], 7.vi.1966, Niedl (CK); 2 & (both diss., figs. 109—111), with written labels “Turkestan,

Akfasch, 60 km NNO Tashkent”, 1500 m, 15.v.1974, at Eremurus, M. Kraus (CKW, ML).

Doubtful specimens. — USSR: | Q (diss.), Turkestan: Bokhara, Tschardschui 1913, H. Veth (BM

1925—553); 1 Q (diss.), with four labels: green square/yellow disk with 2206/8 on reverse side/Museum

Paris/Turkestan/Capus Bonvalot/Melecta (MP); 2 Q (diss.), Turkestan, Osch 1905—Korb (MBUD,

ML). — Afghanistan: 1 9 (diss.), NO. Afghanistan 1953, Nuristan, 2200 m, Kamdesh, 28.iv.1953, J.

Klapperich (MBUD).

This is the first of a small series of puzzling taxa, described consecutively by

Radoszkowski at the close of the last century. M. sibirica is the earliest species in

this group, characterized in 1890, and for that reason deserves particular attention.

Three others were described three years later and are being discussed in the next

pages. For M. eczmiadzini Rad., see under /uctuosa; the identity of M. rutenica Rad.

has been established elsewhere in the present article.

As to M. sibirica, it will follow from the above references, that the author

possibly described two species under the same name, both originating from

Irkutsk and from a comparison of the diagnoses it is not at all clear whether

Radoszkowski had the same specimen before him at the time of describing them.

The present individual, fixed by Enderlein as the type, corresponds with the very

brief 1893 description, but not exactly with the earlier and more fully described

sibirica. For the sake of clearness, both characterizations are here given in full.

1890. ““Melecta sibirica n.sp. Nigra; capite thoraceque niveo-variegatis;

abdomine subopaco, segmentis 1—4 niveo-maculatis; alis subfuscis; scutello

bidentato. Long. 9 11—12 mm. Regue de feu Eckert des environs d’Irkutsk.

Femelle. Noire. Téte garnie de poils noirs; le chaperon porte une touffe; on voit

autour des antennes deux taches et derrière la tête une lisiere de poils blancs de

neige. Thorax densement ponctué, sa partie antérieure garnie de poils longs

blanchâtres; sous les ailes les touffes sont formées de ces mêmes poils; on voit des

taches pareilles de chaque côté de l’écusson, qui est bidente. Abdomen nu,

presque opaque; sur la base du premier segment on voit quelques poils gris; les

deux premiers segments portent de chaque côté une petite tache blanche et le

troisième en porte deux de chaque côté (spacings are mine), qui sont éloignées

du côté, le quatrième porte deux taches rondes; toutes ces taches sont formées de

blancs couchés; ventre nu. Pieds garnis de poils noirs; les jambes portent des

taches blanches. Ailes faiblement enfumées.”

1893. ‘M. sibirica n.sp. Femelle. Par sa stature se rapproche du Pseudomelecta

diacantha; abdomen plus long et moins large que celui du M. luctuosa, avec lequel

elle a beaucoup de ressemblance par la disposition des poils blanchâtres sur la tête

et sur le thorax, et par la petitesse des dents de l’écusson. Taches des quatre

premiers segments abdominaux sont d'un blanc de neige, petites, non allongées, et

garnissent les bords postérieurs des segments; sur les deux premiers segments ces

LIEFTINCK: Palaearctic Melecta 239

taches sont disposées très près des côtés, sur les segments suivants, elles en sont

assez éloignées. Long. 13 mm. Irkoutsk.”’

But for the absence of paired white spots on each side of tergite 3 (a surprising

statement!), the present individual also fits Radoszkowski’s first description of

sibirica. But since the whereabouts of the last described female — if there really

were two specimens — remained unknown, I have accepted the author’s own type

selection as the correct one, there being no other specimens named as such in the

Berlin museum.

As stated in the 1893 description, this bee, by the dullness of its finely and rather

closely punctate abdominal tergites, somewhat recalls Pseudomelecta diacantha

(Eversmann). The white tergal spots consist of very short, fine and closely set,

appressed feathery hairs, those covering the sides at base of tergite 1 are, however,

erect, tufty and much longer. It is undoubtedly a true Melecta, agreeing fairly

closely with /uctuosa, but differing from this by having a distinctly less shiny, more

closely punctate abdomen (see above); also by the placement of the white spots,

which on tergites 3 and 4 are situated more inward, removed from the bend by a

distance almost equal to their own diameter. The neuration is that of luctuosa,

while the antennae, tarsal claws and other parts of the body are not different in

shape from /uctuosa. Only the apex of the pygidial plate is exposed and this also is

shaped similarly to that of the latter. Length 12.3 mm approx., fore wing 9.0 mm.

Of all species hitherto known, the males here associated with sibirica resemble

M. turkestanica Radoszk. most closely. The latter averages larger in size but is,

nevertheless, probably related. I have little to add to the key characters as given in

comparison with the male of that species. M. sibirica can be at once distinguished

from turkestanica by the shorter pile, more slender form of the hind basitarsus, and

the flat, much smaller white abdominal spots. These males are, however, also very

similar in size, general appearance and abdominal spots to certain bees here

assigned to M. leucorhyncha (itself a very variable species), especially to

populations from the Ukraine, eastern Mediterranean islands and Turkey, sharing

with these also the almost colourless wing membrane. The two males from Ramit

(figs. 103—108) and those from Akfasch (figs. 109—111) are also reminiscent of

our new species megaera, but apart from its superior size and details of structure,

the latter can be recognized by having a more even, less fluffy, vestiture and

slightly darker wings.

I am unable to distinguish the supposed female of sibirica from those of brevipila

spec. nov., and certain populations of leucorhyncha. Lastly, in comparison with the

type of M. eversmanni Radoszk. (fig. 122), the pygidial plate of sibirica is broader at

base and more suddenly narrowed apically, the swollen and bluntly carinated tip

being a little downbent.

Melecta turkestanica Radoszkowski

(figs. 114—122, pl. 2 figs. 8— 10)

Melecta turkestanica Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 181—182, fig.

34 a-c, i (& genit., 2g Tachkend (Boc-cy). — Popov, 1967, Trans. Zool. Inst. Akad. Nauk Lenin-

grad 43: 207 (¢ Iran, no descr.).

240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Type material. — USSR: E Uzbekistan: 1 G (diss., figs. 114—117) pl. 2 fig. 8,

labelled “Boc-cy 23 Mrs” (written), “Tach ken” (print), “Turkestanica”

(Radoszkowski’s writing), “Type” (print on orange), ““Melecta turcestanica Radosz.

Type”, rev. Dr. Enderlein. Holotype, evidently selected by Enderlein (MNB),

confirmed.

Further material. — USSR (Turkestan) or Afghanistan: 1 g 1 9 (diss, figs. 118—122 & pl. 2 figs.

9-10), both with small purple cadre (= captured in April), a small written label “Alai’’, and the usual

printed museum labels “Muséum Paris coll. J. Pérez 1915” (MP).

The two males before me are from different localities and therefore described

separately. The Alai male differs from the type in that the long white scutellar

hairs are not restricted to just behind the spines but surrounding (and concealing)

them at all sides; the white mid tibial pad is better defined and the hind basitarsus

is a little more outbent. In spite of these differences I believe the two to be

conspecific. To enable future recognition both are here photographed (pl. 2 figs.

9—10), the Alai specimen along with its female, which undoubtedly belongs to the

same species. The key characters are partly based also on the larger-sized male

from Alai.

Here follows first a brief description of the unique type and only authentic

example. I have not seen the specimen from Iran recorded by Popov, which may

or may not be conspecific.

Male (holotype). — Labrum hidden from view, apparently slightly longer than

broad, with long erect blackish bristles. Mandibles black. Maxillary palpi 7-

segmented (sic), 2—5 subequal in length and longer than | and 7. Pubescence on

disk of clypeus long, decumbent, silky white; hairs alongside also long, black.

Antenna thick, scape clothed with very long white hairs; 3 subequally long to 4,

both a trifle longer than broad, the flagellar segments almost square; rhinaria

deeply impressed, present on 3—13, horseshoe-shaped, but oval on 3, circular on

13. Head above and aside strongly, contiguously punctate; long raised white hairs

not very dense, becoming black laterally, except white at temples. Posterior part

of mesonotum black, thorax for the rest entirely white, hairs mostly concealing the

surface. Scutellar spines rather long, conical, pointed, with tufts of very long white

hairs just behind each of them. Fore and hind femora black, the posterior fringe at

fore femur longer than its diameter, tuft fringing mid femora posteriorly very long

and white, that on hinder pair much shorter and more compact. Tibiae closely

punctate; white external pad of mid tibia isolated but not very conspicuous, much

longer than broad, lacking thin fringe of longish hairs posteriorly; outer face of

hind tibia for the greater part clothed with short white hairs but devoid of spine-

like black setae. Hind basitarsus not markedly concave externally, broadest at

about halfway length, the outer border straight, the inner (lower) distinctly

outcurved (convex). Tarsi yellow-brown; inner rami of all tarsal claws very slender

and but little shorter than outer. Wings almost hyaline, neuration brownish yellow.

Abdominal tergites rather shining, sparsely superficially punctate inclusive of

the hind margins of 1—S; sternites similar, pubescence not very dense, consisting

of short, black decumbent hairs which are longest on posterior sternites. Tergite 1

clothed with very long raised white hairs, most densely so, longest and tufty, at the

LIEFTINCK: Palaearctic Melecta 241

sides; pile on dorsum of 2—5 short, black, sparsely intermixed with longish hairs at

sides of tergites, the white pubescent spots of regular shape. Tergite 7 rapidly

tapered with slightly concave sides, hind margin emarginate; dorsal surface

smooth and shining, dark reddish brown, except the apex which is colourless, the

tips of the side angles very slightly upturned (fig. 114, ventral view). Hind margin

of sternite 6 little prominent, rounded, median impressed area shallow, suboval,

lighter in colour and less hairy than the rest.

Parts of the extruded genital organs were figured by Radoszkowski, but could be

further dissected out and examined. They are here redrawn, together with

sketches of tergite 7 and the apical sternal plates. It must be emphasized that the

drawings and description of the gonostyli (“branche du forceps conique — non

arrondie, mais pointue a l’extrémité — avec sa partie postérieure richement garnie

de longs poils”) are not quite correct, the tip of the left gonostylus being more

rounded than that of the right one, as shown in fig. 117.

Total length 12.0 mm approx., wing expanse 24.3 mm.

Male (Alai, pl. 2 fig. 9). — Labrum of the usual shape, slightly longer than broad,

Figs. 118—122. M. turkestanica from Alai; 118, dorsal view of ¢ tergite 7; 119, ventral view of same (sa-

me specimen); 120, sternites 7 and 8 (same specimen); 121, partial ventral and dorsal views of genital

capsule (same specimen); 122, dorsal view of 9 pygidial plate. — Figs. 123—126. M. megaera; 123, ex-

ternal view of right hind basitarsus (¢ holotype, Polemedia, Cyprus); 124, dorsal view of tergite 7 (same

specimen); 125, lateral view of right gonostylus (same specimen); 126, dorsal view of pygidial plate (9

Polemedia Hills, Cyprus)

242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

closely coarsely punctate lacking a definite median ridge, anterior margin broadly

rounded. Entire dorsal surface of head and thoracic sclerites dull, very closely,

contiguously punctate. Antenna strong, scape fringed all around with long raised

white hairs, the apical ones characteristically recurved; 3 about one and one-

fourth as long as its apical breadth (100 :76), somewhat longer than next ones,

which are subequally long. Silky white patch covering clypeus broad, thick and

conspicuous, all hairs long and straight, lying flush with surface and even

somewhat downcurved, tips projecting well beyond anterior border; paraocular

bristles deep black. Thorax, including sides of propodeum and ventral sclerites

clothed with long, fluffy white pubescence, only metapleurae posteriorly, a poorly

indicated T-shaped median area traversing scutellum anteriorly, and a narrower

zone (representing short stem of T) between scutellar spines, black.; long white

tufts behind wings confluent with still longer and curly hairs surrounding scutellar

spines, the latter much shorter, triangular, acuminate. Wings entirely hyaline,

veins yellowish brown. White pad of mid tibia well defined, elongate, covering

outer face from near base to near apex. Hind femur throughout closely punctate;

pubescence black, moderately long raised hairs fringing posterior ridge much

shorter than diameter of femur. Outer face of hind tibia white on basal half, for the

rest black; few short black spicules between punctures on somewhat shiny ground.

Hind basitarsus slightly but distinctly outcurved, broadest at about midway length

(length-breadth ratio 100 :37), both upper and lower borders convex, the former

lacking fringe of long raised bristles (profile!); hairless ventrobasal pit-like

concavity to the inside of lower ridge distinct, elongate-oval, shiny and punctate;

mid and hind basitarsi white externally, except at extreme base, the remaining

tarsal segments being all white; claws reddish.

Tergal plate 7 rather narrow, tapering, disk a little hollowed out, punctures

largest and setiferous only on basal portion, lateral ridges poorly indicated, a little

swollen and hairy with well-marked crescentic emargination, the rounded tips

yellow-brown, slightly upturned, impunctate. Sternite 6 with very low lateral

tubercles; 7 and 8 as in fig. 120. Genital capsule relatively small, 1.8 mm long (incl.

gonostylus), gonocoxal angle completely rounded, feebly bi-angulate, course of

diverging borders thereafter straight (fig. 121).

Size larger, length 16 mm, fore wing 13.2 mm, wing expanse 28.5 mm.

Female (Alai, pl. 2 fig. 10). — Undoubtedly conspecific with the last described

male. There are few characters additional to those given in the key which may help

to distinguish it from other regional species. M. turkestanica is more stoutly built

than sibirica, with much lighter wings and larger, more regularly arranged,

subrectangular tergal spots, while the body pubescence is longer throughout.

Though similar in general appearance to the eastern Mediterranean megaera,

these two species are probably not at all closely related, the females especially

differing so much in the shape of the pygidial plates that megaera can not possibly

be confounded with other large-sized species (cf. figs. 122 & 126).

Length 15.6 mm, fore wing 12.3 mm.

LIEFTINCK: Palaearctic Melecta 243

Melecta megaera spec. nov.

(figs. 123— 126)

Type material. — Cyprus: | & (diss., figs. 123—125), labelled “Polemedia,

9.3.50, No. 5” in G. A. Mavromoustakis’ handwriting. The specimen is the

holotype (MNB).

Further material. — Cyprus: 1 ¢ (diss.), same label as holotype, No. 4 (ML); 1 9 (diss., fig. 126),

Cyprus, “P. Hills, 14.3.50, No. 3” (do.) (MNB); 1 9 (diss.), Limassol, Yermasoyia, 13.3.1979 (2 days

later than 1 9 M. tuberculata sp.n.!), L. A. Janzon (NRS). — Rodos I. (Greece): 1 9 (diss.), Rhodus,

Hedenb. (written), “1904 Friese det.” (print, but no name) (NRS); 4 9 (all diss.), Rhodos, Monte

Smith, 21—25.3.78, K. J. Hedquist (NRS, ML). — Crete: 1 9 (diss.), Kreta, Omalos Ebene, 26.iv.1942,

— Greece: 1 Q (diss.), Chalkis, Euboea, iv.1926, Holtz (MNB). — Turkey (Asia minor): 1 9 (diss.),

Kleinasien, Sewdiköib b. Smyrna, 4—19.iv.1917, La Baume (MNB); 2 ¢ (diss.), W. Türkei, Ayvalik,

18.iv.1965, Kl. Warncke (CKW, ML); 1 & 1 9 (both diss.), Türkei, Side, 30.iii.1972, Kl. Warncke

(CKW); 1 9 (diss.), Aksehir, Türkei, 4.v.1960, Rassl, no. 364 (ex CP, CMS); 1 9 (diss.), Smyrna, coll. C.

& O. Vogt acq. 1960 (MA); 1 9 (diss.), Exp. Turkey Mus. Leiden, East Turkey, 2150 m, Erzincan, 20

km N, v.1959 (ML). — Israel: 1 9 (diss.), Palestine, Tiberias, 17.iii.1946, H. Bytinski-Salz (CBS); 1 9

(diss.), Israel, C. Jordan Valley, Deganya A, no. 6333, 27.ii.1939, Y. Palmoni, M. aegyptiaca Rad.? det. J.

D. Alfken 1937, and M. luctuosa Scop. var. (unknown hand) (AID); 1 9 (diss., much worn), Negev Mts.,

11.iii.1935, coll. Hecht (CBS). — Iran: 1 9 (diss.), yellow disk, with 16/40 on reverse side/Museum

Paris/Perse/Aucher 16—40 (MP).

albovaria Er. (unknown hand) (MG). — Central Asia: | Q (diss.), Alai (written), Museum Paris, coll. J.

Russland, with two written labels ‘‘Sarepta, S. Russland, Stein” & ‘‘Sarepta” (together with equally

labelled Q having quite differently shaped pygidial plate! (MNB). One of the other questionable

females labelled ‘‘Alai’’ (MP), is a species totally different from the one attributed to turkestanica (pl. 2

fig. 10), bearing an exactly identical locality label. In the shape of its pygidial plate it comes very near

megaera.

The male of this new species most closely resembles turkestanica (pl. 2 fig. 9) in

general appearance and pubescent pattern, but these two do not seem to be

closely allied (see descriptive key). Both are also rather like baeri and candida (pl. 3

fig. 18); apart from structural differences, one of the features most helpful to

separate these bees from megaera, is found in the relative hair lengths of all body

parts.

The female of megaera runs out in the key to near aegyptiaca, with which it

seems, indeed, more nearly related than with the two just mentioned for

comparison with the male.

As in aegyptiaca and the eastern turkestanica, the long white tufts behind the

wings are more or less confluent with those surrounding the scutellar tubercles,

leaving only a narrow black-haired space upon middle of scutellum, thus differing

from assimilis and most large-sized individuals of italica. Typical males of the latter

(from Italy and France) have the middle of scutellum almost wholly black; these

italica also have much stronger antennae and a deeply emarginate 7th tergal plate.

244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Melecta aegyptiaca Radoszkowski

(figs. 11, 127—145, pl. 4 figs. 19—21, map 1, p. 223)

Egypt). Radoszkowski, 1893 (sub egyptiaca, identity uncertain), Bull. Soc. Imp. Nat. Moscou, new

ser. 7 (3): 182, fig. 35 a-c, i, k (& genit., sine loc.). — Friese, 1895, Bienen Europa’s: 165—166 (pars ?

orig. descr. copied; ¢ Persien, note). Friese, 1925, Konowia 4: 28 (pars !? “Ägypten; Beirut; Südspa-

nien; Algerien”).

Melecta lindbergi Lieftinck, 1958, Comment. Biol., Soc. Sci. Fenn. 18 : 25 (key), 28 fig. 21 (scutellum) &

39 (pygidium) (@ Lanzarote, Canary Is.). Syn. nov.

Type material. — Egypt: 1 9 (fig. 131), labelled “Egypt C:Bra” (print on green),

“Type” (orange), “‘aegyptiac’’ (Radoszkowski’s writing), ‘M. aegyptiaca Radosz.

loc. & type labels, syntype M. aegyptiaca (MNB). — Canary Is.: 1 9 (holotype

M. lindbergi Lieft., figs. 137—139), Lanzarote I., Haria, 19.iii.1949, H. Lindberg

(MH).

Further material (from W to E, roundabout Mediterranean & S Europe). — Morocco: 3 4 (one

diss., fig. 143), Maroc, 622—19 & 623—22 (MG); 3 & (one diss.), Tanger, Vaucher, Cn. Tournier (MG

(MP, ML); 1 ¢ 1 9 (¢ abdomen missing), 1 Sétif, coll. O. Sichel 1867, 9 with note “Mel. luctuosa, 3 Q

A. Seitz, Melecta spec.? det. Alfken (SMF). — Tunisia: 1 g, Tunis, coll. Schmiedeknecht,

ML). — Egypt: 1 & (diss., figs., 131, 135, 140), Cairo Aegypt, M. armata var. mediterranea Grib., det.?

(ZSM); 1 9, Cairo Aegypten, M. dasypyga, det.? ex coll. Univ. Groningen (ML); 3 & (pl. 4 fig. 19),

Schmiedeknecht, Egypten, coll. A. Weis, M. luctuosa Scop., det.? (SMF, ML); | 9, Egypt, Sezira-

Cairo, iv, H. Priesner (ex CP, ML); 1 & 1 9, Aegyptus, Cairo (MBUD); 1 9, Egitto, Asyut, 2.ii.1933,

(MNB); 1 9, Aegypten, Ehrenberg/XIII/725, M. grandis Lep., det.? (old handwriting, MNB); 1 9,

Alexandria, Winter 1902/03, Dr. Osborne, M. aegyptiaca Rad., det. J. D. Alfken, Slg. Alfken 1933

(MNB); 1 ¢ (diss.), Egypte, Melecta orientalis Egyp (on green disk), coll. Romand (MG). — Ethiopia:

1 9, Abyssinia, Dr. Rüppel, M. luctuosa Scop., det. H. Friese 1900 (SMF). — Israel: 1 g (diss.), Israel,

Jordan Valley, Zerga R. Colony, ca. 100 m below sea level, Mid Mar. 1952, “Bee + Bombilid” (sic),

Trevor Trought, Crocisa sp., det. I. H. H. Yarrow (BM). — Syria: 2 g, Syria 1899, Morice,

M. aegyptiaca, det. Friese 1904 (MNB). — Rodos (Greece): 3 ¢ (1 diss.), Rhodos-Kattavia & Gaduras

road crossing, 5.iv.1971, V.S. v.d. Goot (MA). — Greece (mainly continental): 3 & 2 9 (diss.), Corfu

(Kerkyra), Schmiedeknecht 1912, coll. A. Weis, one with M. luctuosa Scop., det. Schmiedeknecht

Griinwaldt (CWG, ML); 1 9, Graecia, Levadleia (Levadhia), 4.vi.1966, W. Linsenmaier (CL); 4 9,

Graecia, Peloponnesus, Alt Korinth, 11, 13 & 19.v.1964, Max. Schwarz (CMS, ML). — Hungary: 1 &,

Simontornya, Hung. occ., 24.v.1933, leg. Pillich, at Anchusa officinalis, M. albovaria, det. Alfken 1933,

“zur armata-Gruppe gehörend, sicher nicht M. /uctuosa”, Alfken (MBUD); 1 &, Simontornya, Hu.occ.,

(MBUD); 1 9, Dabas 1855, leg. Metelka, M. luctuosa, det. Mocsary & M. Möczär (MBUD); 3 g,

Apaj-p., Bokor E., M. luctuosa Scop., det. M. Möczär, Csömör, Ujhelyi leg., same identif., and Isaszeg,

Ujhelyi leg., same identif. (MBUD, ML). — Czechoslovakia: | g (diss., figs. 133—134, 145), Slovakia,

Parlzán 1933, A. Hoffer (CTP); 2 ¢ (one diss, (fig. 142, pl. 4 fig. 21), CSSR, Slovak, 2.vi.1965, Sturovo,

LIEFTINCK: Palaearctic Melecta 245

Figs. 127—139. M. aegyptiaca; 127, frontal view of labrum (4 Sturova, CSR); 128, frontal view of left

antenna (same specimen); 129, external view of right hind tibio-basitarsus (same specimen); 130, obli-

que dorsal view of right hind tarsal claw (same specimen); 131, third submarginal cell of right fore wing

partial ventral and dorsal view of genital capsule (¢ Parlzán, CSR); 134, external view of right gonosty-

lus (same specimen); 135, the same of first described 3 (Egypt); 136, dorsal view of pygidial plate, part-

ly exposed (9 Hungary); 137, segments 3—4 of antenna (9 holotype M. lindbergi, Lanzarote, Canary

Is.); 138, lateral view of scutellar tubercle (same specimen); 139, dorsal view of pygidial plate (same

specimen)

246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

leg. M. Kocourek (CK); 1 & 2 9 (1 & diss., fig. 144), Slovak., Obelce, vi & 14.vii.1953, leg. M.

Kocourek (CK, ML). — Italy: 1 ¢ (diss., fig. 141, pl. 4 fig. 20), Lazio, Caffarella, 9.v.1937, no. 53

(INER); 1 9 (diss.), Lazio, Roma, vi.1947, Bisleti leg., M. luctuosa, det. Pittioni (NMW); 1 9, sine loc.,

1903, coll. A. Weis (SMF); 1 9, Carpentras (Vaucluse), 15—23.v.1953, P. M. F. Verhoeff (ML). —

Portugal: 1 ¢ (diss.), Sobreiral [Famalicäo, ca. 20 km NE of Oporto, M. Diniz in litt.] (MUC).

Male. — Labrum distinctly longer than its breadth at base (100: 80 approx.),

slightly narrowed toward apex; surface concave, coarsely rugosely punctate,

lacking a distinct median ridge but frequently with short, shiny impunctate

subapical median streak ending in a minute pointed tubercle; or else, with

somewhat broader, impunctate median area extending from near end of basal

tubercles as far as the end; anterior border otherwise broadly rounded, more

rarely almost straight. Clypeus less closely punctate, punctures at sides isolated on

a smooth surface. Head above dullish, all punctures, including those on para-

ocular area, contiguous and most crowded together upon summit behind ocelli;

frontal line feebly indicated and only slightly raised posteriorly; impunctate juxta-

ocellar area of irregular form, hardly exceeding size of one ocellus; interocellar

distance hardly less than diameter of median ocellus. Antenna (fig. 128), the

rhinaria distinct, well impressed but rather small, oval. Mesonotum and scutellum

closely, deeply punctate, punctures of different sizes, those surrounding median

area of mesonotum smallest, contiguous, the ridges dull; centrally, nearly all

punctures are isolated, deepest and larger than interspaces, the latter but slightly

shiny. Scutellum, propodeum and thoracic sides dull, very closely punctate, the

scutellum feebly biconvex, lacking median line; spines variable in length, usually

robust, acute and spike-like, directed almost straight back, punctate dorsally.

Integument of legs somewhat glossy, closely finely punctate; hind femur lacking

raised hairs of any length interspersed between the short and dense posterior

brush, the latter in profile just visible with a hand lens (x 10); suberect bristles and

thick setae at tibiae and tarsi of moderate length; mid tibial pad distinct, though

nowhere sharply outlined, hind border fringed with some long white bristles (fig.

11). Outer face of hind tibia reticulate-punctate, distal 3/4 of surface hidden under

white pubescence; hind basitarsus slender, more than three times longer than

broad (fig. 129). Inner rami of mid and hind tarsal claws long and slender, over 4/5

length of outer and of equal breadth (fig. 130). Wings relatively long, not much

obscured, bases subhyaline, only centres of submarginal and larger cells as well as

most of the broad papillate border, slightly and diffusely infuscated; third

submarginal cell a little longer than high, its distal side rather strongly elbowed

much more so than the convex proximal side (fig. 131). Abdomen elongate-oval,

broadest at end of segment 2, rather flattened, apex pointed; tergites even more

shiny than in baeri; punctures fine, not very close, most superficial and widely

spaced on tergite 1, closest (though smaller than interspaces) on apical segments.

In profile, the short black semierect setae covering disk of tergites, are all visible

with a hand lens. Raised hairs at base of tergite 1 very long, though sparsely

distributed in middle, becoming more numerous laterally to form long compact

tufts of white, the hindermost hairs decumbent; spots on 2—5 varying much in

shape and size, placed far laterad in a regular row, those on 2 and 3 less defined

LIEFTINCK: Palaearctic Melecta 247

Figs. 140—145. M. aegyptiaca, sternites 7 and 8 of ¢; 140, from Cairo (first described example of that

sex); 141, from Caffarella (Italy); 142, from Sturova (CSR); 143, from Morocco; 144, from Obelce

(CSR); 145, from Parlzan (Slovakia)

anteriorly, the hairs being more raised and dispersed. Sternites progressively more

closely punctate from base to apex of abdomen, punctures on I—2 much smaller

than interspaces, the postgradular areas of all superficially and more sparsely

distributed, hind border of 2—4 often more brownish than black. Sternal plate 6

distinctly narrowed and tapering toward a bluntly pointed apex; lateral ridges

distinct though low, little oblique and hairy, barely visible in profile. Exposed

portion of tergite 7 (fig. 132) with straight converging sides, characteristically

brown-haired. Genital capsule rather large, 1.6 mm approx., gonocoxal enclosure

shaped much as in baeri, mesial borders of coxites at first diverging and concave,

but soon almost parallel as far as the bend, the angle hardly noticeably incurved

(fig. 133). Gonostylus about twice as long as its breadth at base, subtriangular in

outline and tapering gradually to a blunt point; dorsobasal process broadly

attached to base of stylus, its free portion broad, though short and small, tip

248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

bluntly triangular, the marginal bristles long but nowhere curled inward (figs.

134—135).

The most helpful and easily observable recognition marks of aegyptiaca are

found in the shape and hairiness of the 7th tergite of the male and pygidial plate of

the female. Other features are summarized in the descriptive keys, which together

with the illustrations and photographs (pl. 4 figs. 19—21) may suffice to distinguish

the species fairly easily from its congeners. The elongate body form, pale grey-

brown tinge of the fore wing membrane, dense comb-like brush of short hair at the

posterior face of the hind femur, as well as the widely separated white tergal spots,

are distinctive features of the male. Females are less easily determined, but after

uncovering the tongue-shaped brown pygidial plate, the characteristic shape of the

latter will be evident.

Central European aegyptiaca are absolutely indistinguishable from those

collected in the Mediterranean. I am unable to explain the discontinuity of its

range, unless the scarcity of this bee, and unconcerned collecting, are responsible

for the wide gaps existing between the northern accumulation of localities in the

Danube basin area, and the scattered southern habitations: a marked contrast

possibly due to the efforts of several well known entomologists in Czechoslovakia

and Hungary, who since many years paid special attention to the bee fauna of their

countries.

There is little variation in size among the various populations, the body

measuring from 14.5—16.5 mm in length, independent of locality. The noted

difference in stature between the photographed males on pl. 4 figs. 19—21, are

entirely due to the extension of the abdomina in the dissected specimens. The wing

span of these mounted examples measure 29.4 mm (fig. 19), 24.8 mm (fig. 20), 28

mm (fig. 21), and of the holotype lindbergi (not shown), 27 mm.

Distribution. — The scattered distribution is shown on map I, the arrow

indicating its further southward range.

Melecta fumipennis spec. nov.

(figs. 146— 149, pl. 3 fig. 13)

Type material. — Egypt: 1 & (diss., figs. 146—148, pl. 3 fig. 13), labelled

“Egypt, Wadi Gerani, 3.iii.1935, on Stachys, Dr. H. Priesner’’. Holotype (ex coll.

Priesner, ML).

Further material. — Egypt: I 9, labelled “Dep. Agr. Egypt, Giza, 23.iii.1914, coll. Naguib”,

“Melecta fumipennis Alfken 1933” and “Typus” (print on red), det. J. D. Alfken, in Alfken’s

on red) same identification label (MNB); 1 9, Um Assad Egypt, 9.iv.1934, Dr. H. Priesner, Paratypus

Um Assad, Egypt, 9.iv.1934, on Stachys, Dr. H. Priesner, one with Paratypus (print on red) and Melecta

fumipennis m., in Alfken’s writing, det. J. D. Alfken 1936 (ex. coll. H. Priesner, ML). All females are

paratypes.

A large, totally black, short-haired species lacking white spots, with very dark

fore wings.

LIEFTINCK: Palaearctic Melecta 249

Male. — Labrum rather long (length/greatest breadth ratio 100 : 70), broadest at

level of low , somewhat flattened and shining basal tubercles bearing one or two

large punctures; surface otherwise almost flat, coarsely densely striato-punctate,

distal portion with incomplete shiny median line or smooth ridge, anterior border

straight or slightly emarginate (some females!), median line ending in a minute

upturned subapical tubercle; whole surface clothed with longish raised bristly

hairs. Antenna long and strong, exceeding tegula, last segment obliquely

truncated; scape normally curved, longer than 3, the latter a little longer than its

breadth at apex (10: 8) and also somewhat longer than 4 and next segments, which

are subequal, all slightly longer than broad; rhinaria on 3—13 distinct, deeply

impressed, the one on 3 elongate-oval, 4—11 horseshoe-shaped, 12—13 almost

circular. Clypeus closely punctate, punctures circular, most crowded together and

I |

ty]

) M | AIN 1

WHY PAD IU

Figs. 146—149. M. fumipennis; 146, ventral view of tergite 7 (4 holotype, Egypt); 147, sternites 7 and 8

(same specimen); 148, partial ventral and dorsal views of genital capsule (same specimen); 149, dorsal

view of pygidial plate (9 paratype, Um Assad, Egypt)

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

smallest upon basal part of disk, growing larger and deeper distally, surface at

sides shiny with punctures more elongate and much smaller than interspaces;

whole anterior border broadly impunctate and glossy; upper surface concealed

from view by a patch of long appressed pubescence, but sides with equally long

raised hairs. Head above and at temples otherwise coarsely punctate on dullish

ground, punctures almost contiguous; small impunctate (though not polished)

areas just in front of median and on either side of lateral ocelli. All thoracic

sclerites dull, closely coarsely punctate, except a little impunctate area mid-

posteriorly on mesonotum; punctures otherwise deep and contiguous, those on

scutellar areas coalescent. Scutellar spines strong, smooth and shining, slightly

curved with bluntly pointed tips, directed obliquely upward and backward and

projecting well beyond level of almost vertical posterior sclerites. Legs rather long

and slender, without peculiarities; hairless areas dull, but posterior faces of fore

and mid femora slightly more shiny, very closely and rather deeply punctate, those

of hinder pair more coarsely so. Outer faces of tibiae and tarsi rugosely punctate

lacking conspicuously shiny areas, mid and hind tibiae and tarsi, in addition to

short appressed pubescence, with slightly stronger spine-like setae and semi-erect

bristles somewhat sparsely intermixed, especially at margins; setiferous punctures

on mid and hind basitarsi closely set and evenly distributed. Hind basitarsus about

three times longer than broad, slightly but distinctly outcurved, subparallel-sided

in side view with lower border only very little convex. Inner rami of mid and hind

tarsal claws about % lenght of outer and not or scarcely broader than these. Hair

mostly short and dense, but fore and mid femora with much longer and denser

posterior fringes, which are replaced by a very short comb at the blunt posterior

median ridge of hinder pair. Felty pad covering outer face of mid tibia distinctly

flattened and rather obscured, only an isolated and ill-defined elongate-oval

central area remaining dirty brownish white, the surrounding hairs consisting of

much stronger, suberect crowded bristles with shining black denticles more

sparsely intermixed. Wing membrane very dark (see specific key), remaining pale

brownish or subhyaline areas in fore wing are: a broad anterior streak in radial

cell, a well defined narrow stripe in the cells bordering anal vein, and most of the

anal (vannal) lobe; base of hind wing subhyaline except most of the papillate distal

one-third becoming light brown; third submarginal cell slightly less than twice the

size of second, not longer than high, its outer side rather strongly angled.

Abdomen oval, strongly convex, sides of intermediate segments subparallel, last

segments rather abruptly tapered; integument rather dull, but tergite 1 more

shining, very finely, superficially and more sparsely punctate than next ones, on

which punctures, though very small, are more crowded especially at sides, where

all are more widely spaced; hind margins narrowly impunctate; sternites similar,

but punctures larger and more deeply impressed, except those on broad hind

margin of sternite | sparsely and finely distributed. No trace of compact lateral

spots on any of the tergites, the first with thin raised fringe upon middle of basal

portion, the lateral tufts being, however, conspicuous, more compact and longer.

Succeeding segments clothed evenly and not very densely with very short suberect

hairs, longer and bristly ones being present only at sides of tergites and fringing

gradular lines of sternal plates 1—3, the posterior parts of remaining sternites

LIEFTINCK: Palaearctic Melecta 251

clothed more densely with short hairs. Tergal plate 7 rapidly tapered with narrow

tip, the apex itself parallel-sided for a long distance, becoming gradually more

deeply and broadly hollowed out above, its surface shining, carrying large elongate

punctures; hind margin, on either side of an almost semicircular emargination,

with upturned rounded side lobes; no submarginal ventral ridges (fig. 146). Hind

margin of sternite 5 shallowly emarginate, apex of 6 a little projecting, surface

scarcely elevated laterally, the ridges being very long, though more densely hairy

than on median area. Shape of sternites 7 and 8 as in fig. 147. Genital capsule of

normal size, 2.0 mm; gonocoxal enclosure U-shaped in dorsal view, the angles

obtuse, broadly rounded (fig. 148). Gonostylus short, thumb-shaped; dorsobasal

process more or less axe-shaped, its oblique distal border fringed with strong

bristles of great length.

Female. — See descriptive key.

This extraordinary species, both sexes of which are, I believe, adequately

characterized in the keys, has no near allies. Although immediately known by its

sombre outward appearance, male and female being very similar in this respect,

they do not differ appreciably in structure from other regional species (figs.

146—149). I have adopted Alfken’s specific name, which has remained a nomen

nudum in two collections.

A new species of Anthophora from Egypt, also of large size, measuring 19 mm in

length, was described and named A. fumipennis by Alfken (1926: 120), after a single

female. Like the present Melecta, this is an almost totally black species having

black-brown fore wings with bluish sheen. Priesner (1957, Bull. Soc. Ent. Egypte

41: 86—87), examined a second female labelled ““Dabaa (May)”, also in Egypt.

The male is still unknown. It would be of great interest to keep watch for this bee

at the above localities of M. fumipennis, with a view to find out whether the

Anthophora may be, perhaps, the host of its namesake Melecta, the two being so

much alike.

Melecta angustilabris spec. nov.

(figs. 150—154, pl. 3 fig. 14)

Paratypes (CMS, CDB, CBS, ML). 1 & 1 9, Aegypten, Heliopolis, A. Andres 4.ii.1914 (9) and

19.11.1915 (g), the latter with note ‘“Me/ecta sp.n. Labrum lang”, in J. D. Alfken’s handwriting, Sig.

Alfken (MNB); 1 9, Egypt, Min. Agric. Coll., Six Tower Suez Road, 26.iii.1926, R. C. Efflatoun,

Melecta nigra Spin. 9, det. J. D. Alfken 1933 (MNB); 1 9, Aegypten, W.-Riched, 1.iii.1914, A. Andres,

B.S.(BM).

252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

150

Figs. 150—154. M. angustilabris; 150, lateral and frontal view of labrum (4 Fayed, Egypt); 151, ventral

view of tergite 7, tubercles not shown (same specimen); 152, sternites 7 and 8 (same specimen); 153,

Egypt). Scale line 0.5 mm (fig. 152)

A small, strikingly sexual dimorphic species.

“Male (pl. 3 fig. 14). — Basal tubercles of labrum large, elongate, smooth and

shiny, bearing few rugae; whole surface striatopunctate bearing scattered

setiferous punctures, apex with short median crest, distal margin scarcely

upturned, shallowly emarginate with dense fringe of short stiff bristles. Antenna

reaching back to a little beyond tegula; scape of normal length, slightly curved,

with dense lateral fringes of long silvery white hairs, those in front and behind

being less abundant and shorter; 3 in frontal view subequal to 4, in posterior view

distinctly longer; 4 markedly longer than broad and subequal to next segments;

rhinaria on 3—12 (absent on 13), deeply impressed, placed in the long axis,

elongate-oval on 3, subcircular and diminishing gradually in size toward apex,

those on distal segments (11—12) very small. Disk of clypeus closely punctate but

anterior border narrowly bare; upper part at sides of disk with large, widely spaced

punctures, the parts below lateral bends almost polished, with much finer

punctures. The closely punctate vertex with three small polished areas, one in

front of median ocellus, the others beside lateral pair. Mesonotum, pleurae,

scutellum and postscutellar sclerites dull, punctures deep lacking interspaces,

except a fairly large smooth area on either side of mesonotal impression (about the

size of tegula) with few, widely spaced large punctures. Scutellar spines strong,

spike-like, though varying in length, directed obliquely upward and backward.

Legs of normal slender form, hairless areas rather shiny, sparsely superficially

punctate. Outer face of distal portion of tibia somewhat glossy, surface irregular

with large, deeply impressed punctures bearing thick, spine-like denticles. Hind

LIEFTINCK: Palaearctic Melecta 253

basitarsus distinctly curved in posterior view, a little less than four times as long as

broad, in side view slightly expanded about midway length, with (almost) straight

upper and slightly convex lower margin; inner rami of mid and hind tarsal claws

about three-fourth length of outer and not or scarcely broader than this. Vestiture

white on top of head, sides of mesonotum, scutellar areas, and middle of tergite 1;

some dark hairs only anteriorly at sides of clypeus and temples underneath, but

vertex across and beside ocelli with broad, almost hairless band; dark and partly

naked areas also posteriorly on mesonotum, between scutellar spines and medially

behind that level, on which parts hairs are black; conspicuous long white tufts

present wings and to the outside of scutellar spines. Legs mostly short-haired; all

tibiae and tarsi, in addition to short adpressed white and black tomentum, with

long dark bristles sparsely intermixed. External pad of mid tibia well defined

though not quite reaching base of segment; thick black setae strongest at apex of

tibia, the tips remaining partly visible amidst the white tomentum. Upper

(posterior) border of mid and hind basitarsi moreover fringed with long, erect

white and dark bristles. Second submarginal cell of fore wing much shorter than

high, its proximal side straight, the distal one convex; third cell twice the size of

second, a little longer than high, its outer side rather strongly angled. Abdomen

oval, but markedly pointed with tapering end segments; integument of tergites

shining, basal ones finely superficially punctate, on succeeding segments more

closely so, especially laterally on pregradular areas, but all punctures setiferous

and smaller than interspaces; hind margins of 1—2 narrowly impunctate. Sternites

dullish, punctation more even and also closer, especially on 1 and 2. Pubescence

covering sternites consisting of short and fine decumbent hairs, with longish raised

black bristles sparsely fringing gradular lines. Tergal plate 7 (fig. 151, ventral)

trapezoidal, basally almost flat, dull and coarsely punctate, then somewhat

hollowed out and impunctate with slightly upturned sides, apical emargination

obtuse-angulate with narrowly rounded angles; colour of apex rusty brown with

yellowish seam. Sternite 6 with pair of distinct low tubercles, which are just visible

in side view. Shape of hidden sternites 7 and 8 as in fig. 152. Genital capsule (fig.

153) relatively large, 1.6—1.7 mm; gonocoxal enclosure U-shaped, inner border of

gonocoxites at first diverging, then slightly incurved, but gonocoxal angle not

prominent, rounded; gonostylus shaped much as in luctuosa though more rapidly

tapered and bluntly pointed.

Female. — General appearance and body sculpture similar to male, as

described in the keys, the median mesonotal line equally somewhat sulcate.

Scutellar spines distinct, slightly raised, often somewhat downcurved, mostly

hidden under much longer pubescence. Legs slender, all parts not very densely

beset with long dark bristles posteriorly; outer faces of mid and hind tibiae clothed

with dense appressed black hairs and robust, spine-like setae; inner and outer rami

of mid and hind tarsal claws similar in shape, the inner branch about half as long as

outer. All white areas and tergal spots on the body of the male are typically

replaced by dark hairs, the vestiture thus being throughout obscurely brownish

black or black, with very rare exceptions.

Variation. — Only a single specimen in our series of topotypes (from Fayed,

ii.1943) represents a white extreme, approaching the male pattern, as follows.

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Small tufts or fringes of white present upon upper surface of head and antennal

scape, occiput, a narrow collar on mesonotum anteriorly (partly surrounding a

pair of small black dots), the mesepisterna and sclerites behind wings, and laterally

just beside scutellar spines; also a tuft of long raised hairs on either side of tergite

1; transverse postgradular spots at sides of 2 and 3, those on 3 only little shorter

than the interspace. Two or three other females are somewhat intermediate in this

respect, showing few light hairs on mesonotum and tergite 2.

Size small, total length 9.0—11.4 mm; only a single 9 (from Wadi Riched) sallies

out remarkably in being of much larger size, measuring 14.5 mm.

This is another easily recognizable species, apparently restricted to certain more

or less isolated areas in Egypt, and having no near allies. The very pronounced

sexual dichromatism is unique among Palaearctic Melecta.

Although Alfken had already noticed the unusual elongate form of the labrum

more than sixty-five years ago, and made a note of that on the labels (see above),

the species has so far remained undescribed. The photographed holotype male,

with somewhat retracted terminal abdominal segments (pl. 3 fig. 14), measures 9

mm in length and has a wing span of 20 mm, i.e. the average size of both sexes. It

must be emphasized that compact pubescent spots are present at the sides of

tergites 2—4 or 2—S in all females; these are shaped similarly to those of the male,

but all markings are brownish black or black instead of white and therefore not at

all clearly discernible.

Melecta guichardi spec. nov.

(figs. 77, 155—163)

Melecta luctuosa var. ebusana n.var., Friese, 1925, Konowia 4: 28 (pars: Ibiza only!).

Type material. — Turkey (Asia minor): 1 & (diss., figs. 155—158), labelled

“Turkey, Trabzon, Hamsikoy, 1245 m, 24.v.1962, Guichard & Harvey. B.M.

1962— 299”, Holotype (BM).

Further material. — Turkey (Asia minor): | & (diss., genit. wanting), Kleinasien, Sewdikoib/Smyrna,

(MNB). —Israel: 2 4, Palestine, Smith coll., Mrs. Farren-White (BM); 1 & (diss., ant. and legs partly

missing), Israel, Jerusalem, 25.v.1937, at Satureia, leg. Kugler (CBS); 1 9, Israel, Ein el Qilt (Kelt),

Lindos, 2.iv.1970, A. C. & W. N. Ellis (MA). — Romania: | & (diss., figs. 159—162), Dobrutscha 1882

(written), 13 (written on red), M. luctuosa Scop., det. Friese 1893 (MBUD). —Spain (Balearic Is.): 1 9

(diss.), Ibiza: 20.iv.1883/11, Friese, M. luctuosa v. ebusana Fr., det. Friese (MNB).

Male. — Labrum squarish, almost flat, anterior border straight, very slightly

upturned; surface shining, coarsely rugosely punctate, with raised black hairs only

scatteredly distributed, not very long. Mandible-bases finely closely punctate,

except upper one-third impunctate and glossy. Maxillary palpi normal, 5-

segmented, | very short, 2—4 slender, subequal in length to one another, 5 shorter

LIEFTINCK: Palaearctic Melecta 255

than preceding. Clypeus slightly convex in profile, throughout closely deeply

punctate lacking interspaces, except more widely spaced ones on anterior half of

exposed dorsal surface. Whole upper surface of head more deeply punctate on

dullish ground; narrow, impunctate, minutely striated areas in front of median and

beside lateral ocelli. Antenna rather long and strong, exceeding hind margin of

tegula, black; scape curved, subequal in length to 3—4 combined, clothed on

either side with fringe of very long finely branched white hairs; 3 short, little longer

than 4, about 1% as long as its breadth at apex, 4—13 shorter, intermediate

segments squarish, 9—12 even a trifle broader than long; rhinaria deeply

impressed. Thorax segments coarsely, closely and deeply punctate, all punctures

contiguous save for small, irregular, finely chagreened, impunctate median area

just behind mesoscutal line, the latter indistinct, obliterated, not impressed;

esi

Figs. 155—163. M. guichardi; 155, dorsal view of tergite 7 (4 holotype, Trabzon, Turkey); 156, sternites

7 and 8 (same specimen); 157, dorsal view of left half genital capsule (same specimen); 158, laterodorsal

view of left gonostylus (same specimen); 159, external view of right hind leg (4 Dobrutscha, Romania);

160, dorsal view of tergite 7 (same specimen); 161, sternites 7 and 8 (same specimen); 162, laterodorsal

view of left gonostylus (same specimen); 163, dorsal view of pygidial plate (first described 9, Rodos)

256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

transverse sulcus separating mesonotum and scutellum deep; scutellum only little

convex, median concavity shallow, tubercles widely separated, robust and spike-

like, directed almost straight back, punctate; all posterior and lateral parts of

thorax coarsely, contiguously punctate. Anterior half of tegulae strongly punctate

on shiny ground, surface suddenly becoming dull, very finely reticulated and

interspersed with scattered shallow punctures posteriorly. Legs rather strong,

throughout finely closely punctate on rather shiny ground, black. Mid tibia

somewhat swollen, broadest beyond middle, outer face convex; inner hind tibial

spur slightly nodded at about halfway length; hind basitarsus distinctly outcurved

and concave externally, broadest slightly beyond middle, upper border straight,

lower border convex, length-breadth ratio 100 : 33— 33.7. Wing membrane, except

fore wing basally, distinctly brownish, ends of marginal cells and centre of

submarginals more definitely infuscated; third submarginal cell equal in length to

its height, distal side markedly angled, more abruptly so than proximal side.

Abdomen short, oval, the apex rather abruptly pointed, integument black.

Tergites smooth, rather shiny, moderately strongly and deeply punctate, setiferous

punctures largest on basal part of tergite 1, though much smaller than interspaces

on all segments, becoming increasingly more widely spaced, smaller and more

superficial distally, the hind border of all tergites impunctate. Punctation of

sternites much more even and closer, except hind margins narrowly impunctate.

Pygidial plate of tergite 7 well marked off, brown, sharply black-ridged laterally,

sides below ridge vertical, extending from near apex anterad onto the disk, the

latter narrow, straight in side view, but surface at first somewhat convex, then

concave with feeble median impressed line as far as the apex, the latter narrow,

slightly upturned and emarginate; surface rather shiny, proximal part transversely

tessellate basally, the distal portion smooth, strewn with few large deep punctures;

oblique ventral keels subacute (figs. 155 & 160). Sternite 6 projecting medially, the

tubercles distinct though low and hairy, the thin shallow area separating them

almost three times as long as broad. Sternites 7—8, figs. 156 & 161. Genital capsule

relatively long and narrow, gonocoxites hardly expanded distally (fig. 157).

Vestiture (see key) long, predominantly white, but on head and thorax nowhere

quite concealing the surface. Sparsely distributed and black on disk of labrum,

along lower margin of mandibles, lateral faces of clypeus, the ocellar area, and low

down on temples behind eyes. No defined fan-like patch of white on disk of

clypeus: instead of this all hairs slanting upwards, those on paraclypeal, frontal and

occipital areas directed straight up and also white. Long pile covering pronotum,

basal half of mesonotum and upper portion of mesepisternum likewise white, but

lower part of sides and metepisternum black, as also the scutellum and vertical

sclerites of thorax; white spots are present on each side on dorsum just beyond

tegulae, behind wings at sides of propodeum, and long tufts immediately below

scutellar spines. Pile of legs not very long, except the usual white tufts fringing fore

and mid femora posteriorly, and scanty dark hairs at coxae and trochanters. Outer

faces of tibiae and tarsi all white, all hairs comparatively short and fine. Silky white

external pad of mid tibia nowhere sharply outlined, the hairs gradually becoming

thinner toward base: proximal one-fourth of the dark integument often exposed

between the short and scanty white pile, the passage oblique and ill-limited; pad

LIEFTINCK: Palaearctic Melecta 257

itself interspersed with robust, acuminate, deep black denticles, its anterior and

apical borders also beset with strong suberect black setae, especially toward apex.

Hind femur clothed all around with short, raised dark hairs which become longer,

more closely set (rather brush-like) postero-basally toward the blunt median ridge;

white pubescence covering outer face of hind tibia and basitarsus short,

decumbent, not very dense on distinctly shiny ground, the tibial hairs intermixed

with strong, suberect, spine-like black denticles, which are often partly hidden by

soft white hairs. Only few longish bristles projecting beyond posterior margin of

mid and hind tibiae and basitarsi. Abdominal tergites with short, erect black setae

distinct, though much shorter than in species like italica and tuberculata. Band-like

tergal spots 2 and 3 broadest and slightly concave anteriorly; postgradular areas of

sternites 3—5 thinly clothed with longish, straight, and but slightly raised silvery

hairs, especially conspicuous upon middle of sternites.

Female. — Differs from the male in some respects, as described in the key,

though mainly in sexual distinctions.

A delicate, slender species richly adorned with white, the hairs covering head

and thorax rather long and silky. Antennae relatively of great length but not very

strong. Easily distinguished from other small-sized species by the almost entirely

white-haired outer faces of mid and hind tibiae and tarsi, and also by the much less

defined mid tibial pad of the male, the depressed silvery hairs covering most of the

abdominal sternites of both sexes being also quite characteristic. Mandible, fig. 77.

The conspicuously broadened arms of the 7th sternite of the male and narrowly

tongue-shaped female pygidial plate, are reminiscent of the albifrons alliance, for

which reason I believe that guichardi, together with the somewhat allied fulgida,

leads over structurally to group | of the albifrons assemblage.

Evidently a somewhat scarce species.

Melecta nivosa F. Morawitz

(figs. 164—167)

Melecta nivosa Morawitz, 1893, Horae Soc. Ent. Ross. 28: 56—57 (9 Dshisak). — Popov, 1955, Trudy

Zool. Inst. Akad. Nauk USSR 21: 325 (locality record). — Lieftinck, 1972, Tijdschr. v. Ent. 115: 284

(generic status corrected).

Type material. — USSR: 1 9 (holotype), labelled ‘‘Dschisack iv.5 Glasun. [ow]

Coll. Morawitz” and “Melecta nivosa F. Mor. 9” in F. Morawitz’ writing (ZIL).

Further material. — USSR: & (diss., figs. 164—166), TransCaspi G. Turcmenien E. König (print),

Kilti-Tschinan (written) Coll. F. Morawitz (print), nivosa & F. Mor. (Morawitz’ writing) (ZIL); 1 9

(diss., fig. 167), Station Kuniu-Mazar, W. Bukhara, 28.iv.1912, Golbek (print, transl. from Russian),

(ZIL); 2 9, Farab/Buchara 13.iv.1913, Coll. Wollmann, nivosa det. Wollmann, and Balgakum bei

Djulek, Turkest., S. Malischew, nivosa det. Wollmann (ZIL); 1 9, Kara-Boges, 40 km N (illegible)

Arwata, 8.v.1953, nivosa det. Popov (ZIL).

The original diagnosis (in Latin) and description (in German) are based on a

single female from Dzhizak in Uzbekistan, NE of Samarkand. I found several more

individuals, including one male, in the Zoological Institute at Leningrad, one of

258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

either sex being still before me. Alle are undoubtedly conspecific; the following

characters of male and female are best given separately in addition to those given

in the keys and Morawitz’ description of the female.

Male (unique). — Labrum squarish, hence shorter than in female, but the

slightly concave surface similar; punctation dense, median ridge replaced by a

short, irregular, impunctate and somewhat shiny streak near apex. Maxillary palpi

slender, 6-segmented (4—5 apparently fused on left palpus!), the ultimate joint

short. Silvery patch covering whole upper surface of clypeus composed of long

and straight, decumbent hairs slightly exceeding anterior border; bristly hairs at

sides all black. Ocelli separated by a distance equal to diameter of median (largest)

one. Antennal scape white-fringed at all sides, but hairs are longest laterally. Head

closely contiguously punctate leaving no interspaces, black and almost hairless

across vertex and postocellar area, for the rest clothed with long raised white hairs

interspersed with black only at temples. Median mesonotal line neither sulcate nor

visibly impressed (cf. female!). Thoracic sclerites throughout dull, closely deeply

punctate, all white-haired, only mesonotum posteriorly and middle of scutellum,

as well as narrowly between spines and propodeum, black. Spines long, robust and

blunt, at least three times longer than their width at base, somewhat raised though

slightly but distinctly downcurved and directed almost straight back when viewed

from above; white tufts below spines conspicuous, considerably longer than these.

Legs moderately strong; all femora closely, finely punctate on somewhat shiny

ground; fore and mid femora with the usual long white posterior fringes, but

hinder pair clothed evenly all over with short, raised black hairs, fringed

posteriorly with very few longish dark bristles. Outer faces of all tibiae and tarsi

clothed with short depressed white hairs; marginal bristles short, partly brownish.

Mid tibial pad occupying almost whole surface, few black bristles only along

anterior and posterior margins, and some suberect, dark external spicules

intermixed toward end of tibia. Hind basitarsus rather broad, widest about midway

Figs. 164—167. M. nivosa; 164, dorsal view of tergite 7 (4 Kilti-Tschinan); 165, sternites 7 and 8 (same

specimen); 166, external view of right gonostylus (same specimen); 167, dorsal view of pygidial plate (9

Kuniu-Mazar)

LIEFTINCK: Palaearctic Melecta 259

length, upper border almost straight, the lower convex in side view (length-breadth

ratio 100 : 33 approx.). Mid and hind tarsal claws reddish, inner rami about %

length of outer. Wing-bases subhyaline, veins light brown except costa of fore wing

almost black; fore wing membrane distinctly infuscated toward apex, light to

darker brown, especially evident at junction of main anal veins midway length of

wing, the brown colour also filling out whole marginal cell and beyond; third

submarginal cell a little longer than high, its distal side moderately angulated.

Abdomen strongly tapered; white marks fairly well defined, composed of long

decumbent hairs; white collar on tergite | complete, narrowest and partly invaded

by black from behind at middle, spots on 2—5 occupying whole lateral surface,

those on 2 broadly, on next tergites only narrowly, interrupted by black medially,

all of them broadly rounded off inward. Sternal faces as described in the key, the

conspicuous white fringes exceeding hind borders; sternite 6 black-haired, the

tubercles distinct, elongate, broad and convex but scarcely visible in profile.

Tergal plate 7 occupying most of the surface, dark reddish brown, with slightly

upturned tips (fig. 164). Terminalia as described in the key and as in figs. 165-166.

Female. — Head as well as thoracic sclerites closely contiguously punctate;

mesonotal and parapsidal lines both distinctly though shallowly sulcate, punctures

on mesonotum irregular middorsally, separated in places by shiny areas much

broader than their own diameter. Scutellum broadly sulcate; spines robust, as in

male, shorter than spinal tufts, flattened, curved a little inward, raised obliquely

upward and backward, punctate. Thoracic pubescence shorter than in male,

especially anteriorly not longer than tegulae, black (and thin) on middle of

scutellum and (very dense) on parascutella. Legs slender; femora moderately

closely punctate and shiny, clothed sparsely with very long, partly dark, hairs;

tibiae and basitarsi (except black knees) white externally, fringed thinly with very

long white hairs posteriorly, outer surface of middle and hinder pair strewn with

exceptionally strong, black spine-like setae; inner rami of all tarsal claws half as

long as outer, strongly curved, acuminate. Pygidial plate very little downcurved,

surface flat, dullish and finely tessellate, apex with blunt, distinctly raised median

ridge, as in fig. 167. Body length 13— 14 mm.

This is an easily recognized species which differs from all its congeners by the

abundance of white pilosity covering most parts of the body. The male is highly

remarkable by the reduction (abbreviation) of the mid and outer hind tibial spurs,

a unique feature within the genus. On comparing the females in the Leningrad

collection, I noticed that the type as well as two others of that sex, were more

extensively white-haired than the one described above. The latter is a somewhat

worn individual with frayed wings whose body hairs are partly rubbed off. To

emphasize the uniqueness of the pubescent pattern exhibited by a fresh individual

of nivosa, it may be found interesting to attend to Morawitz’ own account of the

type:

“Das Gesicht ist wie auch die übrigen Teile des Kopfes mit langen

schneeweissen Haaren dicht bekleidet. Auf der Oberlippe sind den weissen einige

schwarze Haare beigemischt. Der Fühlerschaft ist weiss behaart, die Geissel

abgeflacht, deren mittlere Glieder nur wenig länger als breit erscheinen. Der

260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Brustkorb zeigt überall sehr dichte und lange schneeweisse Haare; nur auf dem

Schildchen stehen schwarze. Das Mittelsegment ist gleichfalls weiss behaart. Die

glänzenden schwarzen Flügelschuppen sind sehr fein und spärlich punktirt. Die

glashellen Flügel sind dunkel pechbraun geadert, der Aussenrand derselben

schwach rauchig getrübt, in der Radial- und den Cubitalzellen sind Spuren

dunkler Flecken vorhanden. Das erste Abdominalsegment ist mit längeren, die

beiden folgenden sowohl an den Seiten, wie auch deren hintere Hälfte mit

kürzeren, überall sehr dichten schneeweissen Haaren bedeckt. Auf dem zweiten

Segmente ist die helle Binde mitten unterbrochen, auf dem dritten aber nur mitten

dreieckig ausgeschnitten. Die beiden letzten Segmente sind vollständig

schneeweiss behaart. Die Ventralplatten sind glänzend, 2—5 am Grunde sehr breit

weisslich dicht tomentirt und mit wenigen dunkeln Haaren besetzt. Die Beine sind

weiss behaart, alle Schienen und Tarsen aussen dicht schneeweiss befilzt; die

Schenkel tragen an der hinteren Fläche lange schneeweisse Haare.” (loc. cit.: 57).

The male of nivosa is here characterized for the first time. (The specimens

enumerated above are not strictly syntypes, but since all were obviously compared

with the holotype, may be considered homotypical (or plesiotypical), as formerly

understood).

Melecta corpulenta F. Morawitz

(figs. 168— 172, 189, pl. 3, fig. 15)

Melecta corpulenta Morawitz, 1875, in Fedsch. Reise Turkestan 5: 140 (2 Zarafschan Valley, 18 May,

Oalyk rift, 2550 ft.). [Kyzylkum, Uzbekistan]; Morawitz, 1880, Bull. Acad. Imp. Sci. St. Petersb. 26:

371—372 (9 only, not ¢ ! addit. descr., Tjan-Schan, Dr. Regel leg.); Morawitz, 1895, Hor. Soc. Ent.

Ross. 29: 39—40 (addit. descr. 9 Tschuli, A. von Semenow); Morawitz, ibid.: 43 (compar. notes with

M. glasunowi Mor. nov. spec.). — Popov, 1955, Trudy Zool. Inst. Akad. Nauk SSSR 21: 325 (not

seen; compar. notes with M. baerii (Rad.) and M. nivosa Mor., and distrib. in USSR). — Lieftinck,

1972, Tijdschr. Ent. 115: 284 (generic position corrected).

Melecta baeri: Kokuév, 1909, Trudi Yaroslavskogo estest-istor. Obshchestva [Proc. Nat. Hist. Soc. Ja-

roslav] 2: 105 (distrib. notes, part., see also M. baeri).

Type material. — USSR: 1 9, under drawer label “‘corpulenta Mor. n. sp.”, with

small pin-labels “18” (print), ‘‘Zarafschan Valley (transl. from Russian, print), red

square, with “M. corpulenta Mor.” (in Morawitz’ handwriting). Lectotype M.

corpulenta Morawitz, by present selection (ZMM). See M. transcaspica F.

Morawitz!

corpulenta Mor., det.? ex Coll. Wollmann (ZIL); 1 9, Karatangeb. bei Djulek Balamurun,

Koshantschikow, M. corpulenta Mor., det. Wollmann (ZIL), 2 & (1 diss., figs. 168—171 & pl. 3 fig. 15),

both with two written labels in F. Morawitz’ handwriting: Margilan [Margelan, between Kokand-

Ferganal, M. robusta F. Mor., and Margilan, Mauser, M. robusta F. Mor., coll. F. Morawitz (print in

Morawitz (print in Russian), corpulenta Mor. 9 (in Morawitz’ handwriting) (ZIL, 1 9 ML); 1 9,

Krasnowodsk [E-coast Caspian Sea], Coll. Morawitz, M. corpulenta Q F. Mor. (in Morawitz’

handwriting (ZIL); 1 9, Samarkand [Uzbekistan], gold disk, ‘“‘M. corpulenta Mor. Typ. 9” (in

Morawitz’ writing) (ZIL); 1 9, Serbent Glasunow, M. corpulenta Mor., det. Morawitz (ZIL); I &,

“Masgiku (?) Menzer (?), coll. F. Morawitz”, M. robusta & F. Mor., det. F. Morawitz (ZIL); 1 9,

LIEFTINCK: Palaearctic Melecta 261

189), Baluchistan, Quetta, 5,500 ft., iv.1931, Capt. D. Harrison, B.M. 1932—353 (BM).

Doubtful specimens. — USSR: 1 9 (diss.), Uzbekistan, Kara-Tepe (Samarkand), 1.vi.1959, J.

Kohousek Igt., Melecta baeri Rad., det. Dr. Z. Padr (KU). — Turkey: 1 9 (diss.), with written label

“Mesopotamia Malatia 1886” [error for Malatya, in E Central Turkey?], ‘‘74b/100” (black-rimmed

square), “11““ (on red) (MBUD). — Israel: 1 9 (diss.), Jordan Valley, Jericho, 24.iii.1923, P. A. Buxton,

leg. Bytinski-Salz, one with M. aegyptiaca Rad? det. Mavromoustakis (CBS, ML).

This is the first described species of a remarkable little group of short-haired

Melecta with partly exposed thoracic dorsum and a relatively long first abdominal

segment.

As follows from the above, the type of M. corpulenta is a female in the Moscow

museum. Through the kindness of Prof. A. N. Zhelokhovtsev, I was enabled to

examine and redescribe this unique specimen during my visiting there in August,

1968. Shortly afterwards, I had occasion to consult the Melecta collection in the

Zoological Institute of the Academy, at Leningrad. Here an assortment of about

twenty bees, dispersed over three boxes, was found under the drawer label

“corpulenta”. These had been assembled and provisionally arranged by Morawitz

who, realizing their heterogeneity, identified and labelled only few of them. Many

years later, the same bees were examined also by the late V. B. Popov (1955), who

assigned a number of them to their proper genera and expressed his views on the

identity of some others. Unfortunately, the specific status of most specimens as

contained in the Leningrad collection was left undecided as not one of them was

definitely identified or labelled. It soon became evident that Morawitz had

confounded part of the available specimens of M. corpulenta with M. baeri

(Radoszk.), M. transcaspica F. Mor., 1895, and even with M. glasunowi F. Mor.,

1895 (now synonymized with Paracrocisa kuschakewiczi Radoszk., 1890, see

Lieftinck, 1972), a species described by him subsequently. The confusion was

initiated by Morawitz himself who, in 1880, mistook a male of a different species

— and probably of M. transcaspica as well — for corpulenta (see below). He aptly

remarked on the female of the latter (loc. cit., 1875, in Russian) “abdomen

covered with rather dense black down and accordingly wholly mat, almost as far as

tergite 5...”. And again in 1880 (in German): “Der Hinterleib ist bei frischen

Exemplaren mit einem sehr dichten schwarzen Tomente überzogen und erscheint

dann vollkommen mat ...”, etc., a definition applying perfectly to corpulenta.

Morawitz’ characterization of the supposed male, however, does not fit, as he

wrote: “Gesicht mit Einschluss des Clypeus, der vordere Rand des Mesonotum

sehr breit, die Meso- und Metapleuren, so wie auch das erste Abdominalsegment

lang und dicht weiss behaart, das fünfte jederseits weiss befiltzt, das letzte sehr

schwach ausgerandet. Die Beine wie beim Weibchen, die Mittelschienen und die

Metatarsen des dritten Beinpaares aber aussen vollständig weiss befiltzt. Im Tjan-

Schan von Dr. Regel gesammelt.” (loc. cit., 1880: 493—494). The last mentioned

male is possibly a mis-identified specimen of M. baeri.

The reasons for the existing confusion are manifold and can be easily explained.

As stated before (see p. 140), sexual dimorphism greatly hampers a correct

association of the sexes. Apart from some males, all doubtful specimens are

solitary females caught at random in places scattered all over West and Central

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Asia. Besides exhibiting an almost identical pubescent pattern, they hardly differ

from each other structurally and also seem to vary individually. This is exactly

what has happened with the museum specimens of the present taxonomic group

and annectent forms resembling them. Moreover, in the past, individuals of either

sex belonging to four or more species were obviously collected indiscriminately,

but may have occurred together in a given locality. It must be admitted that even

at present the identity of several specimens has to remain uncertain.

The following descriptive notes are based on Morawitz’ material of both sexes

enumerated above, comprising also 2 males from Margilan, labelled robusta (a

nomen nudum) by Morawitz, and | female from Quetta.

Male. — Labrum little longer than its greatest breadth at some distance from

base, narrowing gradually distad and lacking a median crest; length-breadth ratios

40 : 35, and 40 : 26 at apex; surface closely, coarsely punctate, punctures large,

mostly contiguous; border straight or shallowly emarginate, apex with indication

of a short upturned tubercle. Antenna moderately strong, reaching to a little

beyond tegula; segment 3 only about one and one-fifth as long as 4, which is a trifle

longer than broad and equals the succeeding segments. Summit of head dull;

vertex behind ocelli and postocular area very closely punctate, all punctures

contiguous; paraocular area just beside lateral ocelli impunctate, smooth and

rather shining, the outer half of the distance toward eye-margin with large,

circular, well-spaced punctures on equally shiny ground; a crescentic area just in

front of median ocellus also with few punctures on dullish surface. Punctures on

dorsum of thorax of different sizes. White hair tufts below scutellar spines fully

three times longer than these. Wing venation as in pl. 3 fig. 15; veins brown,

Figs. 168—172. M. corpulenta; 168, dorsal view of tergite 7 (4 Margilan); 169, sternite 7 (same speci-

men); 170, sternite 8 (same specimen); 171, partial ventral and dorsal view of genital capsule (same spe-

cimen); 172, dorsal and right lateral view of pygidial plate (9 Pendshikent)

LIEFTINCK: Palaearctic Melecta 263

membrane of fore wing lighter than in both honesta and amanda, subhyaline,

becoming gradually light smoky brown beyond cells, darkest in marginal and at

apex of radial cells. All sternites of abdomen dull, closely punctate and black-

haired, except 1—4 with crescent-shaped postgradular areas somewhat more

shiny, finely tessellate, with few superficial punctures. Apex of tergite 7 broadly

trapezoidal, disk almost flat, impunctate, either hairless and rather shining, or

naked only on both sides of a pubescent median streak (2 males from Margilan);

lateral ridges swollen and hairy, posterior border brown, bare and somewhat

upturned, with crescentic emargination and rounded angles (fig. 168). Raised

ventral tubercles at apex of sternite 6 elongate, distinct though low, clothed

densely with short decumbent hairs. Sternites 7 and 8 as in figs. 169—170. Genital

capsule normal, about 1.5 mm long, inner border of distal portion of gonocoxites

almost straight, enclosing a narrowly U-shaped space, the gonocoxal angle not

incurved, more evenly rounded than in honesta; gonostylus narrower and a little

longer than in that species, but dorsobasal process similar.

Female. — The type, with most others, shares the group characters summarized

in couplet 15 of the key. The most striking features are the dullish, closely

punctate abdomen with its fine, appressed, sharply outlined white tergal spots.

Even on tergite 1, the latter are completely isolated, composed of very short

matted hairs. Nevertheless even corpulenta exhibits great variation in the shape and

size of these markings. On tergite | of the type they are almost triangular, pointing

toward each other along base; those on 2 are large, more widely separated,

subquadrangular with broadly convex inner border, on 3 more transverse, less

deep and more approximated, while on 4 they are small and subcircular. Other

females have spots shaped much as in candida (pl. 3 fig. 18), though made up of

much shorter hair. The pygidial plate in our small series also varies somewhat in

shape and texture, the deflecting angle of the nodded apex being not exactly

identical in all (figs. 172 & 189). A number of specimens could not be determined

with absolute certainty, for which reason they are listed above under “Doubtful

specimens”. The variation is best demonstrated by three undoubtedly conspecific

females from Israel, all collected — at diffent times — near Jericho, yet differing

between themselves, not only in the presence or (near) absence of lengthwise

rugae on the sides of the propodeal triangle, but also in the varying distinctness of

juxta-ocellar pits. Lastly, one perfectly fresh example from Kara-Tepe

(Uzbekistan), has tergal spots almost of equal size and of exceptionally small size,

being almost circular on 1-2, while those on 3 are transverse though only little

prolonged inward. Otherwise this individual conforms to corpulenta from Israel

and Turkey. All of them can be distinguished from similar-looking bees by the

markedly sulcate median mesonotal line. Unfortunately no males are available of

any of these aberrant specimens.

In this connection it is of interest to observe that the solitary female from Quetta

corresponds very closely with typical corpulenta and consequently can not possibly

be associated with the unique male of our new species candida (pl. 3 fig. 18) which,

though bearing exactly the same printed locality label, was collected one year

earlier in a different month. It is undoubtedly specifically distinct! (See under that

species, the female of which is still unknown).

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

The measurements of the lectotype are: length 1.45 mm, fore wing 12 mm. It

agrees very closely with the female from Quetta, still before me.

Melecta honesta spec. nov.

(figs. 173—178, pl. 3 fig. 16)

Type material. — Turkey (Asia minor): 1 & (diss., figs. 173—178 & pl. 3 fig. 16),

“Türkei, Mut, 28.v.1967, J. Gusenleitner” (print). Holotype (CG).

Further material. — Turkey (Asia minor): 5 9 (diss.), same labels as holotype, Mut, 9—13.vi.1965 (1

9, CG), 10.vi.1968 (1 9, fig. 178, CG), 19.v.1970 (1 9, ML); 1 Q (diss.), Asia min., Mut, 275 m,

Turcia, Karapinar, 995 m, 15—17.vi.1969, G. Friedel (CG). — Egypt: 1 9 (diss.), Amrieli (?, written),

17.iv.1914, coll. Alfieri/Egypt, Melecta spec., Alfken det. 1928, Crocisa dimidiatipunct Spin., det. J. D.

Alfken 1933, Crocisa nubica Lep., det. J. D. Alfken 1939, all identification labels in Alfken’s

handwriting (MNB). — Israel: 1 & (diss., figs. 173—174, Palestine, Jerusalem, 25.v.—10.vi.1941,

Melecta caesarae Friese? (both in H. Bytinski-Salz’ writing), and Paracrocisa sinaitica (pencil writing,

det.?) (CBS). — Libya: | ¢ (diss., fig. 175), Cyrenaica, R. U. Agrario, CIRENP (?) V, Geo C. Kruger

(print), ““Pseudomelecta spec. nova”, and “aff. Ps. (Melecta) aff. glasunowi J. Mor.” (both labels in J. D.

Alfken’s writing), det. J. D. Alfken 1933 (BM).

Male. — Labrum short and square, but broadest at middle (ratio 50 : 50), distal

border nearly straight; surface but little concave, coarsely contiguously punctate

lacking a median ridge, but apex with minute upturned median tubercle; bristles

short, black. Silvery white hairs covering disk of clypeus shorter and less compact

than in amanda; anterior border and sides bare, the former punctate, the sides

striated. Antenna moderate, a little stronger than in amanda, reaching as far as end

of tegula; scape with lateral fringes of white also shorter than in that species;

segment 3 about one and one-fourth as long as 4, which is a trifle longer than

broad, equalling the succeeding segments in length. Vertex closely punctate, but

small, smooth shiny areas are present in front of median and on either side of

lateral ocelli. Mesonotum, scutellum and parascutella with large, closely set, deep

punctures leaving no interspaces, the ridges separating them dull; punctures

smallest and most crowded alongside slightly impressed parapsidal lines; median

mesonotal line indistinctly sulcate. No median scutellar line; spines short, similar

to amanda, but punctate all over. Long pubescence covering head, thoracic

sclerites and legs, comparatively short and thin, the integument on most parts

remaining visible; hairs at temples, thoracic sides, sternal faces, tufts behind wings,

and sides of propodeum, all white; hair upon scutellar lobes and parascutella deep

black, the white tufts surrounding apices of scutellum short and thin, not much

longer than spines. Legs slender; hind basitarsus subparallel-sided, length-breadth

ratio 100 : 36.3. Outer faces of tibiae predominantly white, of tarsi mixed black and

white; hairs short, decumbent, pile covering hind tibiae not very dense, leaving

black-tipped setae shining through; integument of distal one-fourth almost

hairless, the underlying surface rugosely punctate and somewhat glossy. Tarsi

black, claws dark brown, inner rami of middle pair about half as long as outer, on

the hind legs slightly shorter. Wing venation as in pl. 3 fig. 16; membrane of fore

wing much darker than in amanda, as described in the key. Sternites for the greater

LIEFTINCK: Palaearctic Melecta 265

Figs. 173—178. M. honesta; 173, ventral view of tergite 7 (3, Jerusalem); 174, sternites 7 and 8 (same

specimen); 175, sternites 7 and 8 (4 Cyrenaica); 176, sternites 7 and 8 (4 holotype, Mut, Turkey); 177,

partial ventral and dorsal view of genital capsule (same specimen); 178, dorsal and right lateral view of

pygidial plate 9 paratype (Mut, Turkey). Scale line 1 mm

part dull, closely punctate and clothed with short decumbent hairs, longer bristles

being sparsely distributed only at gradular lines; small white spots present on

either side at apex of sternites 2—5. Apex of tergite 7, fig. 173; upper surface,

proximal to somewhat upturned naked tubercles, with smooth, rather glossy,

finely transversely wrinkled median area, the sides clothed with longish, mostly

depressed, brown bristles. Raised ventral tubercles at apex of sternite 6 distinct,

though small and not very prominent, mostly covered with dark hairs. Sternites 7

and 8, figs. 174— 176; arms of 7 straight, relatively broad. Genital capsule normal,

1.5 mm long, inner border of distal portion of gonocoxites not strongly concave,

the space enclosed more broadly U-shaped than in amanda; gonocoxal angle,

though slightly prominent, rounded off (fig. 177); gonostylus broad, widest basally,

apex broadly rounded, dorsobasal process broad at base, apex short,

subtriangular, free margin beset with long bristles.

Length 14.0— 15.0 mm approx, fore wing 10.2—11.5 mm.

The two males from Israel, besides being somewhat larger in size, are a little

266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

different from the holotype by having less obscured fore wings, approaching

amanda in this respect, but the abdominal markings are more nearly quadrangular,

as they also are in the females that were collected simultaneously with the type. On

dissecting out the copulatory apparatus (with its hidden sternites), the latter

proved to be very similar in form to those of the Turkish specimen. Moreover, the

Israeli males could not possibly be associated with the questionable females from

the same country which I assigned to corpulenta (antea), because in them the

abdominal spots are placed farther apart and definitely more rounded in shape.

Female. — Differs from the male by having much darker wings, still more

conspicuous, subquadrangular silvery marks, which at the sides of tergites 1 and 2

are nearly of equal size, standing out brightly in contrast with the deep black

ground colour of the rest of the abdomen. It differs from our small series of

amanda females also by the more squarish form of the first two tergal spots, those

on segment | of amanda being distinctly more prolonged inward and oblique than

in honesta. Otherwise the two species are much alike and undoubtedly closely

related.

Without the possibility of consulting the photographs of male corpulenta, honesta

and amanda (pl. 3 figs. 15, 16 and 17, respectively), it would be difficult to obtain a

good impression of the peculiar “facies” of these remarkable and rather aberrant

bees. As mentioned in the key and shown in the pictures, the great length of the

first abdominal segment in proportion to the second, — already referred to in an

earlier paper (Lieftinck, 1972: 264—265) — is the most striking feature separating

them from all others; this is accentuated by the outstanding white lateral patches,

which do not consist of partly raised tufts, as is usual in regional Melecta, but are

made up of sharply delimited marks of much shorter, decumbent, silky hairs

shaped similarly to those adorning the next segments. More recent investigations

showed, however, that these characteristics are not tenable as subgeneric criteria,

since there are a number of species showing transitions to normal (e.g. alecto and

candida), the position of transcaspica being intermediate in that the tendency

towards equalization in length of the basal segments is hardly present and only

seen in the female.

M. honesta and amanda are e ‚ıdently very locally distributed and rare species as

only few specimens of each have become known.

Melecta amanda spec. nov.

(figs. 76, 179—184, pl. 3 fig. 17)

Type material. — USSR: 1 ¢ (mounted, diss., figs. 179—182 & pl. 3 fig. 17),

“Turkestan” (typewritten on blue-rimmed label “Coll. Gribodo”) (blue print).

Holotype (MCG).

(print), “Collection A. Weis” (print), ‘“Melecta luctuosa Scop. var. albovaria Er. Turkestan”

(Schmiedeknecht’s handwriting) (SMF); 1 9 (diss.), with small written label ‘““Sarepta’’ [S Russia] and

“ex coll. Radoszkowski Inst. Zool. P. A. N. Kraków 25/27” (small print), ‘‘Pseudomelecta baerii Rad.”

(written) and on same label “ex coll. Radoszkowski” (print) (IZK).

LIEFTINCK: Palaearctic Melecta 267

Figs. 179—184. M. amanda; 179, external view of right hind leg (¢ holotype, Turkestan); 180, ventral

view of tergite 7 (same specimen); 181, sternites 7 and 8 (same specimen); 182, partial ventral and dor-

sal view of genital capsule (same specimen); 183, dorsal view of pygidial plate (Q Djarkent, Turkestan);

184, dorsal and left dorsolateral view of same (9 Turkestan)

Male. — Labrum somewhat longer than broad at base, apex narrowed, length-

median- and subapical breadth ratios as 40 : 33 : 26; apical border straight, surface

coarsely punctate, punctures of different sizes, contiguous and partly coalescent,

distal % with feebly indicated median ridge; bristles short, black. Disk of clypeus

thickly clothed with long, decumbent silvery hairs completely concealing the

surface, sides striato-punctate. Scape of antenna with long and dense fringe of

white, these hairs longer than diameter of scape; segment 3 little longer than 4,

which itself is about one and three-fourth times longer than broad and equal in

length to the succeeding segments. Hairless areas on upper surface of head with

punctures of different sizes on rather shiny ground: area in front, behind and partly

also at sides of ocelli with larger, almost contiguous punctures, those on

paraocular area smaller anteriorly, more crowded than in honesta; a fairly large

area on either side of lateral ocelli and a smaller, crescentic space just in front of

median ocellus, smooth and shining. Punctation of mesonotum, scutellum and

parascutella much as in honesta, but all interspaces and ridges distinctly lustrous; a

pair of small, shiny impunctate spots also upon dorsum of mesonotum, just before

level of tegulae; median mesonotal and parapsidal lines distinctly sulcate.

Scutellar spines short and small, placed transversely, much broader than high,

268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

directed straight back and forming an isosceles triangle in dorsal view; surface

punctate, but tip smooth and shining. Long pubescence covering head, thoracic

sclerites and legs markedly longer and denser than in honesta, tufts at sides of

propodeum mixed white and black; hair upon parascutella, disk of scutellum and

spines, black and also rather longer, the white tufts underneath spines

conspicuous, greatly exceeding the latter in length. Legs stronger than in honesta,

hind basitarsus subparallel-sided, length and breadth ratio 100 : 47.3. Outer faces

of all tibiae and tarsi almost entirely white, the hairs short, dense and decumbent,

evenly distributed and silvery, especially on hind basitarsus, the latter nearly three

times as long as its greatest breadth (fig. 179). No black-tipped spicules visible

within the compact pad of mid tibia and few only on hind tibia, the former with an

isolated dark hair-line anteriorly. Claws reddish brown, inner rami of mid and

hinder pair normal, one-half to two-thirds length of outer. Wing venation as in pl. 3

fig. 17; veins light brown, membrane hyaline, on fore wing becoming gradually

light smoky brown beyond the cells, darkest in marginal and at apex of radial cell.

Measurements: length 13.8—15 mm, fore wing 11.7—13 mm.

This is the third in a small group of conspicuous Palaearctic species differing

from the rest by a combination of characters summarized in paragraphs 15—17

(3) and 8—10(Q) of the keys. The female is marked similarly to the male (pl. 3 fig.

17), although all tergal white patches are a little less broad. Our specimens differ

from honesta in the less obscured fore wing membrane and the presence of white

tufts under the scutellar tubercles. I have no doubt about the correct association of

the sexes.

Melecta candida spec. nov.

(figs. 17, 185—188 & pl. 3 fig. 18)

Type material. — Pakistan: | & (diss., figs. 17, 185—188 & pl. 3 fig. 18), with

printed label ‘Baluchistan, Quetta, 3500 ft., x.1930, Capt. D.Harrison, B. M.

1932—353”. Holotype (BM).

Further material. — None. For a 9 carrying an identical locality label, see M.

corpulenta Morawitz (fig. 189).

Male. — Similar in general appearance and clear-cut white abdominal spots to

members of the M. corpulenta group, and possibly allied to these. Distinguished

from all by the lustrous abdomen and somewhat longer pubescence.

Labrum short and square, broadest at middle, closely rugosely punctate lacking

interspaces; basal tubercles relatively small, dullish, microscopically tessellate;

anterior border straight, with minute raised median tubercle at apex. Antenna

moderately strong, reaching beyond tegula, 3 about 1% as long as its breadth at

apex (100 : 72) and somewhat longer than 4, which equals the succeeding segments

in length, all being distinctly longer than broad. Clypeus closely punctate, the

anterior border narrowly, very finely, transversely wrinkled, matted white hair

covering dorsal surface not quite reaching front margin. Head above closely,

deeply punctate on shining ground, punctures on summit behind ocelli lacking

interspaces, elsewhere mostly separated by one puncture width; a conspicuous,

LIEFTINCK: Palaearctic Melecta 269

Figs. 185—188. M. candida (Pakistan); 185, external view of right hind tibio-basitarsus (¢ holotype, Qu-

etta, Baluchistan); 186, dorsal view of tergite 7 (same specimen); 187, sternites 7 and 8 (same speci-

men). — Fig. 189. M. corpulenta, dorsal and right lateral view of pygidial plate (@ Quetta, Baluchistan)

polished and slightly concave, area on either side adjoining lateral ocelli and a

subtriangular, finely chagreened, impunctate area also in front of median ocellus.

Mesonotum closely deeply punctate, punctures contiguous leaving shiny ridges;

median line distinctly sulcate anteriorly. Scutellum feebly bituberculate lacking a

median line; spines strong, spike-like, directed straight back, but tips slightly

downcurved, punctate dorsally. Raised dorsal thoracic pubescence not very

dense, consisting of grey-white hairs nowhere entirely: concealing the surface, not

much longer than diameter of tegula, though longer than in members of the

corpulenta group; pubescence at sides of thorax and underneath much longer and

denser, all white. White tufts surrounding scutellar spines long though not

definitely outlined; scutellum and parascutella black, the closely punctate, rather

shiny scutellum remaining well visible under the raised hairs. Legs less slender

than in alecto, shaped much as in alcestis and honesta, the more robustly built

amanda having stronger legs than all of them. Punctation and nature of

pubescence similar to these species. Mid tibial pad, fig. 17. Femora more dullish

than in alecto and also less compressed laterally, the posterior ridge blunt and

incomplete, poorly developed only toward apex. Hind basitarsus (fig. 185)

distinctly more outbent and hollowed out externally than in alecto, and also

somewhat more expanded in side view. Inner rami of mid and hind tarsal claws

only half as long as outer, a little broader than in the allied species of the group.

270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Wings darker than in both alcestis and alecto, but much less obscured than in the

type of honesta: basal one-third of fore wing membrane, and that of the hind wing

entirely, hyaline, for the rest smoky light brown; third submarginal cell shaped

similarly to alecto, its distal side moderately strongly angulated, more convex than

proximal side. Abdomen rather shiny, as in alcestis, but integument of all tergites

decidedly more finely and closely punctate, though punctures are slightly less

crowded than in alecto. Pubescent pattern as in pl. 3 fig. 18; snow-white lateral

spots present on tergites 1—6, almost as compact and sharply delimited as in

alecto, in agreement with the slightly more widely spaced punctures; spots on

tergite | more oblique in dorsal view than in alecto, with numerous suberect

longish hairs all along base, which become decumbent and very dense laterally;

spots on 2 a little larger and more rounded inward than in alecto, those on 3—5

transverse, more regularly rectangular, but distance separating them as in that

species; spots on 6 very small. Raised black hairs on disk of tergites extremely

short, as in alecto, visible only under a strong lens. Sternal pilosity likewise very

short and depressed, with few longer suberect postgradular bristles, all black;

sternites moreover with a regular row of small, roundish, white hair-spots placed

on either side at some distance from the tergal margins. Sternite 6 dark brown,

apex broadly rounded, the apical tubercles distinct and hairy, but lower and more

apart than in alecto, the less sclerotised interspace being oval in outline. Pygidial

plate of tergite 7 brown, more rapidly tapered than in that species, disk almost flat,

rather shiny and hairless, the lateral rims hardly swollen, tip slightly hollowed out

with upturned, broadly emarginate hind margin; ventral ridges shaped much as in

alecto. Gonocoxal enclosure V-shaped, the inner borders of gonocoxites almost

straight, gonocoxal angle obtuse, broadly rounded off, not incurved; stylus with its

dorsobasal process shaped as described for alcestis.

Size moderate; length 14 approx., fore wing 10 mm.

Female. — Unknown.

This new species runs out in the key to near baeri, hence far removed from the

earlier defined taxa of the corpulenta group, whose members are, indeed, very

similar superficially. M. candida is, I suppose, more nearly related with alecto and

alcestis, all three being distinguished from the former, among other characters, by

the relatively shorter first abdominal segment, as mentioned in the key. For the

same reason it is impossible to regard the female from Quetta as the opposite sex

of the present species (see under M. corpulenta).

Melecta alcestis spec. nov.

(figs. 190— 194, pl. 2 fig. 11)

Type material. — USSR: 2 g (one diss., figs. 190—194), labelled “Jiz. Rusko

Toscoe” (i.e. south Russia), one with large label Melecta (print) “‘/uctuosa var.

eczmiadzini Rad.” (Friese’s writing), and both with ‘Melecta luctuosa v.

eczmiadzini’’ (unknown handwriting). Holotype alcestis (figs. 190—192, 194 & pl. 2

fig. 11), in NMP (Prague); topotypical paratype & (ML).

Doubtful material. — Israel: 1 ¢ (diss.), with written label “found in the car between Elat and

Jerusalem, 8.iii.1964'’, leg.? (CBS).

LIEFTINCK: Palaearctic Melecta 271

Male. — Labrum broadest about midway length, length-breadth ratio 100: 74;

anterior border convex, slightly produced, apex upturned with raised, shiny

marginal tubercle; coarsely deeply punctate, tubercles and disk beyond middle

with irregular, somewhat shiny, impunctate area. Antennal segment 3 short, only

little longer than broad at apex and only a trifle longer than 4, which is hardly

longer than broad, equal in length to succeeding segments. Clypeus closely

punctate, except anterior border narrowly smooth; tips of depressed silky white

hairs upon dorsal surface not or hardly exceeding anterior margin. Head above

shining, closely deeply punctate, punctures not contiguous, larger than

interspaces, except anteriorly behind antennae, upon summit behind ocelli, and on

paraocular area and temples, where punctures are partly coalescent. An irregular

shiny streak in front of median ocellus with few punctures. Mesonotum and

scutellum coarsely deeply punctate, all punctures confluent, leaving no

interspaces; median and parapsidal lines faintly indicated, not impressed; sulcus

between mesonotum-scutellum deep. Scutellum moderately biconvex, lacking a

median line; spines rather long and slender, directed straight back and hardly

downcurved. Propodeum dull, coarsely punctate, lacking impunctate areas.

Thoracic pubescence long, dense and silky, the sides and ventral sclerites all

white, but hairs nowhere entirely concealing surface; parascutella and middle of

scutellum black, the white tufts below spines much longer than these; lateral

propodeal tufts also long, white. Legs rather slender, all parts predominantly white

externally. Interpunctate areas of hind femur smooth and shiny; matted hair and

posterior fringe as described in the key, with very few longer bristles intermixed;

hind tibia and basitarsus reticulate-punctate; basitarsus subparallel-sided, about

À. >

GA LL IT

Figs. 190—194. M. alcestis (S Russia); 190, external view of left tibio-basitarsus (¢ holotype, Jiz Tos-

coe); 191, dorsal view of tergite 7 (same specimen); 192, sternites 7 and 8 (same specimen); 193, sterni-

tes 7 and 8 (second, topotypical specimen); 194, dorsal view of left half genital capsule (3 holotype)

DIR TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

three times longer than broad (fig. 190). Mid tibial pad distinct and compact

though not sharply defined, with some raised white hairs projecting posteriorly.

Inner rami of mid and hind tarsal claws about three-fourth length of outer. Fore

wing membrane lightly infuscated, smoky brown, especially toward apex; third

submarginal cell as long as high, distal side moderately angled, ar in alecto (pl. 2

fig. 11). Abdomen more expanded basally than in alecto, moderately lustrous,

finely, rather densely punctate, punctures much smaller than interspaces;

pubescence rather long and dense. Short black, suberect setae covering disk of

tergites well visible with a hand lens in profile. White tergal spots conspicuous and

broad, the raised hairs on | very long, condensed at sides but not sharply defined,

the depressed spots on 2—3 also ill-limited anteriorly, and with sparse admixture

of longish white bristles at gradular lines. Sternal vestiture consisting of almost

depressed, mainly black hairs intermixed with white: 2—5 with transverse gradular

bands of white, diverging posteriorly, one on each side of the median line. Sternite

6 black, apical ridges very low, hairy. Tergite 7 shaped much as in alecto but more

hairy and closely punctate; disk flat, margins broad and slightly swollen, apex a

little narrower and more hairy in paratype than in holotype (fig. 191); oblique

ventral ridges short, thick and densely hairy. Sternites 7—8, fig. 193. Genital

capsule, fig. 194. Gonostylus thumb-shaped in side view, but only twice as long as

broad at base, convex externally, apex evenly rounded; dorsobasal process broad

at base, curving suddenly upward to form a setiferous distal process which, though

much smaller, is similar in form to the stylus.

The sketches of the two — evidently topotypical — south Russian males clearly

demonstrate how different these sclerites can appear in simple glycerine

preparations mounted in exactly the same way. Also the apices of tergite 7 are not

quite alike in the two specimens, the whole plate in the paratype being a little

narrower and more densely hairy. Yet I am convinced that they are conspecific,

because in other respects the males are practically identical. The third male, from

Israel, may or may not belong here, the main incongruity consisting in the fact that

the gonocoxal angle is evenly and broadly rounded instead of a little incurved, as it

is in the type (fig. 194), a character otherwise of great constancy and importance.

More material is much needed to definitely establish its identity.

Female. — Unknown. I have not succeeded to discover a suitable partner for

this species among the many Melecta left unnamed.

A medium-sized species, superficially resembling festiva, but immediately

distinguished therefrom by the extremely short hairs covering the hind femur

posteriorly, and the less incurved gonocoxal angle. The conspicuous, oblique,

silvery hair bands fringing the graduli of sternites 2—4(5), one on each side of the

median line, are also characteristic, as is the relatively slender form of the legs.

Possibly allied to alecto spec. nov., which also occurs in Israel and is of the same

size. However, M. alcestis can be separated from that species by the presence of

longish black and white bristles at the tergal graduli (wholly absent in alecto), less

sharply defined abdominal markings, and also by the longer overall pubescence.

LIEFTINCK: Palaearctic Melecta 273

Melecta alecto spec. nov.

(figs. 195—201)

Type material. — Iraq: | & (diss., figs. 195—198), labelled “IRAQ: Kurdistan,

Shaglawa nr. Sala Huddin, N of Kirkuk, 19.v.1957, L. G. Higgins, B. M.

1957—391”. Holotype (BM).

Further material. — Israel: 1 g (diss.), Palestine, Mar Saba, 15 mi SE of Jerusalem, 15.iii.1922, P. A.

Buxton, Pres. by Imp. Bur. Ent. Brit. Mus. 1923—530, Melecta luctuosa, det.? (ex BM, ML); 1 & (diss.),

Palestine, Jerusalem-Jericho road, 29.iii.1943, H. Bytinski-Salz (CBS). — USSR: 1 & (diss., much

worn), 2 9 (diss., figs. 199—201), Turkestan: Ost-Buchara, Tschitschantan, coll. Hauser 1898 (print),

one pair labelled ‘‘Melecta sp.”, det. D. B. Baker (NMW); | g (soiled and worn), O. Turkestan, Narin,

E. A. Böttcher, M. ruthenica Rad., det.? (MNB). — Syria: 1 9 (diss.), Syria 1899, M. ashabadensis Rad.,

det. Friese 1900 (MNB). — Turkey (Asia minor): | 9 (diss.), 724/Brussa [Bursam, NW Turkey] (ML).

Male (holotype). — Labrum distinctly shorter than in alcestis: little longer than

its greatest breadth at middle, coarsely rugosely punctate lacking a median crest;

anterior border in frontal view almost straight, somewhat swollen and upturned,

with pair of minute, more or less confluent, marginal tubercles. Antenna of

moderate strength, segment 3 little less than one and one-fourth as long as its

breadth at apex (100: 70) and somewhat longer than 4, this and succeeding

segments almost square. Disk of clypeus nearly flat, surface shining, covered with

large, widely spaced punctures much as described for alcestis, only sides closely

punctate. Head above also closely deeply punctate, punctures greater than

interspaces; surface somewhat glossy; ocelli surrounded by a narrow, irregular,

dullish impunctate area prolonged forward to merge into the raised, smooth

frontal crest; some punctures in front of median ocellus more widely spaced than

the rest. Median mesonotal and parapsidal lines very fine, the former scarcely

impressed anteriorly, but more distinctly so posteriorly; mesonotum and scutellum

very closely, contiguously punctate, except distally upon disk of mesonotum,

where large punctures on either side of median line are somewhat more widely

spaced on shiny ground; sulcus between mesoscutum-scutellum as described for

alcestis. Scutellum normally biconvex, lacking a median line; spines triangular,

slightly divaricate, somewhat depressed, very little downbent, punctate. Thoracic

sclerites and propodeum otherwise as described for alcestis, the propodeum dull,

closely rugosely punctate lacking impunctate areas. Legs black, slightly more

slender and glossy than in alcestis, more throughout closely punctate and short-

haired; mid and hind femora more compressed laterally than in that species, all

with distinct though blunt median ridge, most marked at distal half of femur. Mid

tibial pad elongate-oval, well defined, its posterior fringe composed of few short,

widely spaced, black hairs. Hind basitarsus (fig. 195) very slightly outbent, as with

alcestis, but narrower basally and with both sides a little convex, about three times

longer than broad, length-breadth ratio 100: 34. Inner rami of mid and hind tarsal

claws relatively short, about two-thirds length of outer (ca. 67: 100) and a little

broader basally. Wing membrane as lightly infuscated as in alcestis; third

submarginal cell of fore wing moderately angled, length and height subequal, as in

that species, but second cell shorter, with distal side less outwardly convex. Tergal

274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Figs. 195—201. M. alecto (Iraq and Turkestan); 195, external view of right tibio-basitarsus (¢ holotype,

Kirkuk, Iraq); 196, dorsal view of tergite 7 (same specimen); 197, ventral view of sternite 6, showing tu-

bercles (same specimen); 198, sternites 7 and 8 (same specimen); 199, ventral view of tergite 7 (4 Ost

Buchara, Turkestan); 200, sternites 7 and 8 (same specimen); 201, dorsal and right lateral view of pygi-

dial plate (9, supposition, Tschitshantan, Buchara, Turkestan)

punctation of abdomen superficial, punctures fine and widely spaced on more

shiny ground, than in alcestis.

Thoracic pubescence much as in alcestis, but scutellum, between the white tufts

below spines, more broadly black; white hairs covering sides restricted to about

upper two-fifth of pleurae, the remaining parts black with scattered white tufts

sparsely intermixed (hence not at all white as in alcestis). Abdominal vestiture

much shorter than in alcestis, visible in profile only at high magnification; white

tergal spots conspicuous, transverse, placed in regular row, sharply defined even

anteriorly, all more condensed and longer (deeper) than in that species, with hairs

more finely branched; spots on | subtriangular, distance separating those on 1—2

about three times their own diameter, twice that distance on 3 and about equal on

4, spots on 5 smallest. Sternal pile also short, decumbent, with few suberect

postgradular hairs of somewhat greater length; no white spots. Sternite 6 black,

apex projecting somewhat between a pair of blunt hairy tubercles, one at each side

of an elongate, less sclerotised and much less hairy depression, these tubercles just

visible in profile (fig. 197). Tergite 7 (fig. 199) broadly trapezoidal, sides slightly

outwardly convex in dorsal view, the rims rather swollen and a little upturned, not

LIEFTINCK: Palaearctic Melecta 275

very hairy, apex shallowly emarginate with rounded angles; oblique ventral ridges

distinct, somewhat curved, the crests bare. Sternites 7—8, fig. 200. Gonocoxal

enclosure U-shaped, the inflexed margins (inclusive of gonocoxal angle) rather

strongly incurved (concave) mesially, but angulation well rounded; stylus

elongate-triangular in outline, twice as long as its breadth at base, apex bluntly

pointed; dorsobasal process broadest at base but shorter than in alcestis and

tapering gradually to a broadly rounded apex. Size moderate, total length 14—15

mm approx., fore wing 10 mm (holotype Iraq and male Turkestan).

The two males from Palestine (Mar Saba and Jerusalem-Jericho rd.), are

practically identical structurally and unquestionably not conspecific with the one

from the same country referred to alcestis with some misgivings (antea).

Female. — There is nothing of interest to complete the description in the key.

The two specimens labelled ‘““Tschitschantan” and Brussa, are in much better

condition than the male from the first-mentioned locality bearing an identical

printed label. I take it that all males and females are conspecific, but here also,

more material from all countries involved is greatly to be desired.

The females measure: length 15—15.5 mm, fore wing 11.5 mm.

Melecta transcaspica F. Morawitz

(figs. 10, 75, 202—214, pl. 4 figs. 22—24)

Melecta transcaspica Morawitz, 1895, Horae Soc. Ent. Ross. 29: 39 (¢ “Bei Hodscha-kala, von Pome-

ranzow gesammelt’’).

Melecta caesareae Friese, 1925, Konowia 4: 28—29 (9 “Caesarea; Palastina’’). Syn. nov.

? Melecta octomaculata Radoszkowski, 1876: 124(9 Egypte).

Melecta octomaculata Friese, 1925, Konowia 4: 29 (¢ “Kar-Boghaz, Taur. cil. & g Mersina, Kleina-

sien’’). Nom. preocc. Syn. nov.

Type material. — USSR, Turkmenia: | ¢ (diss., mounted, figs. 202—205, & pl.

4 fig. 22), labelled ‘“Chodshakala P.” (F. Morawitz’ handwriting), “Melecta

transcaspica F. Mor. 2” (id.), “ce. F. Morawitz” (print in Russian). Lectotype M.

transcaspica F. Morawitz, by present selection (ZIL). — Israel: 1 9 (pl. 4 fig. 24),

labelled ‘‘Caesarea, Palast.” (written), “Type” (print on red), Melecta caesareae Fr.

Kl. As.” [sic], in Friese’s handwriting. Holotype M. caesareae Friese (MNB). —

Turkey (Asia minor): 1 g, labelled “Kar-Boghaz, Taur. cilic.” [S Peninsula,

opposite Cyprus] “Holtz 1897” (print), “Type” (print on red), "Melecta

octomaculata Fr. Kl. As. u.” (in Friese’s hand). Holotype M. octomaculata Friese,

by present confirmation (MNB).

Further material. — USSR: | 4, sine loc., but with pinned gold disk [sic], & ‘*107626/corpulenta Mor.

Typ. 4”, in F. Morawitz’ writing (ZIL); 3 3, Uzbekistan, Okr. Kopala Semieretsj (illegible), 13.v.1910,

Sj..tn..kov (illegible), M. transcaspica Mor., det.? (ZIL); 1 9, Uzbekistan, Buchara occ., Hum-Kala

1892, Melecta robusta F. Mor., in F. Morawitz’ handwriting, M. transcaspica Mor., det.? (ZIL); 1 & 1 9

(diss., in perfect condition), sine loc., but with printed label “21”, both with Melecta spec. g, det. H.

Hedicke 1934 (NMW). — Iran: 1 & (diss.), Persien (written), ‘‘M. aegyptiaca Rad., m. Typ. vergl.

[ichen].” (Friese’s writing), det. Friese 1893, and ““Melecta not aegyptiaca”, in D. B. Baker's writing

(NMW). — Lebanon: | £ (diss.), Nd. Liban., Cedern b. Becharré, 1900 m, 3—6.vi.1931, Zerny (print)

(NMW); 1 & (diss.), Lebanon, Nr. Cedars Hotel, 6400 ft., 15.vi.1944, H. B. Cott (BM). — Israel: 1 9

(diss.), W. Jaar, Golan, 1.vi.1970, H. Bytinski-Salz (CBS); 1 9 (diss.), Palestine, Judean desert, Ma’ale

276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Haadumim (formerly Khan Chadrur), 200 m, 18.iv.1943, H. Bytinski-Salz, M. caesareana [sic] Fr., det.?

(CBS). — Turkey (Asia minor): 1 & (diss.), Turquie, 11—14 km avant Ciftehan-Ulukisla, 1150—1300

m, 28.vi.1976, R. Desmier de Chenon (INRA); 1 g (diss.), Turkey: Ankara, Ravli, 1000 m. 30.vi.1962,

Guichard & Harvey (BM); | 9, Turkey: Ankara, above Hasanoglan, 1500 m, 29.vi.1962, Guichard &

Asia min., Eski-Tshehir, vii.1906, Dr. Lendl (print) (MBUD); 1 9 (diss.), Asia min., Konya, colli

steppici ad W, 6.vii.1962, A. Giordani Soika (print) (MP); 1 9, Armenia, Monastero Chérard,

Beli, Kiefernstufe, 11—1200 m, 23.vi.1967, Reinig (SMF); 1 9 (diss.), Turkey, Konya, 22.vi.1969, W.

Linsenmaier (CL); 5 3, Türkei, Madensehir/Konya, 20.vi.1973, K. Warncke (CKW, ML); 1 ¢ (diss.),

Türkei, Yesilhisar/Kayseri, 19.vi.1973, K. Warncke (CKW); 1 g, SW Türkei, Sarkishla/Sivas,

Seidenstücker (CWG); 1 &, Asia min., Merrina, 1901 (MNB); 1 9, Kleinasien, Brussa, leg. Ihirk

(MNB); 1 g (diss.), Türkei, Icel:Sertavul, 1400 m, 9.vi.1978, Max. Schwarz (CMS).

A large, robustly built species.

Male. — Disk of labrum coarsely rugosely punctate, punctures large,

contiguous; basal tubercles low, rather shiny, with few large irregular punctures;

an incomplete obtuse median ridge present (a.o. distinct in lectotype) or wanting,

usually restricted to distal one-fourth or less and highest, ending in a raised

tubercle, at the upturned apical border, the latter feebly convex in frontal view.

Mandible-bases finely striatopunctate, the upper portion smooth and shiny, as in

most other Melecta. Disk of clypeus closely punctate, all punctures contiguous and

partly confluent, smaller than on labrum, those at sides more widely spaced,

striatopunctate on smooth ‘surface; silky white patch of decumbent hairs fan-like,

restricted to dorsal face, tips of hairs barely reaching anterior border. Antennal

segment 3 longer than its breadth at apex (100: 71) and a little longer than 4, which

is only slightly longer than broad, the succeeding segments squarish. Head above

closely deeply punctate, especially anteriorly in front of ocellar area and upon

middle on summit of vertex behind ocelli, where punctures leave no interspaces; a

broad line running from median ocellus to frontal carina as well as a fairly large

flat space adjoining (slightly posterior to) each of the lateral ocelli, smooth and

impunctate, the latter occasionally with few scattered punctures. Punctation on

mesonotum dense, irregular, punctures differing in size: small and contiguous all

around middorsal area, on which they are more irregularly distributed and partly

smaller than interspaces, the latter smooth and rather shining. Median mesonotal

line markedly impressed posteriorly but sulcus punctate; parapsidal lines poorly

indicated bij fewer punctures. Scutellum, parascutella, sides of thorax and

propodeum, dull, very closely contiguously punctate. A small, slightly lustrous

impunctate spot at mid-base of propodeal triangle invariably present, but varying

Figs. 202—214. M. transcaspica; 202, ventral view of tergite 7 (4 lectotype, Chodshakala); 203, sternites

7 and 8 (same specimen); 204, partial ventral and dorsal views of genital capsule (same specimen); 205,

external view of right gonostylus (same specimen); 206, the same (4, Persien); 207, ventral view of ter-

gite 7 (same specimen); 208, sternites 7 and 8 (same specimen); 209, partial ventral and dorsal view of

genital capsule (same specimen); 210, dorsal view of apex of tergite 7(¢, Lebaron); 211, apical portion

of sternite 7 (same specimen); 212, whole sternite 8 (same specimen); 213, sternites 7 and 8 (4, Tur-

key); 214, dorsal view of pygidial plate (9, no. 21, NMW). Scale line 1 mm (figs. 203, 208 and 213)

277

LIEFTINCK: Palaearctic Melecta

278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

in size and shape; lateral edges filled out with 6—8 short longitudinal rugae.

Scutellum but little biconvex, lacking distinct median sulcus; spines robust,

variable in size and shape, short and triangularly pointed in the lectotype, much

stronger and/or more or less curved in others, but in all implanted at a high level,

their dorsal face punctate, almost in line with basal part of scutellum. Legs robust,

integument dull, all segments closely, finely punctate. Hind femur with low, hairy

median longitudinal carina restricted to distal one-third; punctation very close on

smooth ground, diameter of punctures hardly less than interspaces, except

ventrally toward apex irregular and widely spaced on flattened, elongate-

triangular area. Mid tibial pad, fig. 10. Outer face of hind tibia rather convex,

whole surface coarsely, irregularly punctate with short, erect black spicules

abundantly interspersed. Hind basitarsus rather broad, widest at middle, shorter

than tibia, upper border straight, the lower somewhat convex; markedly

outcurved, the external face hollowed out, closely punctate, clothed with short,

fine, decumbent black hairs which are silvery at middle; no marked impunctate

infra-basal pit on hind basitarsus. (The shape and depth of the external hind

basitarsal concavity is rather variable: in lectotype and a male from Lebanon, for

instance, the joint is less outbent in posterior view than in a second Lebanese

specimen and one from “Persien”, while the length-breadth ratios also vary

somewhat (e.g. 100 : 32.5 in lectotype, 100 : 37 in a Turkish specimen. Yet all males

presently recorded are undoubtedly conspecific). Inner rami of mid and hind

tarsal claws relatively short, only half as long as outer and a little more broadened

basally than usual. Wings much more obscured than in many other species:

membrane of fore wing dark smoky brown only in basal portion of median cell, all

along anal vein as far as end of first discoidal cell posteriorly, in most of the anal

lobe, and along the dark central areas of the outermost cells. Abdomen above

rather dullish, distinctly less shining than in M. baeri, candida, aegyptiaca, and the

smaller-sized species pertaining to the group having short-haired hind femora.

Punctation dense and fairly strong, punctures smallest and most crowded together

on postgradular parts of tergites on which they are equal in size (though less

superficial) to those on disk of tergite 1; all punctures smaller than interspaces.

Gradular lines at sides of tergites 2—5(6) fringed thinly with rather long black

bristles, present also and more numerous at the corresponding sternal plates.

Sternites otherwise very closely finely punctate, the broad, almost impunctate and

smooth mid-posterior crescents accordingly more contrasting than in the equally

large-sized male of baeri; sternite 6 not projecting, simply rounded, its ventral

ridges hairy, very low and unapparent. Pubescence short, not at all concealing the

surface, the suberect tergal hairs just visible with a hand lens (x 10); lateral spots

snow-white, well defined, transverse, arranged as in pl. 4 fig. 22, the long raised

hairs at base of 1 arranged archwise, narrowest and thin medially, becoming

compact and spot-like laterally, the broad crescentic posterior area remaining

dark and short-haired. Spots on 2 transverse, somewhat comma-shaped, their front

margin characteristically a little incurved and less sharply delimited anteriorly

than behind, the apices of the spots pointing more or less inward; spots on 3—5

more nearly rectangular, decreasing gradually in size posteriorly, those on 5,

though small and subcircular, nearly always present. Exposed portion of tergite 7

LIEFTINCK: Palaearctic Melecta 279

(figs. 202, 207, 210) broad, trapezoidal, sides slightly outwardly convex or more

nearly straight in dorsal view, the lateral rims continuing all along border,

becoming gradually more swollen and upcurved toward apex; disk hollowed out

between a pair of low, closely approximated median ridges (rarely unapparent),

which diverge toward base of tergite; surface dull, irregularly striatopunctate and

sparsely hairy; oblique ventral ridges well developed, rather broad. Sternites 7—8,

figs. 203, 208, 211—213. Genital capsule large, 2.0—2.2 mm; gonocoxal enclosure

widely U-shaped, mesial borders of gonocoxites at first markedly concave, then

curving outward, the angle evenly and broadly rounded. Gonostylus thumb-

shaped, its dorsobasal process small, rather variable in shape, its base attached to

stylus for a fairly long distance (figs. 204, 206 & 209).

Female. — First recorded — and quite insignificantly described — as M.

caesareae Friese (pl. 4 fig. 24), a species which turns out to be synonymous with

transcaspica, agreeing in all respects with other females of the latter from Israel.

It differs from the male by having still darker wings, more deeply sulcate

mesonotal line, and shorter body pubescence. With honesta it shares the rich snow-

white abdominal ornamentation, in strong contrast with the dull, very finely

punctate abdomen; distinguished from honesta at a glance by the more normal (i.e.

shorter) length of the first abdominal segment.

The complications with reference to the taxonomic position and nomenclature

of transcaspica, are explained under corpulenta (p. 260). The type of M.

octomaculata Radoszkowski, from Egypt, is a wrongly sexed male, either of

aegyptiaca or transcaspica. For a transcription of the original diagnosis, see under

that species in the Appendix.

With the aid of the photographs of both sexes (pl. 4 figs. 22—24), the key

characters, and the structural details figured, this fine deep black, dark-winged

Melecta is one of the more easily recognized species in the Eurasian fauna. There is

little variation in the shape and size of the white body markings in comparison with

many congeners. The slightly curved, more or less comma-shaped transverse spots

on tergite 2 are characteristic for both sexes.

Distribution. — From Transcaspia (Turkmenia) and Iran in the east, ranging

westward into the eastern Mediterranean. Apparently fairly common where

found, especially in Asia minor, in which country transcaspica was met with by a

number of hymenopterologists not especially interested in bees.

Melecta solivaga spec. nov.

(fig. 215)

Type material. — Algeria: 1 9 (diss., fig. 215), with green disk (meaning caught

in June), with on reverse side ‘‘872/80”, and “Museum Paris” (print), ‘‘Ouargla

1894 Weisgerder” (written), i.e. oasis about 480 km S of Constantine. Holotype

(MP).

Female (holotype). — Body elongate, abdomen broadest across end of segment

2, tapering gradually, length-breadth ratio as 10 : 6. Labrum squarish, greatest

280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

breadth little before middle, at that point slightly exceeding its median length;

apex thin-margined with distinct crescent-shaped emargination; no median crest;

surface (smooth basal tubercles excepted) strongly punctate on shiny ground,

clothed with numerous very strong, raised black bristles. Face not very prominent;

clypeus squarish in frontal view, with numerous punctures smaller than somewhat

shiny interspaces, anterior border polished, impunctate. Head behind face more

closely punctate, all punctures small, subequal in size to their interspaces.

Antenna normal, 3 distinctly longer than 4 (ratio 10: 6), the latter subequal to next

segments, all a trifle longer than broad. Anterior surface of thoracic dorsum and

sides finely, contiguously punctate, punctures on posterior part of mesonotum,

scutellum and hindermost sclerites larger, deeper and isolated, those covering

dorsal convexities on each side of median sulcus scattered on brilliantly shining

ground; parapsidae also distinctly impressed distally. Transverse suture between

mesonotum-scutellum deeply recessed. Scutellum exposed, hairless, feebly

biconvex on either side of shallow concavity, rounded off posteriorly and but

slightly overhanging posterior sclerites; spines robust, triangular, flattened and

bluntly pointed, directed abruptly upward and backward. Legs moderately strong;

femora smooth and shiny, sparsely, not strongly punctate, fringed posteriorly with

few long raised hairs; hind femur with incomplete, blunt median ridge. Outer faces

of mid and hind tibiae and basitarsi clothed with appressed pubescence

interspersed with small, suberect spinulose setae and fringed with erect bristles,

which are longest posteriorly. Inner rami of all tarsal claws broadly triangular and

flattened, less than one-third length of main branch, claw-like. Tibial spurs

normal, all evenly and but slightly curved, outer spur on hind tibia only little

shorter than inner. Wing venation blackish brown; membrane of fore wing very

Fig. 215. M. solivaga, dorsal and left lateral view of pygidial plate (Q holotype, Ouargla, Algeria)

LIEFTINCK: Palaearctic Melecta 281

dark brown with low purplish blue reflections, subhyaline basal areas much as

described for the equally dark-winged fumipennis spec. nov., but brown slightly

lighter in tint roundabout cubital cell-centres and along distal margin; hind wing

also lighter brown. Venation normal; third submarginal cell a little shorter than

high. Integument of all tergites throughout extremely finely reticulate-punctate on

somewhat shiny ground, most superficially chagreened on postgradular areas;

sternal surfaces similar, except broad and brightly polished postgradular annules

of 1—4, but whole sternal plate 5 dull, caused by fine reticulation. Black and white

pubescent pattern well defined; hair covering head and thoracic segments longest,

though only partly concealing surface, black on mouth-parts, clypeus and summit

of head, with long tufts of white just behind antennae and along occipital border,

but hairs at temples and underneath all black. Thorax with conspicuous elongate

white patches dorsally in front of tegulae and a median tuft filling out mesonotal

sulcus, on each side of which are small, deep black spots of shorter raised hairs, all

hiding the surface; also small tufts of white just in front of black parascutella, large

patches of raised white hairs on each side behind wing bases, and slender white

tufts projecting caudad from beneath scutellar spines. Posterior sclerites and sides

of thorax black, save a smallish white central spot upon middle of mesepisterna.

Outer faces of mid and hind tibiae white in proximal half. Abdomen rather dull;

short pubescence covering tergites consisting of minute, closely set, black

decumbent hairs; sides of 1—4 marked with white: all spots placed far laterad in

regular row, those on | and 4 smallest, subtriangular or oval, respectively, those on

2 and 3 distinctly transverse, nearly rectangular, more widely distant across

dorsum than their own transverse diameter; all spots compact, depressed, made up

of not very long hairs. Sternites all black. Dorsal and ventral bristles fringing

graduli sparsely distributed and relatively short, but more numerous and longest at

sides of tergite 5. Pygidial plate triangular, basal two-third of disk finely

reticulated, gently convex and somewhat shiny, thereafter broadly sulcate

between slightly raised side margins and base of crest, which is smooth, more shiny

and rounded off (fig. 215).

Male. — Unknown.

The decision of giving a new name to a single female Melecta is mainly based on

the extraordinary shape of the pygidial plate of this species. The type is an aged

specimen with frayed wings and parts of the black tergal pubescence rubbed off.

Additional features, which may also serve to the recognition of the still unknown

male, can be summarized as follows. A large, dark-winged species with relatively

small, subrectangular white tergal spots, which at sides of tergite | are made up of

short hairs; hind wing nervellus (cu-v) coinciding with point of origin of Cu-M

fork; and great number of short, acuminate, spike-like setae covering outer faces

of mid and hind tibiae of female.

A peculiar bee, easily known also by its size and deeply impressed median

mesonotal sulcus, as mentioned in the key (p. 180).

282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Melecta baeri Radoszkowski

(figs. 12, 216—221, pl. 2 fig. 12)

Pseudomelecta Baeri Radoszkowski, 1865, Horae Soc. Ent. Ross. 3: 56, pl. 1 fig. 3 (col. pict., whole in-

sect, “Gouvernement d’Orenbourg”, vide Popov, 1955!). — Popov, 1960, Entom. Oboz. Akad.

Nauk SSSR 39: 239 (no type selection!). — Lieftinck, 1972, Tijdschr. Ent. 115: 282—284 (note on ge-

neric position).

Melecta baeri: Popov, 1955, Trudi Zool. Inst. Akad. Nauk SSSR 21: 325 (type not studied, but compar.

notes with M. corpulenta Mor. and nivosa Mor., all transferred to Melecta; type loc. of baeri discus-

sed). — Kokuév, 1909, Trudi Yaroslavskogo estest. — istor. Obshchestva [Proc. Nat. Hist. Soc. Ja-

roslav] 2: 105 (distrib. notes; see M. corpulenta).

Melecta baerii: Friese, 1895, Bienen Europa’s 1: 155—156 (key 9 gd), 168—169 (not seen! orig. descr.

Type material. — USSR: 1 9 (figs. 216—218, lectotype P. Baerii Rad., by

present selection), with pinned gold disk and labelled “Cauca Mlokos” (bad

capital print), “Baerii 94.30” (Radoszkowski’s writing), “ex coll. Radoszkowski

Inst. Zool. P. A. Krakow 25/27” (print), “Pseudomelecta baerii Rad., det. ex coll.

Radoszkowski” (print), “Typus” (written on red) (IZK).

Further material. — USSR: 2 g (both diss, figs. 12, 221, one mounted, pl. 2 fig. 12), “Turcomania,

Krasnowodsk” (print) (MBUD, ML); 1 9 (diss.), “Krasnowodsk” (written), ‘‘coll. F. Morawitz” (print

in Russian), “Melecta corpulenta F. Morawitz” (in Morawitz’ handwriting) (ZIL); 1 9 (diss., mounted,

Turkestan” (written, white disk), ‘M. baerii Radosz.?, det. C. Ritsema” (ML); 1 9 (diss.),

Krasnovodsk/Ahnger/158, M. albovaria Er., ex coll. & det. Pittioni (BM).

Doubtful material. — USSR: 1 3, small-sized (diss.), “Turcomania/Krasnowodsk” (print) (MBUD).

This conspicuous bee is a true Melecta. Though one of the earliest described

Eurasian species, its relation to others has given rise to much speculation and

confusion with similar large-sized members of the genus, like corpulenta F. Mor.

and transcaspica F. Mor. In an attempt to find out more about the status and

distribution of baeri, Popov supplied interesting information. The following

sentences are partly translated from Popov’s meditations expressed in his own

language. In a first article (1955), the author already concluded that baeri was

wrongly placed in Pseudomelecta, as is evident from the superficial description and

colour picture of this bee. He also pointed out that Kokuév (1909), while

discussing the distribution of baeri, was mistaken by referring three females from

Boz-Dara (leg. Shelkovnikov) to that species. In point of fact, these individuals

came from Krasnowodsk and — at least one of them — should be referred to M.

corpulenta F. Mor.: they were labelled as such by Morawitz himself. Friese’s

statement at the end of his description of Paracrocisa caesareae (1925), that baeri is

a short-haired bee, had to be rejected, because both the picture and description

clearly point to a more hairy species. Popov further explains that the original

habitat of baeri, viz. “Orenbourg government”, before its incorporation with the

Kirgisian district in 1868, extended very far toward the south and included the

Caspian and Aral lakes. He remarks in passing that Eversmann also understood

the region to be that extensive. In view of this and considering also the large size of

Radoszkowski’s female, Popov at the same time considered the possibility that

baeri could be identical with M. corpulenta or nivosa, a conjecture which later

LIEFTINCK: Palaearctic Melecta 283

proved to be untrue, though both species were correctly placed in Melecta. The

former is distributed from the Ryn Sands in the north (in coll. Morawitz) as far as

Bajgakum in the Syr-Darja (in coll. Malyshev); nivosa is rather smaller in size but

otherwise agrees more closely with baeri and is also known from Bajgakum (in

coll. Malyshev). So it appears that both occur in the southern part of the former

Orenburg government. The above statements can now be confirmed, in so far that

at least some of Morawitz’ females from Ryn Sands (Rynpeski) and Krasnowodsk

are indeed true corpulenta (see under that species). In a next publication, dealing

with Radoszkowski’s so-called “types” in the Zoological Institute at Krakow,

Popov (1960) argues that a “careful re-examination of the bees in that collection

described and/or illustrated by him as such, very often turned out to be not types,

for example his Pseudomelecta baerii and Andrena oulskii” (loc. cit: 239). Popov’s

hesitation to decide upon the status of M. baeri, thereby renouncing the selection

of a lectotype, can be easily explained by the lack of any available specimen

corresponding exactly with the original diagnosis and colour picture. Of course,

this only caused the existing uncertainties to persist, but at the same time justifies

the belief that Popov’s concept of baerii was less confused than were

Radoszkowski’s ideas in general. The matter was further complicated by the

existence of three more females, turned up recently in other collections, which

inevitably also deserve attention in the present context. Though resembling the

bees presently regarded as baeri, they are even more extensively white-haired than

any of the others. One of them, here called ‘spec. A”, was found in a consignment

of Melecta sent to me on loan recently by Dr. Konigsmann of the Berlin museum,

while two others (“spec. B”, one of which unnamed, see below) had apparently

remained unnoticed in the Berlin and Leningrad collections, respectively. The

neglected (or overlooked) ‘spec. A” shows a white pattern surprisingly similar to

Radoszkowski’s portrait of “baerii”’, so much so in fact that it might even have

served the artist as a model for that species! Unfortunately, it is exactly this

particular specimen “A” which lacks a clear locality record, the two identification

labels being also confusing. Apart from that, this bee is a good deal smaller than

the one described, viz. 13.5 mm as against 16 mm for baeri in the original diagnosis.

The two other females (‘‘spec. B’’), though undoubtedly conspecific, are again

different from all others. For these reasons, and also because the males are still

unknown, I am reluctant to describe them as new. Instead, it would seem enough

to make up the whole by transcribing their labels only, as follows:

bispinosa Evm.” (Eversmann’s writing), ,,PseudoMelecta Beeri [sic] Rad”.

(unknown handwriting), “Type” (print on red) (MNB). Obviously a distinct,

undescribed species (see above).

Aulis-Anak/Syr-Darja lake (orange) and “coll. Kokuev” (print); 9 (idem),

“Dsungarei [Dzunggarskiye, SE USSR frontier area] Sandwüste ... (illegible)

27.6.25, leg. Beik”, and “M. arm. v. grandis” (pencil, det.?) (MNB). These two are

undoubtedly conspecific, resembling “spec. A” and M. baeri nob., but differing

structurally from both. Like the former an undescribed species.

Summarizing the above, and assuming that Radoszkowski mixed up at least two

284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

species brought to his attention, I have decided to accept his own unmistakable

type labels as settling the matter, selecting the specimen in his own collection as

lectotype and leaving the identity of “spec. A and B” undecided.

The six good specimens here characterized (one male mounted and

photographed, pl. 2 fig. 12, and one female with gold disk and Radoszkowski’s

type labels), are very similar to one another and unquestionably conspecific. The

status of a much smaller female in the Budapest museum must remain doubtful.

M. baeri runs out in my keys to near candida (3) and grandis (9), but resembles

transcaspica most closely in general appearance. Differs from the last in details of

structure, by having lighter coloured wings, more regular abdominal maculae and

a more strongly outcurved hind basitarsus.

Male. — Labrum almost square, a fraction longer than broad, the widest point

slightly before halfway length; apical one-fourth with smooth median carina,

better pronounced and more sharply acute, the anterior border less distinctly

upturned, than in transcaspica. Disk of clypeus less coarsely, more superficially

punctate, punctures not contiguous, those at sides isolated and separated by more

than one puncture width on shiny surface; silky white patch flat, somewhat more

compact and broader, covering upper parts of sides as well, the hairs longer,

Figs. 216—221. M. baeri; 216, frontal view of labrum (@ lectotype, Orenbourg); 217, left hind tarsal

claw, oblique dorsal view (same specimen); 218, dorsal and right lateral view of pygidial plate (same

specimen); 219, the same (9, “Siberia occ.’’); 220, dorsal view of exposed portion of tergite 7 (¢, Tur-

comania); 221, sternites 7 and 8 (4 Krasnowodsk, Turkmenia), scale line | mm

LIEFTINCK: Palaearctic Melecta 285

reaching anterior border. Antenna slightly more slender than in transcaspica, all

segments including 3 a little longer (length-breadth ratio of the latter 100 : 60).

Head above closely punctate but most punctures smaller, more evenly distributed,

the interspaces where present dullish, finely tessellated, especially on paraocular

area, the impunctate spots in front of median and beside lateral ocelli small and of

irregular form; frontal line distinctly raised, subacute, running up as far as anterior

ocellus. Punctation and sculpture of mesonotum, scutellum and propodeum as

described for transcaspica, the interspaces (where present middorsally on

mesonotum) smooth and shining; scutellar spines robust though equally variable in

shape; no distinct median scutellar line. Legs shaped much as in transcaspica,

expect that all tarsal segments appear to be somewhat stronger, especially the hind

basitarsus being more markedly outcurved than in that species (length-greatest

breadth ratio 100 : 35.7); inner rami of mid and hind tarsal claws longer, about %

length of outer (fig. 217). Mid tibial pad, fig. 12. Hind tibia below fringed with long,

raised, widely spaced marginal bristles, the margins of hind basitarsus with shorter,

suberect setae; outer face of hind tibia clothed with depressed plumose white hairs

interspersed with thick black spicules. Wing membrane much lighter than in

transcaspica: hind wing entirely and fore wing for more than its basal half, hyaline;

a spot at extreme base, an elongate dot at apex of median cell, as well as the entire

marginal cell, rusty brown (with diffuse marginal extension distal to it), all the rest

of membrane but little infuscated (pl. 2 fig. 12). Abdomen definitely more shining

than in transcaspica, all setiferous punctures smaller, more widely spaced, the

integument accordingly more plainly exposed, than in that species. Black and

white vestiture much alike in the two species, except that the long white hairs

covering most of tergite I are all of the same length, not subinterrupted in the

median line; also the compact white lateral spots on tergites 2—S are more

regularly rectangular (cf. pl. 2 fig. 12 and pl. 4 fig. 22), those on 2 (—3) not at all

excised anteriorly or pointing inward. Sternites of all segments much more glossy

and at the same time less contrastingly punctate and hairy before and behind;

gradular bristles also shorter and more depressed. Sternite 6 as in transcaspica, but

ventral ridges rather longer and more protuberant, just visible in profile. Exposed

portion of tergite 7 broad, trapezoidal, sulcate, shaped similarly to transcaspica,

though differing markedly in that the longitudinal ridges are better marked off,

more widely distant, shiny and hairless (fig. 220). Genital capsule large, 2.0 mm

long, gonocoxal enclosure more narrowly U-shaped than in transcaspica, mesial

borders of gonocoxites at first diverging and a little concave, at about halfway

distance toward the angle parallel, then very little converging and finally again

curving outward, the angle slightly better pronounced and less evenly rounded.

Gonostylus somewhat shorter, broader basally, and more distinctly tapering

toward apex, which is either subtruncated or rounded, clothed with relatively

short bristles; dorsobasal process more abruptly angled at apex than in transcaspica

and attached to stylus almost at right angle.

Wing expanse of first described mounted male, 30 mm.

Female. — Differs from the male in the usual variegated black and white

pubescent pattern of head and thorax. In addition to the key characters, the

following details are taken from the lectotype and are worth recording.

286 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Mandibles wholly black, with the usual, though rather narrow, inner tubercle

about midway length. Occipital ridge with dense fringe of long white hairs curving

outwards on either side at the rounded hind angles; long pubescence lower down

along orbits and on temples, black. Mesonotum with pair of distinct, oval, black

dots placed in the long axis, separated by a distance greater than transverse

diameter of one spot. Median mesonotal line very fine but distinctly sulcate. A

pair of conspicuous white tufts, one each to the inside of and just behind middle of

tegula; large tufts of much longer and thinner white hairs beneath and partly

surrounding scutellar tubercles, the latter short, conical, directed slightly upward

and straight backward, much shorter than white tufts, which are almost coalescent

with equally long white hairs behind posterior wing base. Propodeum only laterally

beset with white hairs, which are directed sideways. Sides of thorax with large

white hair spot covering upper portion below mesospiracle, for the rest black.

Basal three-fifths of mid and hind tibiae white externally, the tips of not very

strong, black spike-like setae visible through decumbent white pilosity. Wing

membrane unusually light: only faintly smoky, especially along margin beyond

pterostigma. Abdomen short-haired, but tergal marks conspicuous, well defined,

the hairs longest and forming large semi-erect tufts at sides of tergite 1, those on

2—4 transverse, subrectangular, broadest on 3. Longish black bristles at graduli

and hind angles of tergites 2—5. Sternites all black, clothed with very short hair.

Length 16.5 mm, fore wing 12.5 mm, expanse 29 mm.

Three of the Krasnowodsk females still before me agree very nearly with the

lectotype, although in one of them black anterior mesonotal dots are wanting, the

pattern resembling Radoszkowski’s picture in this respect.

Melecta grandis Lepeletier

(figs. 9, 222—230, pl. 5 figs. 25—30, map 2, p 303)

Selected references

Melecta grandis Lepeletier, 1841, Hist. Nat. Ins. Hym., 2: 443—444 (pars, 9 only! Oran, ‘‘comme para-

site de l’Anthophora Espagnole” [= A. hispanica (F.)]. — Lucas, 1849, Explor. Sci. Algérie, 3, Hym.:

211—212 (descr. 9 3), pl. 9 fig. 1la—e (sex?, whole insect, mouthparts & ant.; Algérie, Oran, sec.

Lepeletier). — Pérez, 1883, Actes Soc. Linn. Bordeaux, 37 (secr. 4, t. 7): 303—305 (notes, Algérie).

— Gribodo, 1893, Bull. Soc. Ent. Ital. 25: 406—407 (? pars, notes, sine patria). — Friese, 1895, Bie-

nen Europ.: 159 (? pars, Südeuropa, Nordafrika). — Alfken, 1914, Mém. Soc. Ent. Belg. 22: 235 (flo-

wer record, Algeria).

Melecta luctuosa: Gribodo, 1924, Boll. Mus. Zool. & Anat. Univ. Torino, 39 n.s.: 39(9 Cyrenaica).

2 Melecta n.sp.? 6 M. luctuosa n.var.? Dusmet y Alonso, 1915, Mem. Real Soc. esp. Hist. Nat. 8 (7a): 330

over drawer label “M. grandis Lep. Oran, coll. St. Fargeau”; lectotype M. grandis

Lep. by present designation (MP); 1 ¢ (diss., figs. 9, 225—226), in bad condition,

left hind leg partly missing, Algeria, with old white written label “Oran” and green

printed museum label “coll. J. Pérez 1915”, first described (“allotypic”) & M.

grandis Lep. by present selection (MP).

LIEFTINCK: Palaearctic Melecta 287

right hind leg missing (BM). — Morocco: | & (diss.), Maroc, Moyen Atlas, between Azrou &

Timahdite, 1800 m, 8.vi.1973, M. A. Lieftinck (ML); 1 g (diss.), Maroc, Midelt, 24.v.1947, J. de

Beaumont (CB). — Tunisia: 1 & (diss., pl. 5 fig. 29, figs. 227—229), Médinin, 3.95 (green label), coll. J.

Pérez 1915 (MP); 1 9, Gafsa 1910, P. Chrétien (on green), 19.3 (red ink), Melecta luctuosa Scop., det.?

(MP);2 £ 1 9, Tunisi, Belv ledere), 12.3.1882, and Tunisi dint., Bab Alena, 8—10.3.1882, G.& L. Doria,

coll. Schmiedeknecht (MNB); | 9, Tunis merid., Melecta aegyptiaca Fr. [sic], ex coll. & det. J. D.

ML); 4 9, Cyrenaica, Cyrene, 1800 ft., 26.iii—3.iv.1954, K. M. Guichard (BM, ML); | 9, Tripolitania,

Uadi Caäm, 5 aprile 1953 (MA); 1 9, Cyrenaica, R. U. Agrario, Ain Raru, 10120, iii.1926, M.

1934 (MNB); 1 9 Frfd [Frauenfeld], Alex [andria], M. grandis, det. Kohl (NMW); 2 9, sine patria, Coll.

Palestine, Bir Rechme, 13.iii, H. Bytinski-Salz (CBS). — Spain: 1 9, Andls [Andalusia], coll. J. Pérez

1915 (MP); 2 9, Andalusien, Chiélana, 22—23.iii.1890, Korb (MNB, ML); I &, “Andalusien, 11.2.

Spain (prov. Murcia), Cartagena, J. Caceves (written), M. luctuosa Scop. & (unknown hand), Melecta

grandis &, det. Alfken, and M. luctuosa Scop. var. albovaria Er. 9, det.? ex coll. J. D. Alfken (MNB). —

Sicilia (Italy): 1 ¢ (diss., pl. 5 fig. 30 & fig. 230), Sicilia 1878, Mann (print), grandis, det. Kohl (NMW).

The female lectotype (pl. 5 figs. 25—27) is an old, worn individual with frayed

wings, its missing body parts having been eaten away by Anthrenus. Nevertheless it

is clearly recognizable as the specimen from Oran dealt with in the original

description of grandis. However, one of the supposed males, likewise from Oran, is

a misidentified example of italica Radoszkowski, photographed on pl. 6 figs.

33—34. This bears a vertically pinned blue disk, which would also prove its

origin from one of the old collections kept in the Paris museum (see under M.

italica). To avoid nomenclatural difficulties, it seems justified to consider the

second male from Oran a syntype, because this is an indubitable specimen of

grandis. On the other hand, I gather from certain comments upon the genus given

by J. Pérez in his publication (1883), that some at least of his presumed examples of

grandis are not that species but outsized individuals of M. albifrons albovaria, which

occurs all over southern France and in many parts of North Africa as well. This

inference was corroborated on a later occasion, when all specimens in the Paris

museum had been re-examined. Still arguing on the assumption that only a single

species was involved, — even true grandis being considered a variety of ‘‘armata’’,

— Pérez was of opinion that, like the last, considerable size differences also exist

in Algerian populations of grandis. That statement is undoubtedly wrong, because

we now know for certain that several of his specimens (including medium-sized

females) were misidentified, at least three distinct species having since been

recognized in the available material of “grandis”; moreover, exceptionally small

females of grandis are unknown.

Male (pl. 5 figs. 29—30). — Sufficiently characterized in the key. Head and

thorax dull, very closely contiguously punctate, except on middle of mesonotum.

Legs not very strong, closely finely punctate on somewhat shiny ground. Mid tibial

pad, fig. 9. Outer face of hind tibia reticulate-punctate, lacking impunctate areas

toward apex; inner face broadly sulcate and hairy along full length. Hind

288

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Figs. 222—230. M. grandis; 222, external view of mid tarsal claw (9, Zavia, Cyrenaica); 223, the same,

lateral view (same specimen); 224, dorsal view of pygidial plate (same specimen); 225, ventral view of

tergite 7 (first described “allotypic’ 4, Oran, Algeria); 226, sternites 7 and 8 (same specimen, scale line

1 mm); 227, sternites 7 and 8 (4, Medinine, Tunisia); 228, partial ventral and dorsal views of genital

capsule (same specimen); 229, external view of right gonostylus (same specimen); 230, sternites 7 and 8

(3 Sicily). Figs. 226, 227 and 230 drawn on the same scale

LIEFTINCK: Palaearctic Melecta 289

basitarsus long and slender, subparallel-sided, borders very little convex in profile

view. Genital capsule large and compactly built, 2 mm long. Gonostylus short,

broadest at base, tapering rapidly, tip bluntly rounded; dorsobasal process of

characteristic shape (fig. 229).

Female. — Labrum longer than its breadth at base but narrowing distad (ratios

as 100 : 75 : 54 approx.), distal border, gently upcurved, a little projecting and

swollen medially, as in male, whole surface deeply, coarsely rugoso-punctate

lacking interspaces; basal tubercles large, brilliantly shining with few large,

irregular punctures roundabout; raised bristles long, black. Head above closely

punctate on slightly shiny ground, as in male; no impunctate juxta-ocellar areas of

any size. Suberect hairs covering clypeus long, predominantly or wholly black,

some white only upon middle; distal one-third or less bare, closely punctate.

Antenna only little less strong than in male, segment 3 markedly longer than 4

(ratio varying between 100 : 75—86), and also longer than in male, but the

remainder scarcely more elongated than in the other sex. Scutellar tubercles

variable, usually short and straight, much shorter than surrounding long white

tufts. All thoracic sclerites, including sides of propodeum, hidden from view by

dense black and white pubescence, only centre of mesonotum, scutellum and

vertical postscutellar sclerites remaining visible under much thinner raised hairs.

Propodeal triangle dull, finely chagreened, smooth and somewhat shiny, usually

bisected by a short, fine median line. Wings obscure, fore wing darkest in distal

cell-centres, apex of radial cell, entire marginal cell, and along anterior border

beyond the latter; length of third submarginal about equal to its height. Abdomen

more expanded and flattened than usual, broadest about midway segment 2,

tapering rapidly and pointed. Hairs at basal edges of tergite | somewhat raised and

subinterrupted at middle by thin basal collar of longer erect hairs. Pygidial plate

rather long en slender, but less than twice as long as its width at base; sides straight

or feebly outcurved in dorsal aspect; margins hardly raised as far as the

constriction, apex expanded, slightly spatulate, with low median ridge; plate very

gradually a little downbent from base to apex, surface dull, very finely transversely

wrinkled, impunctate; colour dark brown, lateral margins and whole apex

obscured (fig. 224).

Affinity. — M. grandis is one of the larger, sturdily built and less common

members of the genus. Males are easily recognizable by a number of structural

characters while females can only be confounded with oversized individuals of the

allopatric M. tuberculata and some poorly known eastern species resembling it only

superficially. Like most others, it is an early spring species that has been observed

in the coastal districts of North Africa as early as the first week of February. It

occurs from near sea level up into the lower mountain zone to about 1800 m alt., at

which height males were still on the wing in June.

Bionomics. — The single male which I caught in Morocco was taken at about 10

a.m. while gathering nectar from the attractive lilac flowers of the Boraginacea

Anchusa azurea Miller, growing abundantly in clusters beside the road, in the hills

near Timahdite. The Melecta kept company with males and females of Anthophora

290 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

robusta (Klug), a big, fast-flying polylectic and locally common species with a very

long proboscis; I encountered this Anthophora in several places, a.o. near Midelt,

at the foot of the High Atlas mountains, a locality where Dr. J. de Beaumont had

also taken Melecta grandis on an earlier occasion, so the latter may well be a

parasite of A. robusta. Females of that species were collecting pollen and nectar,

while the males were observed (and filmed) hovering in front of the Anchusa

flowers, taking nectar in the same way to a fairly large Bombyliid and a very

peculiar, excessively long-tongued Nemestrinid fly (Neorhynchocephalus tauscheri

Fischer)'), with brilliant emerald green eyes. All three insects were jointly

extracting nectar from flowering Anchusa on the same spot. Unfortunately I failed

to discover the nesting sites of A. robusta.

I do not know whether Lepeletier’s remark regarding the possible host of M.

grandis in Algeria, the large and conspicuous Anthophora hispanica (F.), is indeed

founded on field observations made by his son, who first discovered the species.

Perhaps the statement is merely conjectural and based on the corresponding

superior sizes of these bees. Otherwise actual data on the parasite-host

relationship and their biology are quite unknown. It is of some interest to note that

A. hispanica has repeatedly been taken on Ibiza, of the Balearic islands, but has

never been observed in one of the more frequently visited islands of the group,

neither has M. grandis been collected in any of the Balearics.

For its distribution, see map 2.

Melecta prophanta spec. nov.

(figs. 18, 231—236)

Type material. — Canary Is: 1 ¢ (holotype, diss., figs. 18, 231—236),

Lanzarote I., Famara, 4—15.ii.1979, W. N. Ellis & R. T. Simon Thomas (MA).

Large species, total length 18 mm approx., fore wing 12 mm. Stature and

pubescent pattern most closely resembling M. transcaspica and baeri, but

abdominal markings even more transverse and prolonged inward.

Male (unique). — Labrum a trifle longer than its greatest breadth across basal

tubercles, a little narrowed toward apex (breadth ratios 100 : 67), anterior border

straight, angles broadly rounded. Squarish silvery patch lying flush upon clypeus,

the tips of long hairs exceeding anterior border. Antenna (figs. 231—232)

markedly shorter and thicker than in aegyptiaca, 4—12 square instead of somewhat

longer than broad, with larger, more deeply impressed rhinaria; long fringes on

either side of scape white. Ocelli closely approximated, distance separating them

less than one-half their own diameter. Punctation of head and thorax as in

aegyptiaca, but pubescence, though shorter on all parts, relatively long and

') Nemestrinidae are known to be parasitic upon the larvae of small Lamellicorn beetles, but in view of

the same semblance in behaviour, body size and striking co-operation noticed at the spot, this parti-

cular species might deposit its eggs at the nest entrances or within the burrows of Anthophora robusta

as well. The fly was kindly identified by my colleague, Dr. P. J. van Helsdingen, of the Leiden mu-

seum.

LIEFTINCK: Palaearctic Melecta 291

predominantly white, e.g.: raised tufts on each side of antennal scape, behind

antennae, fringes at occipital border, and most of thoracic sclerites; black are only

hinder part of mesonotum, scutellum and, narrowly so, metepisternum between

spines, as also whole posterior surface. Scutellar spines straight, slightly raised and

pricker-shaped, punctate, directed straight back, much shorter than long white

tufts beside each of them, the area between spines remaining only narrowly black.

Mid tibial pad (fig. 18) narrow, the white squarely cut off basally, with very few

black spicules and some longish posterior bristles. Outer face of hind tibia with few

raised spicules, the black posterior bristles short. Hind basitarsus wholly black,

slightly less than three times as long as its widest point about midway length (ratio

100 : 35.7), moderately outbent, outer face concave, evenly closely punctate,

short-haired, with few longish bristles at lower level posteriorly. Inner rami of mid

and hind tarsal claws about % length of outer. Tergites of abdomen distinctly more

shiny than in aegyptiaca, all punctures setiferous, much smaller and more

superficial, the setae shorter, more finely branched. Snow-white marks very

conspicuous, more approximated, compact and better defined, than in aegyptiaca,

the hairs markedly shorter; 2—5 all transverse, 2 and 3 three times broader than

deep, especially spots on 2 somewhat hollowed out anteriorly. Tergite 7 tapering

Figs. 231—236. M. prophanta (3 holotype, Lanzarote, Canary Is.); 231, antennal segments 3—8 (scale

line 2 mm); 232, antenna (scale line 0.5 mm); 233, dorsal view of tergite 7; 234, sternites 7 and 8; 235,

dorsal view of genital capsule (scale line 1 mm); 236, right lateral view of gonostylus, with dorsobasal

process

292 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

rapidly, apex narrow, slightly emarginate (fig. 233), the oblique ventral ridges low,

flattened and hairless. Sternites all black; ventral ridges at apex of 6 low, barely

visible in profile, densely black-haired, the enclosed median area elongate, with

sparse setiferous punctures on somewhat shiny ground. Sternites 7—8, figs. 234,

apices of both strongly setiferous. Genital capsule 2.3 mm long, gonocoxal angle

subrectangulate, evenly rounded; gonostylus much less rapidly tapering in full

profile view than shown in fig. 236, which shows the curious squarish form of the

basodorsal process in side view.

Doubtful female (unique). — Morocco: | 9 (worn, wing borders damaged),

High Atlas, Oukaimeden, 2600—2800 m, 8.vii.1976, J. Gusenleitner, M. luctuosa

albovaria, det. J. Heinrich (CG).

Stature, size and tergal markings very similar to male, length 17.5 mm approx.

Labrum shaped similarly, but with distinct, subacute, median crest extending

anterad for little less than two-thirds its whole length and terminating in a small

tubercle (barely indicated in male); anterior border slightly but distinctly concave.

Basal half of mandible smooth and shiny externally, impunctate (male, finely

longitudinally striato-punctate). Head above as in male, closely punctate, vertex

lacking impunctate areas. Antenna more slender, 3 one and one-fourth as long as

next flagellar segments, all being a trifle longer than broad. Mesonotum and

scutellum deeply, contiguously punctate, as in male, the median mesonotal line

not impressed, reaching back to a level halfway length of tegulae, very finely

bilineate; parapsidal lines similar, though considerably shorter. Scutellar tubercles

directed obliquely upward, triangular, very slightly curved and punctate, as in

male. Propodeum dull, closely punctate throughout. Legs strong, size normal;

distal portion of inner faces (behind median carina) of mid and hind femora almost

impunctate, the carina of hinder pair distinct, almost complete, subobtuse. Wings

only little darker than in male; distal side of third submarginal slightly less

markedly angled. Abdomen much as in male, tergites smooth and shiny, finely

superficially punctate (microsetae rubbed off on disk of tergites). Pydigial plate

shaped much as in festiva, about twice as long as its width at base, almost straight in

profile, only the broadened apex a little downbent; disk flat, very finely

transversely wrinkled, with few scattered punctures at extreme base only; colour

black. — Pubescence moderately long and dense (rubbed off in places); depressed

patch of longish white hairs upon middle of clypeus not nearly reaching anterior

border, raised upon whole vertex, and fringing occipital border. Mesonotum with

the usual white areas and a pair of subrectangular black spots anteriorly; scutellum

mostly black, a few long white hairs only below the tubercles; sides and under

surface black, upper portion of mesopleurae with large patch of white. Legs

moderately hairy, as in most species; robust, suberect, shiny macrosetae at outer

faces of mid and hind tibiae interspersed with longish bristles and short decumbent

fine hairs. White are: basal one-third of fore tibiae and about basal half of mid and

hinder pair; for the rest all black. Compressed inner rami of mid and hind tarsal

claws about two-fifths as long as outer. White lateral marks of abdominal tergites

2—4 very similar in shape and size to those of male, all transverse spots broadest

on 3, about three times broader than deep; spots at sides of 1 isolated,

subquadrangular, a little diverging posteriorly, consisting of longer, somewhat

LIEFTINCK: Palaearctic Melecta 293

tufted, suberect hairs, the remaining spots compact, decumbent, though not raised

above level of integument. Sternites fringed with longish black bristles at gradular

lines; no white hairs.

This conspicuous new species superficially resembles other fairly large-sized

congeners, like aegyptiaca, baeri, festiva and tuberculata, but can be distinguished

from all by a combination of characters. Its nearest ally would seem to be

aegyptiaca. Apart from the differently shaped copulatory organs, the male

disagrees with the latter by having much more enlarged white tergal spots, a

shorter labrum, stronger antennae, broader and more outbent hind basitarsi, and

by the very differently shaped 7th tergite. An additional feature separating the two

is the sculpture of the outer face of the broad basal portion of the mandible, which

in aegyptiaca is smooth and shiny with a few scattered punctures distally, whereas

in prophanta this part is wholly finely striato-punctate. The differences between

prophanta and the other species just mentioned are best understood by consulting

the key and illustrations. It is impossible to associate the male of prophanta with the

female of my “lindbergi”’, described from the same island of the group, but which

has now proved to be synonymous with aegyptiaca, the male of which is wholly

different from that of prophanta. The same applies to the supposed female of the

latter, of the high mountains in Morocco, which is immediately distinguished from

aegyptiaca by the shape of its pygidial plate. Hence we are confronted with two

evidently related species occurring side by side in the same island of the north-

eastern group of the Canaries. This is surprising and certainly needs confirmation.

It is of interest to note that Dr. S. Erlandsson recently sent me a single pair of yet

another Melecta, taken in Gran Canaria (San Bartolome, 9.iv.1973, leg. T. Palin),

representing an undescribed species. I am reluctant to characterize these as new,

because both specimens are in too poor condition to serve as types of a new taxon.

At all events these discoveries indicate that the bee fauna of the Canary Islands is

of high quality and unexpectedly rich in species. For some further remarks, see

under M. caroli (p. 324).

Melecta festiva spec. nov.

(figs. 6, 16, 237—265, pl. 6 figs. 31—32, map 2, p. 303)

Melecta luctuosa var. albovaria subvar. eczmiadzini: Friese, 1895, Bienen Europa's: 163—164 (key: Kau-

kasus).

Type material. — France: 1 & (diss. fig. 237, holotype) and 1 9 (diss.,

paratype), Callian (Var), 22.v.1963, W. Linsenmaier (both ex coll. Linsenmaier,

ML).

Further material. — Switzerland: I & (diss.), Wallis (Valais), Vesperterminen, S of Visp, 1250 m,

8.vi.1972, on Thymus serpyllum (figs. 16, 244), M. A. Lieftinck (ML); 1 9, Wallis, Useigne, 26.vi.1924,

Th. Steck (NMB); 2 9, Valais, Sierre, vi.1949 and v.1950, W. Linsenmaier (CL & ML). — Italy:1 9,

Piemonte, with square label / Susa (written) 108, ‘‘Melecta punctata F. Lep. notata Illig. Klug. Ped. 9”

(old writing), Spinola collection, supplement (MT); 2 & (one diss., figs. 241, 247), Tirol, Bozen 1886 F.

Kohl, M. luctuosa v. albovillosa (sic) and v. albovaria Er., det. Friese 1893 (NMW). — France: | 9,

Nyons (Drôme), coll. J. de Gaulle (MP); 1 9 (figs. 239—240), Vallouise (Htes Alpes), ca 400 m,

294 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

18.vi.1939, ‘‘/uctuosa, white variety with convex plate anale”, P. F. M. Verhoeff (ML); 1 9 (fig. 238)

Bses Alpes, v.1891, coll. J. Vachal (MP); 1 ¢ 3 2 (diss.), Provence, Callian (Var), 17—18.vi.1930 &

8.vi.1931, Th. Steck (NMB); 1 &, 1 9, Callian (Var), 10.v.1959 & 2.v.1963, W. Linsenmaier (CL); 1 g,

Les Arcs (Var), 18.iv.1939, W. Fassmidge (BM); 1 &, Beauvallon (Var), 7.iv.1967, O. W. Richards

(BM); 1 & (diss., figs. 242, 245), Grimaux (Var), 15—24.iv.1968, G. Barendrecht (MA); 1 G (diss.), St.

Aegulf (Var), 22.v.1954, W. Linsenmaier (CL); 2 9, Cap Camerat (Var), 5.v.1976, H. Wiering (MA); 1

9, Massif de Moures (Var), 8.v.1965, ‘‘flew together with 2 g 2 9 Habropoda tarsata (Spin.) along

Figs. 237—248. M. festiva; 237, right hind tarsal claw, oblique dorsal view (¢ holotype, Callian, S Fran-

ce); 238, the same (9, Basses-Alpes, S France); 239—240, right mid (239) and hind (240) tarsal claw (9

Vallouise, Htes-Alpes, S France); 241, dorsal view of tergite 7 (g, Bozen, Italy); 242, sternites 7 and 8

(3, Grimaux, S France); 243, the same (4, Castilia, Turkey?); 244, apices of sternite 8 (upper) and 7

(lower) (&, Vesperterminen, Wallis); 245, partial ventral and dorsal view of genital capsule (¢, Gri-

maux, S France); 246, dorsal view of left gonostylus (same specimen); 247, lateral view of left gonosty-

lus (4, Bozen, Italy); 248, dorsal view of pygidial plate (9, Gard, S France)

LIEFTINCK: Palaearctic Melecta 295

loamy wall at roadside near overgrown excavation”, S. J. van Ooststroom (ML); 1 9, Montauroux

Q (diss, fig. 248), Mas Méjean (Gard), 29.v.1962, H. Wiering (MA) | 9, La Fontaine du Buis (Gard), 3

km S, 14.v.1971, R. Desmier de Chenon (MP); 1 9, Vernet-1.-B. (Pyr. or.), Col de Juell, 7.vi.1963, W.

H. Gravestein (MA). — Spain: 1 9 (diss.), Arganda, SE of Madrid (= Montarco olim), 11.v.1964, W.

Linsenmaier (CL); 1 &, Cuenca, Motilla, 8.iv.1955, I. H. H. Yarrow (BM). — Portugal: I ¢ (diss.),

Sobreiral (probably nr. Famalicäo, 20 km NE of Oporto, M. Diniz in litt.) (MUC). — Albania: 1 9

(diss.), Alban. Exp. 1918, Hodzha bei Prizren, 15.v, M. luctuosa, det. Maidl (NMW). — Romania: 1 &

(diss.), Mehädia (E of Belgrad), leg. Pavel/17/1, Melecta albovaria Erichs., det. Mocsáry, luctuosa v.

albovaria calabrina Rad., det. Friese 1893 (MBUD). — Bulgaria: 1 9 (diss.), Sliveu, Pavlasz (MBUD).

— USSR: SW Russia, Gruziaya ? [Georgia]: 1 ¢ (diss., pl. 6 fig. 31 & 259—260), “Kauk. 1885”

(written), “M. eczmiadzini Rad. mit Typ. vergl lichen)”, det. Friese 1893, “Typus” (print on orange),

labels of H. Friese, with additional label “holotype M. alecto spec. nov. in litt. Lieftinck” [not M. alecto

spec. nov, huj. op.!] (MNB); | g (diss.), S. Russia, Crim (written), Sig. Alfken (MNB); 1 9, Armenia,

18 km. Erevan, 11.vi1.1963, A. Giordani Soika (MP); 1 2 (diss.) 39 (id.), Turkestan, Gouldscha/Ferghana

labelled in identical handwriting ‘‘Mesopotamia Malatia 1886” (err. pro Malatya, in E central Turkey!)

and ‘‘Mesopotamia Castilia 1886”, respectively, 3 moreover with “12” (written on red) and ““74b/125”

(black-rimmed cadre), the 9 with ‘‘74b/100”’ (same cadre), £ with Melecta luctuosa v. albovaria Rad.,

det. Friese 1893 (both MBUD; 1 & (diss., figs. 256—258), Sultan Dag, 1500 m, 22.v.05, ex coll. C. & O.

Vogt (MA); 2 & (both diss.), Turkey, Mut, Sertavul, 1300 m, 7.vi.1968, and Asia minor, Cardak,

1—2.vi.1967 (5 9, fig. 265), 7.vi.1968 (1 9), same loc., 1600 m, 22.v.1970 (1 2), all J. Gusenleitner (CG,

23 1 9 (diss., figs. 253—255), Türkei, Ostl, Sirnak/Siirt, 4.vi.1977 (9), E. Uludere Hakkari, 5.vi.1977,

Turquie, Ulukisla, 1500 m, 1 km avant Col de Caykavak Gegidi, “fond de combe très riche en fleurs”,

As. Türkei, Ürgüp, 17—19.vi.1976, J. Heinrich (CJH); 1 g, Asia minor, Beysehir See, NW Ecke,

5.vi.1964, H. Hamann (CMS); 2 Q(diss.), Turkey, Erzurum, 20 km Ispir-Ikizdere Rd., 1700 m,

1 —2.vi.1962, Guichard & Harvey (BM); 1 g(diss.), Turkey, Erzurum, 15.vi.1967, H. Oxizek (CKW); 1

Q(diss.), Turkey, Ankara, Polatli, 800 m, 2.v.1962, Guichard & Harvey (BM). — Rodos I (Greece): 2

(ML); 4 9, 10 km N of Malona, 11.iv.1970, A. C. & W. N. Ellis (MA). — Israel/Jordan: I Z(diss.),

Jerusalem, 800 m, 20.iii.1975, K. M. Guichard (BM); 1 2 (diss.), Judaean Highlands, Artüf (?),

12.iv.1923, P. A. Buxton, M. ashabadensis Rad., det. B. Uvarov (BM), | 9 (diss.), Eshtaol, Kesalon

Bytinski-Salz, M. aegyptiaca, det. Mavromoustakis (CGS); 2, Ramat Gan, 6.iii.1942, M. luctuosa var.,

det. Bytinski (CBS) and same loc., 4.v.1951, P. M. F. Verhoeff (ML); 1 9, Jericho, 9.iv.1943, H.

loc., 1700 m, 11.vi.1976, D. Simon (1 g), same loc., 1800 m, 11.vi.1976, A. Freidberg (1 3), same loc.

at Satureia, leg. Kugler (CBS); I 9, Palestine, Jerusalem, 25.v.1937, J. Wahrman (CBS). — Greece: 3

& (all diss, figs. 249—252, Graecia, Peloponnes, Chelmos, 1900—2100 m, 1—4.vi.1962, Max Schwarz

Peloponnes, Athen, Acropolis, 21.v.1962, H. Hamann (CMS); I & (diss.), Euboea, Steni, iv.1926, Holtz

(MNB); 2 ¢ (diss.), Griechenland, Euböa, Stehni, 800 m, 4—11.v.1956, and Frioni, Olymp, 1000 m,

296 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Daan & V. van Laar (MA); 1 9, Cumani, Brenske, M. luctuosa Scop.? det. Friese 1893 (MBUD); 1 9

(diss., fig. 255), Graecia, Peloponnes, Vytina, westlich Tripolis, 20.v.1962, W. Linsenmaier (CL).

Doubtful material. — USSR: 2 2 (diss.), Süd-Turkestan, Digai, 18.vi.1913, leg. K. Küchler (SMF,

(NMW). — Crete: 4 9 (identity uncertain), Creta Bird, Herakleion 1906 (3 9) & Anoya, 2.vii.1906 (1

(MA) and Cyclades (MCG).

A richly spotted species, averaging a little smaller than tuberculata but

resembling that species fairly closely in general morphology and pubescent

pattern.

Male. — The following differences are worthy of note. Maxillary palpi (both

sexes) 6-segmented, relative lengths of separate segments variable, but 2—S

usually subequal and always longer than 1 and 6. Antennae as described for

tuberculata except that segment 3 is relatively shorter and only little longer than 4.

Punctation on disk of mesonotum coarser, the punctures slightly larger, lacking

interspaces and even more or less coalescent in places; scutellar spines variable

though frequently a little downbent and generally longer than in tuberculata. Mid

tibia relatively shorter, less markedly broadened from base to apex; outer face of

hind tibia rugose, more coarsely tessellate-punctate, with 9—12 irregular black

spicules shining through the pubescence. Hind basitarsus similarly outbent but a

little shorter and less slender than in tuberculata, about three times as long as the

greatest breadth at a point slightly beyond half-way its length; surface reticulate-

punctate but, like tuberculata, mostly concealed from view by short, appressed

white hairs. Inner ramus of fore tarsal claw about three-fourth length of outer,

those of mid and hind legs much shorter, at most a little over one-third length of

main branch (fig. 237); outer faces of remaining tarsal segments white-haired.

Colour and neuration of wings much as described for tuberculata: shape of

submarginal cells equally variable and not appreciably different, except that the

fore wing membrane is somewhat darker (in the female, on the contrary, usually

less obscured than in tuberculata!). Abdomen more gradually tapered posteriorly,

the paired white pubescent spots on 2—5, though about equal in size and shape,

accordingly more closely approximated, than in tuberculata. Integument of all

tergites still more shining than in the latter, the setiferous punctures finer, more

widely spaced, the raised hairs consequently somewhat shorter and more sparsely

distributed.

Tergite 7 shaped similarly to tuberculata, sculpture of exposed portion equally

variable: coarsely striato-punctate, but median area frequently slightly hollowed

out, transversely striated, occasionally feebly ridged on either side of the middle;

apex subtruncated, smooth and bare, the hind border brownish, shallowly

emarginate, a little swollen with upturned rounded angles; ventrally, the folded

apical rim carries a pair of curved, converging ridges similar to those seen in

tuberculata. Like the tergites, the whole surface of the sternites 1—4 is distinctly

more shining, the punctation finer, more superficial, than in tuberculata, leaving

broader and almost impunctate posterior margins on 2—4; pubescence shorter

LIEFTINCK: Palaearctic Melecta 297

+ AS

RER.

A EN A aN

4 nt

iR Î Veh”

1 | a ;

/ Vis on

Y DT

/ =

F Si

249 |; N

KIN

EE EEE

es:

{

Va 3

\ \

7 \ / \

E ER k, 5

Abe i} val 255 À EN

if AIN

Figs. 249—255. M. festiva (SQ Greece); 249, external view of left hid leg (Z, Chelmos); 250, ventral

view of tergite 7 (same specimen); 251-252, sternites 7 and 8 (2 Z, same locality; scale line 1 mm);

ponnesus)

and less dense, hair fringes in front of gradular lines also thinner, but white

submarginal tufts frequently present at either side of 3—4 (or 5). Posterior border

of sternite 5 straight, surface closely punctate; exposed portion of sternite 6

bluntly triangular in outline, as in tuberculata, closely punctate, the median area

elongate, somewhat hollowed out, thinner and lighter in colour than the side

portions, the latter forming a pair of low, unapparent ridges placed in the long axis

and not visible in profile. Sternal plates 7 and 8 as in figs. 242—243. Basal arms of 7

more broadened, hollowed out and incurved than usual, the anterior margin of

each, about halfway its length from base, with a tiny recurved process; median

plate subparallel-sided and fringed on either side with many setae; its apex

298 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

variable in outline, either rather abruptly truncated with subcircular and narrow

emargination (Wallis, and one from France, fig. 244), or a little longer with more

rounded side-edges and more deeply emarginated (France and Spain, figs.

242—243). Sternite 8 relatively broad in all dissected specimens, up to twice as

broad as its median length, apex suddenly narrowed and a little excised (same

figs.), but in a male from Wallis a little shorter and broader and also more deeply

excised than shown in the sketches. All dissected specimens agree in the form of

the genital capsule and gonostylus: in dorsal view the inner apical angle of the

gonocoxite is sharply pronounced, hooked inward, but the angle itself more or less

rounded; gonostylus not very long and perfectly straight, laterally compressed

with obliquely truncated apex, which in some specimens is slightly emarginate, the

inner and outer (basodorsal and basoventral) processes shaped as shown in figs.

246 —247. Genital capsule (incl. gonostylus) of moderate size, 1.7—1.8 mm.

Female. — Sexual dimorphism less pronounced than in many other species, the

alternating black and white pattern of the thoracic dorsum not as markedly

contrasting is in species like grandis, baeri a.o., the anterior mesonotal black dots

diffused. Labrum squarish, broadest about middle, distal half usually with distinct

carina terminating in a small tubercle, the anterior border slightly emarginate.

Median mesonotal line not impressed but evenly, finely raised; parapsidal lines

short, often obliterated and replaced by an irregular area with few punctures.

Sculpture of propodeum obviously also variable, frequently throughout coarsely,

closely punctate, but triangle occasionally with finely chagreened, somewhat

shiny, trapezoidal impunctate area in front of propodeal pit, its presence or

absence apparently quite independent from locality (e.g. examples from S.

France). Sides of posterior sclerites (black in grandis!) clothed with abundant long

white hairs. Much longer white tufts behind wing bases almost fused together with

smaller ones projecting from beneath scutellar spines, which themselves are

directed obliquely upward and backward, though varying in shape and length.

Inner face of hind femur more sparsely punctate than in grandis, setiferous

punctures rather deep and of different sizes, those toward blunt median ridge

becoming scattered on more shiny ground, carrying long erect hairs. At least basal

two-thirds of outer faces of mid and hind tibiae white, those of tarsi usually also

more or less white; tips of all black spicules visibly projecting from the soft

pubescence. White tergal spots placed more inward and much larger than in

grandis, those at sides of 1 subcircular, with many of the raised hairs at outer edges

shorter than in male, these same spots often almost connected across middle at

base by a thin collar of erect hairs, similar to grandis. Spots on 2—4 more compact

than in male and distinctly transverse, broadest on tergite 3 and subequal in width

to the black interspace. Suberect gradular bristles at tergal sides and sternites

quite distinct, only little shorter than in grandis.

Body size very variable. Some measurements are: ¢ holotype, length 13.8 mm,

fore wing 10.7 mm, wing expanse 24 mm; 9 paratype, 15 mm approx., and 10.5

mm, respectively. Further specimens, ¢ length 11—16.5 mm, for wing 8—11 mm;

25—26.5 mm, respectively.

LIEFTINCK: Palaearctic Melecta 299

With good series of either sex, taken together in one locality, discrimination

between festiva and tuberculata (next to be described), offers no great difficulties,

because the sexual characters of each, particularly those of the males, are quite

specific. Without uncovering the pygidial plates of the females it is, however, less

easy to separate them, so that a number of features apparently only of secondary

A LL

Wr ale :

i SK

SSS SSS

256

Figs. 256—265. M. festiva; 256, external view of left hind leg (4, Sultan Dagh, Turkey); 257, ventral

view of tergite 7 (same specimen); 258, sternites 7 and 8 (same specimen); 259, ventral view of tergite 7

(3 syntype M. eczmiadzini, Caucasus); 260, sternites 7 and 8 (same specimen); 261, dorsal view of tergi-

te 7 (Z, Sertavul/Mut); 262, sternites 7 and 8 (same specimen); 263, sternites 7 and 8 (4, Cardak, Tur-

key); 264, partial ventral and dorsal view of genital capsule (same specimen); 265, dorsal view of pygidi-

300 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

importance, such as the shape and armature of the legs, presence or absence of

white-tufted scutellar tubercles, etc., must also be taken into consideration.

Infraspecific variation is considerable in this species. For instance, on comparing

the sketches figs. 237—248 of morphological structures, with figs. 249—265 of the

same details, some striking differences can be observed between the shapes of the

hidden sternal plates of the male. In a general sense, the former are taken from the

more typical, western populations of festiva, the latter from those occurring in the

eastern parts of the Mediterranean and Turkey. As a matter of fact, however, only

the specimens from Chelmos in the Peloponnesus (S Greece) are a little different

from the rest by their smaller average body size (only 12 mm); unfortunately no

females were taken with them. All other specimens of more eastern distribution

are indistinguishable from western examples. As to the drawings compared, it

must be said that these are selected extremes, to demonstrate differences as well as

local conformities (cf. e.g. figs. 252—254), many intergradations existing.

Distribution. — Discontinuously, from Iberia, S France (terr. typ.), S

Switzerland, NW Italy and, scatteredly, through Greece and the Balkans eastward

via Turkey probably far into Transcaspia and Turkestan (see arrows on map 2).

Melecta tuberculata spec. nov.

(figs. 266—274 & pl. 6 figs. 35—37, map 2, p. 303)

Type material. — Rodos I. (Greece): 1 ¢ (holotype, with Ophrys (?) pollinium

upon middle of frons) | 9, Fileremos, 300 m, 19.iv.1971, M. A. Lieftinck (ML); 4

Fileremos (lalissos), 21.iv.1970, D. C. Geijskes (ML). The latter specimens of

either sex are paratypes.

Further material. — Rodos I. (Greece): 1 ¢, Petaloudes, 500 m, 22.iv.1971, M. A. Lieftinck (ML); 1

9, Epta Piges, ca. 50 m, 7—8.v.1971, M. A. Lieftinck (ML); 10 ¢ (diss.), Profitis Ilias, 800 m, 3.v.1971,

M. A. Lieftinck (ML); 8 ¢ 2 9, Profitis Ilias, 800 m, 20.iv.1970 (6 9), Ataviros, 18.iv.1970 (2 3),

Rodini, ca. 150 m, 22.iv.1970 (2 9), all H. Teunissen (CT, ML); 1 g 4 9, Fileremos, 23.iv, Epta Piges,

Kalithea, 1.v, Faliraki, 25.iv.1976, all H. Teunissen (CT, ML); 7 g (diss.) 4 9, Rhodes, Aegios Isodores,

Q, Stadt Rhodos, Oertzen (MNB); 1 ¢, Rhodus, Hedenb. (NRS); 1 3, Rhodos, Profitis Ilias, 8.v.1975,

H. Malicky (CG); 1 9, Rodi, 17.iv.1928, M. luctuosa Scop. 9, det. Hedicke 1931 (IEB); 1 9, Rhodos,

Chanea, 24.iv.1942, Kl. Zimmermann (MNB); 3 &, Kreta, Omalos Ebene, 1000 m, 26.iv.1942 &

11—16.vi.1942, Kl. Zimmermann, one with M. aegyptiaca Rad., det.? (MNB) | g, Ost-Kreta,

Hochebene Nidha, 1418 m, 11.v.1925, A. Schulz, M. armata v. albovaria Er., det. Alfken (MNB); 2 9,

Ost Kreta, Iraklion, (Gandia), 29.iv.1925, and Kloster Apésanas, Ep. Kantergion, 20.v.1925, A. Schultz,

Kusdas (CG); 5 9, Amnissos beach and Anogia-Axos, 16—23.iv.1972, D. C. Geijskes (ML); 1 g (diss.),

Kreta, Hersonissos, 14.iv.1976, M. van Lookeren Campagne (coll. R. Leijs); 1 9, Crete, Katharo,

Lassethe, 4000 ft., D. M. A. Bate (BM); 2 3, Crete, Arkhanes, 28.iv.1972 and 2 9, Chania, 15.v.1972 &

1.v.1974, K. M. Guichard, one & with 2 orchid pollinia attached to clypeus (BM); 1 g, Creta, v.O., ex

LIEFTINCK: Palaearctic Melecta 301

Mitki, 13.iv.1971, leg. Malicky, K. Kusdas (CJH); 1 & (diss.), Arkanes, 4.v.1972, at Borago officinalis, J.

van der Vecht (ML). — Greece (mainly continental): 1 9, Chalkis, Euboea, iv.1926, Holtz (MNB); 1 &

1 2 (diss.), Athen, Lange 1872, M. luctuosa v. albovaria Er., det. Friese 1893 (NMW); 1 & (diss., fig.

270), Athen, Akropolis, 12.v.1963, Max. Schwarz (CMS); 1 9 (fig. 274), Athen, Melissia, 31.v.1966, W.

Linsenmaier (CL); 4 g 1 9, Graecia, Delphi, 2.iv.1966, W. Grünwaldt, and 1 9, Souli, 5.v.1973, W.

(NMW); 1 9, Olympia, Schmiedeknecht (SMF); 3 9, Griechenland, Olympia, Prioni, 1000 m,

3—13.vi.1956 (9), Gorgopotamos, Oitigebirge, 800 m, 26—28.v.1956 (9) and Euböa, Stehni, 800 m,

4—11.v.1956 (9), all Fr. Borchmann (MKB); 3 9, Greece, M. Veluchi [Velouchi], Emvritania, 1000 m,

15—19.vi & 12—1400 m, 13.vii, Holtz (MNB); 12 g, Centr. Greece, Pilion, Portaria, 3—4.vi.1971, J.

van der Vecht & P. M. F. Verhoeff, flower 51 (ML); 1 9, Greece, M. luctuosa Scop., det.? ex coll. Vogt

acq. 1960(MA); 1 9, Macedonia centr., Shar-Planina, Vratnica, 900 m, 21.vii.1956, F. Daniel (ZSM); 1

3.vii.1958, R. Loberbauer (CG); 1 9, Peloponnesus, Altkorinth, 3.vi.1963, J. Gusenleitner (CG); 8 9,

Peloponnesus, Xylokastron, 28.v.1966 (9), 18 km südlich Tripolis, 4—6.vi.1961 and 19.v.1966 (4 9),

Korinthos, 9—23.v.1962 (3 9), all W. Linsenmaier (CL, ML); 11 9, Peloponnesus, Kalamata,

Kythrion foothills, 6.iv.1950, N. Waloff, with cluster of 8 bright orange orchid pollinia adhered to

crassicornis Fr., det. Enslin, coll. Pillich (MBUD); 1 9, Cyprus, 9.iv.1924, M. luctuosa crassicornis Fr.,

det. Mavromoustakis (MCG); 1 g 6 9, Limassol, 14.iii.1924, 15.111.1927 (¢ M. aegyptiaca, det.?),

3.iv.1928, 15.iii.1931, and 1.v.1932, all G. Mavromoustakis, M. /uctuosa crassicornis Fr., det.

22.11.1932, E. E. Green (BM); 4 ¢ 3 9, Cyprus, Antifanitis, 12.iii.1971 (2 &) and Dhavlos, 10.iv.1971 (2

Karakurt, Arastal, 23.v.1975, Kl. Warncke (CKW). — Lebanon: 1 g, Syria, Beirut/12, Stoll,

M. aegyptiaca, det.? (MNB). — Israel: 1 g, Jerusalem, 25.v.1937, at Satureia, leg. Kugler, and 1 &,

Burgas, Arkutino (Black sea coast), vi-vii.1970, K. Bleyl (CJP, ML). — France: 1 g, Gallia mer.,

Dohrn, Coll. Makl (three printed labels), M. punctata, det.? (MH); | 9, sine loc., coll. J. Pérez 1915

(MP). — Spain: 2 g, Hispania, M. aegyptiaca, det. Friese 1904 (MNB); 1 9, Spain, Burgos (Castilia),

Aranda de Duero, 1.vii.1973, Z. Boucek (BM); 1 & (diss., figs. 271—273 & pl. 6 fig. 35), S. Spain, Jaen,

Vadillo de Castril, S. Cazosla, Mateu-Cobos leg. (ex coll. Verges, ML); 1 9, S Spain, Andalusia, with

A robust black species with well defined white markings, clear-winged (3), or

with fore wing much obscured (9). The following descriptions are based on

specimens of either sex from the Greek islands of Rodos, Cyprus, Crete and

Naxos, which are considered “typical” and very similar to the male from France

and the pair from Spain. Individuals from continental Greece are generally darker

on head and thorax segments than the rest, the female also having darker wings.

Male. — Integument deep black, only mandibles distally often more or less

brown. Labrum subrectangular, only little longer than its width at base, surface

except on top of basal tubercles, very coarsely punctate, punctures partly

302 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

coalescent and often with feeble indication of a median crest about halfway

length; disk slightly concave, free margin not raised, straight or shallowly

emarginate. Maxillary palpi usually 5-segmented, but a short 6th segment

frequently also present; relative lengths variable, 1 and 6 usually the shortest and

3—4 generally a little longer than 2. Mandible-bases smooth and shiny, sparsely

striato-punctate. Clypeus but little convex, closely punctate. Head for the rest

densely, rather deeply punctate, punctures lacking interspaces and often partly

confluent, except a small area just beside each of the lateral ocelli. Antenna

strong, reaching back to end of tegula, scape cylindrical, broadest apically and

somewhat curved, much longer than 3, which is about one and one-third as long as

4, flagellar segments from 3 onward square or almost so; rhinaria distinct on

segments 3—13, elongate on 3, circular on succeeding segments, all deeply

impressed. Punctures on disk of mesonotum deep, circular and of one size, the

interspaces mostly less than the diameter of one puncture; anteriorly, on either

side of parapsidae, scutellar area, propodeum and pleurae the punctures are still

more crowded and coalescent in places; propodeal triangle finely punctate and

frequently partly tessellate or glabrous. Tegulae superficially punctate on basal

half, more scatteredly so distally, surface either smooth and shiny or minutely

striato-tessellate. Scutellum convex, median sulcus shallow, tubercles short,

variable in length and form (occasionally a little curved), usually nipple-shaped but

frequently vestigial or barely indicated. Femora and tibiae not unusually expanded

or armed; mid tibia evenly broadened to beyond halfway length and slightly

convex above; few apical spicules, dorso-apical tooth short, spur almost straight.

Hind tibia normal, outer surface convex, evenly, finely reticulate-punctate lacking

spines, its apex truncated, with some thick marginal setae and very short, bluntly

triangular dorso-apical tooth; spurs normal, the inner one longest, gently curved.

Tarsi thin and slender, depressed, hind basitarsus subparallel-sided, little

expanded, about twice as broad as mid basitarsus, slightly but distinctly outcurved

and 3.4 times as long as its greatest breadth about halfway length, apex truncated

with short dorso-apical tooth and row of strong setae; outer face somewhat

hollowed out, reticulate-punctate but surface mostly hidden under pubescence.

Inner ramus of all tarsal claws about % or a little less length of outer, all slender,

evenly curved and acuminate. Wing veins dark, second and third submarginal cells

variable in shape and length, the third relatively short, not or scarcely longer than

high, but apex strongly angulated; membrane of fore wing subhyaline, the apicial

border gradually enfumed (pl. 6 fig. 35). Abdomen relatively broad, subcordate,

tergites only moderately convex, considerably broader than high, segments

tapering rapidly posteriorly; dorsal surface very shining, punctures small, widely

spaced and superficial, successively more sparsely distributed and finer toward

apex of tergites, and all bearing erect black hairs well visible under low

magnification in fresh examples. Distal portion of tergite 7 rather broad, the

exposed part gradually narrowed from base to apex, which is truncated with

rounded angles, the hind border brownish, a little swollen and bare, usually

shallowly emarginate and slightly upturned; surface flat or somewhat hollowed out

above, proximal portion striato-punctate and clothed with appressed brownish

hairs; in ventral view the folded rim carries a pair of convergent elevated ridges

303

LIEFTINCK: Palaearctic Melecta

saloads DJ2a/aP 9WOS JO SPA093.1 AJI[P907

‘7 dew

VIVINIHAANL *

VAILSAJ ®

SIGNVYD MX

304 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

(fig. 268). Sternites dull, whole surface of all segments finely closely punctate, the

postgradular punctures more widely spaced and smaller than the rest. Posterior

border of sternite 5 slightly, shallowly emarginate; 6 closely microscopically

punctate, sides of exposed portion converging, tapering gradually and bluntly

pointed; this median triangular area continues basad to form an almost

rectangular flat area on each side of which arises a conspicuous ridge-like tubercle

placed in the long axis of the sternite and well visible in profile; these tubercles are

highest apically, reaching back as far as the base of the median triangle, and are

clothed with dark bristle-like hairs; the tubercles are rather shiny and bare in worn

individuals (fig. 267). Sternal plates 7 and 8 somewhat variable in shape and

setoseness, even in specimens from one locality; examples are shown in figs.

269—271. Genital capsule of large size, measuring 2.0—2.4 mm (incl. gonostylus)

in the dissected males; gonostylus short, broadly thumb-shaped, apex strongly

bristled; dorsobasal process small, more or less triangular, ventrobasal process

broad, ridge-like, carrying a dense row of strong setae (fig. 273).

Pubescence. Labrum clothed with rather long and stiff, closely set hairs; stiff

hairs fringing lower margin of mandibles of great length, white and/or black.

Clypeus with brilliantly shining pad of long, depressed silvery hair entirely

concealing the surface, all hairs directed straight forward, not surpassing anterior

border; laterally with long black bristles. Summit of head (vertex and postocular

area) with few short black hairs, pubescence on remaining parts long, erect, pure

white but often mixed with black on paraclypeal and upper part of genal areas as

well as on orbits posteriorly. Antennal scape with short hair in front and behind,

but at either side along full length with tuft of very long raised hairs which are

longer than greatest diameter of scape. Thoracic pubescence conspicuous,

forming a very broad collar of long, dense and raised white hairs with a silvery

gloss; this broad collar extends onto the sides all around tegulae as far back

dorsally as a little beyond the latter posteriorly, the middorsum (inclusive of most

of the scutellum and the sclerites behind it) is black-haired; this transverse dark

area suddenly appears narrower in the median line by the presence of fairly well

defined tufts of long white hairs around the scutellar tubercles, similar white tufts

being present also at the sides of the propodeum and above the hind coxae; ventral

parts likewise white, including the leg bases. Femora black-haired anteriorly, but

fore and mid pairs fringed behind with much longer white hairs, most

conspicuously so on mid femora at which they sometimes attain a length greater

than the breadth of femur; fringe at hind femora thin and much shorter, either

wholly black or white only along distal portion. Inner faces of all tibiae black, fore

tibiae with long white posterior fringe; mid tibia clothed very densely with silky

white pubescence from near base to near apex, except an isolated linear intrusion

of black at anterior border, the raised hairs along posterior border short and

sparse, black. Hind tibia densely white-haired, including posterior carina, but

those at extreme base and postero-apical one-third (or more) black. Outer faces of

hind basitarsus hidden from view by dense, appressed, finely branched, silvery

white hairs, except at extreme base and, narrowly, along posterior border, where

hairs are black. Remaining tarsal segments all white-haired exteriorly, except fore

and mid basitarsi, which are more shining and partly black basally. Abdomen

LIEFTINCK: Palaearctic Melecta 305

Figs. 266—274. M. tuberculata; 266, oblique dorsal view of hind tarsal claw (4 9, Fileremos, Rodos);

267, apex of abdomen, showing tubercles, left lateral view (4, same locality); 268, ventral view of tergi-

te 7(¢, Naxos, Greece); 269, partial view of sternites 7 and 8 (same specimen); 270, sternites 7 and 8

(3, Acropolis, Athens); 271, sternites 7 and 8 (4, Vadillo, Spain); 272, partial ventral and dorsal view of

genital capsule (same specimen); 273, external view of right gonostylus (same specimen); 274, dorsal

view of pygidial plate (9, Melissa, Athens)

306 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

clothed with short, semi-erect black hairs, the disk of postgradular portions in

worn specimens almost or wholly naked; basal part of | profusely covered with

long, raised white pubescence right across tergite, though longest and most

condensed at the sides; sides of 2—5 each with dense, subrectangular, snowy-

white marginal spot composed of decumbent hairs, these spots widely apart and

successively a little smaller from before backward, their limits straight or (usually)

slightly concave anteriorly and a little convex posteriorly, th pair on 5 placed

away from the side margin and separated by a distance suvequal to their own

diameter. Sternites with very short depressed black hairs, 2—5 just before

postgradular line moreover with rather dense fringe of much longer, slightly

raised, bristly setae, which are directed caudad, black, with few white ones

interspersed on 3—4. Vestiture of apical sclerites as described above.

Female. — The long and slender, gradually downbent pygidial plate is one of the

characteristics assisting in the recognition of this sex. It is a more stoutly built bee

than the female of festiva, from which it is most easily distinguished by the lack of

white tufts around the scutellar tubercles.

Variation. — As mentioned before, individuals collected simultaneously with

unquestionable conspectfic males in Bulgaria, central Macedonia and various

parts of Greece (mostly in the Peloponnesus), are noticeably darker than

anywhere else. This applies to both sexes, all of these having much darker, grey-

brown wings. Moreover, the light pubescence on head and thorax in the male is

not pure white but has acquired an ashy grey tint, while in the female most of the

white hairs on these parts are deep black with traces of silvery white only upon

middle at base of clypeus, at the thoracic sides, and behind the wing bases. In

melanistic extremes even these light hair spots may disappear completely; in them

also, at least half of the external white hairs and spots on the legs are replaced by

black, which gives these obscured populations a most peculiar appearance. This is

the more interesting because typical examples from the adjacent islands and more

eastern countries (Crete, Rodos-Kos, Turkey, Cyprus, Palestine), resemble each

other closely in being much lighter.

(The stout body form of tuberculata is not too well brought out in pl. 6 fig. 35,

which shows a more slenderly built male from Spain).

Size variable, averages slightly larger than festiva; 3 holotype, length 16.5 mm,

fore wing 11.7 mm, wing expanse 32 mm approx.; 9 paratype, 17.2 mm and 12

mm, respectively; other specimens (3g 9), length 12.8—18.5 mm, fore wing

10—13.5 mm; wing expanse (pl. 6 fig. 35), 28 mm; other specimens (¢ 9), 27—33

mm.

Distribution. — Discontinuously, in the west ranging from Iberia to S France

(very rare); then again throughout the eastern Mediterranean as far as beyond the

Black Sea and Israel (see map 2). Unknown from Italy and the Tyrrhenian Is. !

M. tuberculata is evidently the species arranged in the Paris museum collection

over the drawer label grandis, but which was called M. bituberculata J. Pérez, a

nomen nudum in the unpublished catalogue: — “Bône 1 ¢ voisin de M. punctata.

307

LIEFTINCK: Palaearctic Melecta

wy 00S

saloads D/92/2W 2Wos Jo susayyed uonnqginsiq ‘€ de W

07 Sk TE OI 0

po: RÉ VNINYOVL 1 8 VOINVLI

LTÉE VHONAHYOONAT 3 V211VLI

2 VNINYOVL'

VHONAHHO9NI11

308 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Anus faiblement échancré, lobes arrondis, peu ponctué à peu près comme 982 (=

pygialis J.P. ou nigripennis Lep.?); plaque anale inférieure portant 2 tubercules

triangulaires”.

Bionomics. — This is also one of the Melecta whose males were observed more

than once to play a role in orchid pollination, and of which females sometimes

exhibit the peculiar habit of collecting earth clumps at the legs (see chapter on

mud-collecting, p. 135).

Melecta italica Radoszkowski, 1876 & 1893, and leucorhyncha Gribodo, 1893

In the next pages I shall have to deal with the status and nomenclature of two

nearly similar — though not necessarily closely related — polytypic species whose

taxonomic and systematic position has so far remained a complete mystery.

Although the original description of italica was published 17 years in advance of

Gribodo’s /eucorhyncha so that there are no nomenclatural difficulties involved,

Radoszkowski’s second treatment of italica appeared almost simultaneously with

that of /eucorhyncha in a much more comprehensive and significant article. The

two names were validly proposed for two common and widespread species

inhabiting vast expanses of land surrounding the Mediterranean Sea, but the status

of both has been ignored ever since they were described and named. As might be

expected as a result of this negligence, subsequent writers gave new specific and

varietal names to them (and a number of non-related bees as well), so that the

necessity of studying the primary types and clarifying the nomenclature of all,

became obvious. Therefore I take this opportunity to straighten out the question

of priority in nomenclature in the two publications mentioned above. This could

be definitely settled in December 1963, by Dr. A. C. Townsend, the then librarian

of the General Library of the British Museum (Nat. Hist.), London, who informed

me at length about the dates of publication. It became clear that the seven (!)

specific names proposed for Melecta by Radoszkowski have priority over the three

varietal names introduced by Gribodo, antedating the latter by two or three weeks

(Dec. 11 and 31, 1893, respectively). The confusion in the nomenclature is briefly

summarized under italica, first to be discussed.

The analysis has been impeded from the outset by a variety of difficulties. Owing

to the complicated nature of their variation and the irregular distribution pattern

of the various populations, discrimination is sometimes decidedly difficult. If local

populations of each had shown greater specific diversity of structure, separation

would have been easier. As it is, however, they exhibit considerable infraspecific

variation not only in the extent of white maculations, wing colour and venation,

but also in numerous details of the male genital organs and form of the female

pygidial plate. Consequently, the characters employed in the keys are few and had

to be generalized for both sexes. All the same, a preliminary investigation led to

the conviction that italica and leucorhyncha are distinct, polytypic species with

broadly overlapping ranges. However, the degree of their geographic variation is

different. While leucorhyncha has developed a melanistic subspecies, taormina

Strand, confined to Italy and the big Tyrrhenian Isles (terr. typ. Sicily), all forms

occurring to the west and east of this central part of the Mediterranean are more

LIEFTINCK: Palaearctic Melecta 309

profusely white-spotted and hardly differ among themselves in this respect.

Nominotypical italica, on the other hand, shares with /. taormina a distinctly

melanistic pattern, a form also restricted to Italy and the Tyrrhenian islands;

elsewhere, however, it exhibits greater plasticity and becomes differentiated into a

number of irregularly distributed clinal populations, the instability being so great

that segregation into subspecies proved impossible. For the present I have

contented myself with the key characters and an enumeration of all material at my

disposal, giving full details of localities where populations of the two taxa occur

together. They could be told apart only tentatively on direct comparison by

weighing one set (or “sum total”) of characters against another. My present

observations on the various phenotypes are merely factual, no attempt having yet

been made to explain the astonishing coincident or near-identical distribution

patterns which so often go hand in hand with analogous colour designs (see map

3). A detailed analysis of all local populations, each on its own, must be reserved

for the future.

Melecta italica Radoszkowski

(figs. 5, 15, 78, 275—294, 358—359, pl. 6 figs. 33—34, pl. 8 figs. 38—39, map 3, p.

307)

wrongly sexed = ¢; ¢ = Eupavlovskia spec., probably E. obscura obscura (Friese). Radoszkowski,

1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 182, figs. 36a-c, g genit. — Lieftinck, 1969,

Tijdschr. v. Ent. 112: 111 (note on synonymy).

Melecta grandis: Lepeletier, 1841, Hist. Nat. Ins. Hym. 2: 443—444 (pars, ¢ only! not M. grandis Lep.,

Q Oran, Algeria).

Melecta quadripunctata Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 184 (9 Ita-

lie). Syn. nov.

Melecta luctuosa Scop. Var. I Meridionalis mihi, Gribodo, 1893, Bull. Soc. Ent. Ital. 25: 409, 410 (9 &

partim! “Piemonte: Torino, Susa; Francia: Montpellier Germania: Mecklenburg; Algeria.” Syn.

noy.

Melecta luctuosa Scop. var. albovaria: Friese, 1895, Bienen Europa’s 1: 163 (“3 Südeuropa’).

Melecta armata var. grandis & M. luctuosa: Alfken, 1914, Mém. Soc. ent. Belg. 22: 235 (pars, ¢ 9 Alge-

rien).

Melecta luctuosa Scop. var. acutivalvis Alfken, 1914, Mém. Soc. ent. Belg. 22: 235 (pars, & only, Alge-

rien). Syn. nov.

Melecta luctuosa var. crassicornis nom. nov. pro M. albovaria Friese, 1895 (nec Erichson), Friese, 1921,

Archiv f. Naturgesch. 87A (3): 168, 176, diagn. 39 “Sideuropa; Chanzik, Amanus Mts. (SE Tur-

key) & Taurus cilic.”. Friese, 1922, Zool. Jahrb. Syst. 46: 207 (repeated remarks, 3 Kaluckova, Ma-

kedonien; “sonst mehrfach aus Südeuropa erhalten, auch von Chanzik im Amanusgebirge (Nordsy-

rien), Tölg leg.” Syn. nov.

Melecta luctuosa var. crassicornis n.var.: Friese, 1925, Konowia 4: 28 (repeated remarks, “Balearen,

Type and syntypic material. — Italy: 1 ¢ (antennae and tarsi partly incomplete,

genitalia dissected out and glued on card by Radoszkowski, relaxed and redrawn

by present author, figs. 275—276), labelled “Italie” (Radoszkowski’s ink writing),

gold disk, “9” (error! print on lilac), ““italica/Type” (print on red square),

“Melecta italica Rad. type” (Dr. Enderlein’s writing). Lectotype M. italica Rad., by

310 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

ink-written grey label, Radoszkowski’s writing?), “Type” (print on red), “Melecta 4

pustulata Type”, Dr. Enderlein’s printed label and handwriting (MNB). Lectotype

M. quadripunctata Rad. by present designation. — Tunisia: 1 & (diss., genit.

examined), labelled “Tunisi dint. I—II—1882, G. & L. Doria” (small print),

Melecta luctuosa Scop. var. meridionalis Grib. & D. Gribodo” (Gribodo’s writing

on pale blue), collezione Gribodo” (print). Lectotype & M./. var. meridionalis

Gribodo by present selection (MCG); 1 9 (diss., fig. 287), labelled “Tunisi Belv.

ledere] 12.111.1882, G. & L. Doria” (small print), “Melecta luctuosa Scop. var.

“Collezione Gribodo” (print). Paralectotype (syntype) M. I. meridionalis by

present indication (MCG). — Algeria: | ¢ (diss.), with printed labels “Alger, Bab

el Oued”’, and written on reverse side ‘‘8.iii.10” “Dr. J. Bequaert’, “Sig. Alfken”,

‘“Melecta luctuosa Scop. f. acutivalvis Alfk.” in J. D. Alfken’s handwriting.

Lectotype M. acutivalvis Alfken, by present selection (MNB). — Corsica: I &

(diss.), labelled “Corsica, Ajaccio, 3.1902, Guglielm., Melecta luctuosa (erased), v.

crassicornis Friese” (Friese’s writing), “Type” (print on dark red). Evidently

holotype? a. crassicornis, selected by Friese (MNB).

Further material. — Italy (continental, S to N): 1 & (diss., fig. 15, pl. 8 fig. 38 & figs. 281), Italia mer.

Zeller (ML); 1 4, S. Italien (Puglia), Lecce, 10.iv.1963, Kl. Warncke (CKW); | &, Brindisi, J. Sahlberg,

Calabria, Antonimina, 1905, Paganetti(NMW); 1 9, Monteleone, coll. Magretti (MCG); 1 g (diss., fig.

278), Monticchio (Basilicata), 4.v.1892, M. luctuosa Sc., det.? (MT); 2 & (one diss.), Vulture

(Basilicata), 20.ii.1890 & 20.vii. (sic) 1891, both with M. armata Pz., det.? (MT); | & (very small),

M. luctuosa var. ruthenica Rad., det. Alfken (IEP); 2 4, Portici, coll. Magretti (MCG); 5 g 1 g (diss.),

Mai 1896 Andriai (?), M. luctuosa v. ruthenica Rad., det. Alfken 1904 (MNB); 1 9, Lazio, Puerta di

Castelfusano, 15.iv.1949, Bisleti leg. (CMC); 1 9, Roma, 8.iv.1927, Giordani Soika, M. luctuosa v.

calabrina Rossi, det.? (CGS); 1 & (diss., figs. 278, 280, 282), Lazio, M. Cimino, 600 m, 18.iv.1949, M.

Comba (CMC); 1 9, Lazio, Marino, 19.iv. & 1.v.1934, M. armata, det. Stocklein 1955 (ZSM); 3 9,

Lazio, Marino, 19.iv. & 1.v.1934, leg. Hartig (INER); 1 9, Capo Circeo, Quarta media basso,

1872, Mann no. 153, M. luctuosa ruthenica Rad., det. Friese (NMW); 3 & 6 9, Castiglioncello nr.

Livorno, 3—11.v.1958, G. Barendrecht (MA, ML); 1 ¢ 2 9, Rimini (Emilia-Romagna), 22.iii.1894

(39) & x. (sic)1893 (9), A. Tosi, one pair with ‘“M. armata Pz. 3 9 et varietates”, det.? (MH, ML); 1

9, Dintorni di Rimini, 3.7 (sic). 1890, Zanghi (ex coll. Pittioni, BM); 2 9, Cattolica [18 km Rimini],

2.v.1958, W. Grünwaldt (CVZ) & 22.iv.1963, Kl. Warncke (CKW); 1 3, Bologna (Emilia), M. Calvo,

30.iv.1931, M. luctuosa, det. Hedicke 1931 (IEB); 4 9, Bologna, Gaibolo, 3.v.1953 & Torren Ravone,

M. leucorhyncha taormina Strand, 10.viii (sic) 1886; one 3 9 with M. luctuosa Sc., one g with M. armata

Pz., det.? (MT, ML); 1 9, Is. Giglio, iv.1900, G. Doria (MCG); 1 9, sine loc., M. luctuosa var., det.?

3 1 9 (fig. 277), Genova dint., Borzoli, 11.iv.1909, G. Doria, “su fiori Brassica oleracea’ (MCG, ML); 2

2, 36/G Borzoli, v.1883, G. Doria M. luctuosa Scop., det. Gribodo (MCG, ML); 1 g, Sestri Levante,

LIEFTINCK: Palaearctic Melecta 311

7.iv.1906, Dr. Uzel (NMW); 1 9 Lago di Garda (Lombardia), S. Vigilio, 13.iv.1942, leg. Hartig (INER).

— France: | 3, purple disk, Tarbes (written; H.-Pyren.), coll. J. Pérez 1915 (MP); 1 g, Tarbfes?], 4.93,

(INRA, ML); 1 g (diss.), Provence (MNB); 1 g (diss.), Provence, St. Rémy, 9.iv.1933, A. Naville (CB);

1 2 (diss.), “Camargue 39” (B.-du-R.), S. Kelner Pillault (MP); 1 g, Marseille (B.-du-R.)... 1848

(written), Schmiedeknecht vid. (print), v. Heyden (print) (SMF); 1 &, Marseille, coll. Ernest André

1914 (MP); 1 g, sine loc., coll. J. A. Snyder, 20.iii.1931, M. luctuosa Scop., det. Schmiedeknecht (sic,

ML); 2 3 3 9, Le Trayas (Var), 16—21.iv.1933, G. Barendrecht (MA, ML); 1 g, Le Lavendou (Var),

24.iv.-5.v.1956, G. Barendrecht (MA); 2 g (diss., fig. 284), Grimaux (Var), 15—24.iv.1968 & 1 g, same

(MP); 1 & (diss.), Cannes (Alpes-Mar.), 4.90, coll. J. de Gaulle 1919, M. luctuosa, det.?; 1 9, Cannes, E.

Saunders coll., M. armata Panz. (pencil writing) (BM). — Corsica: 5 & 5 9, Corsica, Klinckow (NRS,

O. Sichel 1867 (MP, ML); 1 9, Corsica, M. calabrina Rad., det. Friese 1893, and M. luctuosa calabrina,

det. Friese 1904 (MNB); 1 4, Corsica, Melecta luctuosa v. calabrina Rad., det. Friese (MNB); 1 9,

Corsica, Ajaccio, 5.1901, Guiglielm., coll. A. Weis, M. luctuosa var. crassicornis Friese, det. Friese 1900

(SMF); 3 9, Corsica, Ajaccio, 5.1901, 3.1902 and 9.1902, Guiglielm./ 8—9 (MNB, ML); 2 9, Ajaccio,

10.3.06, coll. J. Vachal (MP); 1 9, Corsica, env. Zonza, 8—900 m, Giglio, 30.v.1967, M. A. Lieftinck

(ML); 1 9, W. Corsica, Ajaccio, Parata road, 15.iv.1938, G. Kruseman (MA); 1 9, Ajaccio, 5.iv.1956,

H. Wolf (ML). — Sardinia: | g, Sardinien, Tissi stat. (?), 7.iii.91, M. luctuosa crassicornis Fr., det.

Friese 1904 (MNB); 2 9, Sardegna, Sassari-Ploaghe & Sassari-Sorso (the latter together with | 9 leuc.

(BM); I &, Sicil./722 (print), Sicula N. Sicil. Zelller] (written), and 1 9, Sicil./725 (written), M. luctuosa

v. crassicornis, Fr., det. Friese 1896 (MNB); 1 9, Sicilia, Staud[inger] (MBUD); 1 9, Sicile, coll. O.

Guichard (BM); 1 9, Sicilia, Agrigento, v.1959, C. v. d. Voorn (ex coll. Barendrecht, MA); 1 9 (diss.),

Sicilia, Taormina, Sirinatal, 12.iv.1971, K. Kusdas (CG); 1 9, Sicilia, Zappulla, Aprile 1914, coll.

(NMW, ML); 1 9 (diss.), Sicilia Mann 1858, M. grandis det. Kohl, together with differently identified 9

of M. leucorhyncha taormina (NM W); 1 9, Sicilien, Agrigento, 23.ii.1977, M. luctuosa det.? (CMS); 1 9,

(INER); I 4, Sicilien, Agrigento, 23.iii.1977, M. Schwarz, M. luctuosa, det. Schwarz (CMS). — Malta:

2 4, Malta, 90—126, and Malta, iv.1927, H. C. Harford, M. luctuosa, det. Uvarov (BM). — Balearic

Is., Mallorca: 1 & (diss.), Mallorca, Soller de Mallorca, 1956/57, J. Briedé (MA); 1 g (diss.), Mallorca,

2.4.83, H. Friese, M. luctuosa var. crassicornis, det. Friese 1900, coll. A. Weis (SMF); 2 9, Mallorca,

(sic), det. Friese 1893 (MBUD, ML); I 4, “Sideuropa” [probably Mallorca!] (written on green), ‘‘720”’

(MG); 1 g, Mallorca, Albufera, Breit leg. (NMW); 1 g, Baleares, Palma de Mallorca, coll. Ernest

André 1914 (MP); 3 3 1 9, Mallorca, Son Moro, Monacor, 15.ii.1894, M. luctuosa Scop., det. A.

Compte Sart (CCS, ML). 1 9, Mallorca, Palma, 23.3.56, A. Compte Sart (CCS). — Ibiza: 1 9, Ibiza,

“Behaarung d. 5. Stern. dichter u. kürzer”, and “‘crassicornis Fr.”, det. Alfken (MNB). — Spain: 1 &,

Espana, Cadiz, Castellar de la Frontera, 19.iii.1968, R. Straatman (MA); 1 9, Cadiz, Guadiaro,

4 3, C. Spain, Valladolid, Granja Escuela, Camino Hondo, 26.v.1972 (2 g), and two with addition

“talud I”, 26.iv.1973 (2 3) (CAV); 1 & (diss.), Spanien (two written labels) (MNB); 1 9, Andalusien,

“M. grandis Lep. hisp.” in L. Dufour’s handwriting (MP); 1 9, Zaragoza, 19.7.02, L. Navas (ML); 1 3

(diss., fig. 285, pl. 8 fig. 39, mounted), NE Spain, Llobregat (Catalonia, prov. Barcelona, MP); 2 9

Canet [de Mar? nr. Barcelona], 4.93, E. Saunders (BM). — Portugal: 1 9, W. Portugal, Coimbra, and

312 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

d’ Andrade (MUC, ML); 1 9, C Portugal, Estoril, 20.iv.1952, N. F. d'Andrade (MUC). — North Africa

(W to E). Morocco: 1 & 1 9, Tangier, J. J. Walker, E. Saunders coll., & 11.2.02, Wism. Morocco

(BM); 1 9, Tanger, J. Vachal 1911 (MP); 1 9, N Morocco, Chaouen, 22.v.1971, F. Hüschinger (CE); 1

3, Maroc, Dj. Amsittene, S de Mogador, 13.11.1961, H. Lindberg (MH); 1 9, Maroc, 622/19 (MG); 2

Imouzzer di Kandar, 1350 m, 25—26.v.1968, M. A. Lieftinck (ML); 1 9, same area, Azrou, 1250—1400

m, 20—24.v.1966, M. A. Lieftinck (ML). — Algeria: 1 g (pl. 6 figs. 3334), with vertically pinned blue

disk, labelled ‘“Grandis 3” (Lepeletier’s writing), obviously one of Lepeletier’s & of M. grandis Lep.!

(MP); 5 3, Algérie, coll. J. de Gaulle 1919 (MP); 1 9, Arzew (?) avril 1888, coll. J. Vachal (MP); 1 & 2

Q, Prov. d’Alger, Chellala, Jardin Romanetti, Vauloger 138—97 (MP); 1 9, Bône, 4.3.96, E. Saunders

Rothschild & Hart., and Hamman Meskoutine, v.1911, E.G. B. Meade-Waldo (BM, ML); 1 9, Algier,

M. luctuosa Scop., det.? coll. de Saussure 1910(MG); 2 3 3 9, Alger, Oran, Schmiedeknecht 1910, coll.

A. Weis (SMF); 1 g (diss.), Alger, Bab el Oued (same loc. as lectotype acutivalvis Alfken!), 25.11.1910,

three labels ‘‘/uctuosa Fr., nicht grandis”, ‘M. crassicornis Fr.”, and ‘‘Behaarung d. 5. Stern. lockerer u.

kürzer”, Sig. Alfken (MNB); I 4, Alger, 19.1.1910, Dr. J. Bequaert, M. grandis Lep., det. Alfken 1933,

with addit. pencil-written label “Ypsilon Type”, Sig. Alfken (MNB); 2 9, Alger, 25.iii & 7.iv.1910,

Husseyn-Dey, Dr. J. Béquaert, M. grandis Lep., det. Alfken 1933 (MNB); 2 9, Alger, S.iii.1910, and

5.II & 10.11.1882, arranged in drawer sub M. luctuosa meridionalis (2 3), Tunisia, Cartagina, I—II. 1882,

9), all G. & L. Doria, but none bearing identification labels: probably syntypic for both varieties

(CMS); 1 9, Tunis, Gafsa, 21.4.03, Birö (MBUD); 1 £ (diss, figs. 292—293) 14 9, Tunis, 7.2.03 (3)

M. crassicornis Fr.=italica Rad., det. Alfken, Sig. Alfken (MNB); 1 & (diss.), Tunis, Kairouan, coll.

Santschi, M. crassicornis, det. Friese (MNB); 1 9, Tunis, coll. Schmiedeknecht (MNB); 1 ¢ (together

with & M. leucorhyncha!), Tunis, 3—5.1912, Zobrys, Sig. Alfken (MNB); 1 g, blue disk, Tunis/coll. J.

coll. A. Weis, one with /uctuosa, det. Schmiedeknecht (SMF); 1 9, Tunis, Dr. Vohsen 1903, coll. A.

Weis, /uctuosa, det.? (SMF); 1 9, Tunésie, Dermech/Carthage, 20 m, 6.iv.1965, R. T. Simon Thomas

Tunis, Coll. Graeffe, M. aegyptiaca Rad., det.? (NMW, ML); 1 9, Tunis, März, Coll. Graeffe,

M. armata var. grandis Lep., det.” (NMW); 4 9, Tunis, Coll. Graeffe, III (Kohl’s writing), M. luctuosa

var. albovaria, det.? (NMW); 2 9, Nord Tunis, Spatz, M. luctuosa, det. Friese, and Tunis, iv.1927, R.

Meyer, M. luctuosa, det. Alfken 1935 (MNB); 1 ¢, Médinine, coll. J. Vachal 1911 (MP). — Libya:

Tripolitania, 5 & (1 diss.) 17 9, Tripoli, 4.ii—4.iii.1954 & 30.1.1955 (3 3 8 9), Gargarese, 15.iii.1952 (1

9), Sidi Mesri, 29.1.1954 (2 ¢ diss.), Garian, ca. 2500 ft., 22.ii.1954 (1 9), Sabrata O’ ASL, 27.11.1954 (2

9), Caradin Hills, 4.1.1955 (1 9), Leptis Magna, 5.i & 11.iii.1955 (4 9), all K. M. Guichard (BM, ML); 5

Egypt: 1 9 (diss.), “Aegypten, Kingi, 17.2.1912” (Friese’s hand) (MNB); 2 9, Aegypten, A. Andres,

one with Melecta aff. assimilis Rad., det. Alfken (SMF). — Israel-Jordan: 7 g, O. Jordan, Wald bei

Jerash, 600 m, 15.iii.1959, J. Klapperich (MBUD, ML); 1 g, Jerusalem, J. Sahlblerg] (MH); I &,

Palestine, Jerusalem, 29.ii.1940, leg. Bytinski-Salz, M. luctuosa var., det. Mavromoustakis (CBS); 4 & (1

diss., fig. 294), Palestine, Dahlia, 6.ii.1948, leg. Bytinski-Salz (CBS, ML); 1 g, Palestine, Jerusalem,

“10.8.1946” (sic), leg. Bytinski-Salz (CBS); 1 & 1 9, Palestine, Nazareth, 1800 ft., 16.iii.1920, P. J.

LIEFTINCK: Palaearctic Melecta 313

Barraud (BM); 4 3, Jerusalem, Mt. Scopus-N., Bellevalia, 13.11.1947, one at Rosmarinus (CBS, ML); 2

Q, Israel, Montfort, 7.iv.1950 & 20.iii.1964 (CBS); 1 9, Israel, Bridge Beerotaim, 26.ii.1968, 11—12

Cyprus: | g (diss.), Lagoudera, 15.iii.1971, 3000 ft, K. M. Guichard (BM); 1 & (diss.), Famagusta,

7.1.1932, E. E. Green (BM). — Rodos (Greece): 2 3 (both diss.), Rhodus, 18008 (MNB) & Rhodus

Loew, 18008 (MBUD); 1 & (diss.), Rhodos, Monte Smith, 20.iii.1978, K. J. Hedquist (NRS). —

Greece: 1 & (diss.), Macedonia, Langhadas-Saloniki [Thessaloniki], 3—8.iv.1956, Fr. Borchmann

Evvoia 1.] iii.1926, Holtz (MNB); 1 g, Ionian Is., Sir E. Saunders (BM). — Yugoslavia: 1 9 (diss.),

Istria, Rovigno, v-viii.1911, leg. Anisits S. G. (NMB).

The position of M. italica among its congeners has been ignored ever since

Friese (1895 et seq.) relegated it to the synonymy of Melecta funeraria F. Smith (see

Lieftinck, 1969, sub Eupavlovskia). The suppression of the name was accepted

offhandedly by contemporary and subsequent authors publishing on Palaearctic

Melecta. Even in Schmiedeknecht’s ‘“Hymenopteren Nord- und Mitteleuropas”

(1930) italica still figures as a synonym of M. (rect. Eupavlovskia) funeraria. This

neglect was initiated by Radoszkowski himself, who in the original description

(1876) not only mistook the type male of M. italica for a female but, only one

paragraph further down, added the definition of an indubitable example of

Eupavlovskia, misidentifying this as the male of that “female” italica. Without

giving any explanation, the author corrected the error many years later by

describing and figuring the male genitalia of true italica (1893 : 182, figs. 36 a-c. i),

— only to get mixed up again on a next page of the same publication by supplying

sketches taken from a male which he thought was Melecta grandis Lep.! In fact, the

last mentioned drawings pertain to a specimen of Eupavlovskia, as is evident from

his illustrations (figs. 40 a-c, i & k). The result of all this has been that M. italica

remained hidden and unrecorded in the literature for over half a century. It was

the late G. Enderlein who finally implied that the original specimen preserved in

the Berlin Museum, whose sexual organs had been first dissected out and

illustrated by Radoszkowski in 1893 (loc. cit., fig. 36), should be the original italica.

This was never published, although Alfken, as time went on, may have been aware

of the nomenclatural confusion and its bearing upon a species so well represented

in many private and European museum collections. Nevertheless, as will be seen

from the above census of studied material, a disorderly situation persisted, since

all italica (save its type), brought together for the purpose of the present work,

remained misidentified or were left unnamed. Enderlein’s type designation was

acknowledged by Lieftinck (1969) and can now be substantiated by descriptions

and illustrations.

Synonymy. — M. meridionalis (Tunisia). — As will appear from Gribodo’s

comments on the diagnosis of ““luctuosa Var. I meridionalis”, it is evident that this is

only a profusely white-haired form of italica Radoszk. The author aptly described

a number of features shared alike by meridionalis and leucorhyncha, but failed to

explain why two new varietal names were proposed for the evidently composite

taxon ‘‘/uctuosa’’ sensu Gribodo. The author, overlooking Radoszkowski’s

diagnosis of the much earlier defined italica, mentions a variety of localities for

314 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

meridionalis (see above) but, as Dott. D. Guiglia informed me in a letter (June 14,

1958), specimens from Piemonte, specified in the first place, are altogether

wanting, only North African individuals being available. Thereupon, while

attempting to interpret the present variety, the chief difficulty met with concerns

the selection of an appropriate lectotype. As with Gribodo’s ‘armata Var.

mediterranea”, all individuals named meridionalis (or arranged under that name)

bear near-identical labels “Tunisi leg. Doria...” etc., there being no single

specimen of European origin amongst them. As none of these could be traced, we

may safely assume that all French, Italian or German bees stated to belong here

were confounded with other taxa, or shared out among correspondents outside

Italy. Of the whole series, comprising 3 males and 3 females, I have selected a

lectotype male and a topotypical syntype female from Tunisia, i.e. the only ones

bearing the author’s own written identification labels, albeit that one of the

females figured as leucorhyncha. All are still in fairly good condition.

M. acutivalvis (Algeria). — While engaged in a study of Bequaert’s Algerian

collection of bees, Alfken (1914) got entangled by proposing another new varietal

name of ‘‘/uctuosa’’, viz. acutivalvis n.var. This was based on examples apparently

differing from others called by him ‘“Stammform” of luctuosa. As far as I could

ascertain, no single /uctuosa was amongst the many Algerian specimens

enumerated under that name, some pertaining to albifrons albovaria, several others

to italica, while one male proved to be /eucorhyncha. All names written on Alfken’s

pin-labels will be found specified under “Further material” listed for the

respective taxa just mentioned. Only one of the two males from Bab el Oued

carries the author’s own identification label ““v. acutivalvis, for which reason this

was selected lectotype. It differs in no way from light-haired (i.e. “plus variants’)

of italica occurring in Algeria and elsewhere in North Africa.

M. crassicornis (Corsica). — The male from Corsica was selected and labelled

Type by Friese himself, and by that very fact should be considered holotype. It is a

dark specimen of moderate size with reduced abdominal spots, quite similar to

nominotypical examples of italica from the same island and opposite provinces of

Italy.

M. quadripunctata (Italy). — The brief description of this puzzling bee is based

on a single female recorded from Italy, but lacking a locality label. It was also

erroneously named “4 pustulata’” by Radoszkowski, or in someone else’s

handwriting. Enderlein considered this female to be one of several more types —

formerly in Radoszkowski’s collection — acquired by the Berlin museum. On

selecting and labelling types from this collection, he evidently recognized it as

Figs. 275—287. M. italica; 275, sternite 8 (3, lectotype “Italie’’); 276, lateral view of gonostylus (same

specimen); 277, left antenna, showing rhinaria on segments 3—13 (4, Borzoli, N Italy); 278, ventral

view of apex tergite 7, showing ridges (upper, ¢ Lazio, Italy; lower, ¢ Monticchio, Italy); 279, sternites

7 and apex of 8 (4, Basilicata, Italy); 280, apices of sternites 7 and 8 (4, Lazio, Italy); 281, sternites 7

and 8 (3, “Italia mer., Zeller”); 282, partial ventral and dorsal view of genital capsule (4, Lazio, Italy;

283, outer contour: dorsal view of pygidial plate (9, Bari, Italy); 284, sternites 7 and 8 (3, Grimaux, S

France); 285, sternites 7 and 8 (4, Llobregat, Spain); 286, apices of sternites 7 and part of 8 (£ Maison

Carrée, Algeria); 287, inner contour: dorsal view of pygidial plate (9, syntype meridionalis, Tunisi). All

figures, including 283 and 287, on different scales

LIEFTINCK: Palaearctic Melecta

315

316 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

such, but instead of correcting the mis-spelt name he copied the original and

labelled it “M.4 pustulata Rad. Type, Dr. Enderlein”. While assuming this

specimen to be indeed the mislabelled type of quadripunctata, I have accepted

Enderlein’s type designation, especially because it corresponds closely to the

(incomplete) description of that species. It is a somewhat worn specimen,

measuring only 11.5 mm, yet undoubtedly conspecific with italica. Features of

interest are the small size and predominance of white on the thoracic dorsum,

agreeing in these respects with a few other atypical females listed above as from

Venice, southern France, ““Syra” (?), and one labelled “Attica”. All differ from

M. leucorhyncha in the long pubescence, compact body form and short, broad

subcordate abdomen. The original description being worthless, the next brief

characterization may not be found out of place: — Antenna moderate, segments

4—12 slightly but distinctly longer than broad. Dorsum of head and thorax closely,

almost contiguously punctate, interspaces slightly shiny; punctation of abdominal

tergites finer, less dense on more lustrous ground. Scutellar spines suberect, longer

than broad at base but much shorter than surrounding hair. Body pubescence

throughout long and dense, predominantly black; thin fringe of white only on

inner (anterior) face of antennal scape, just behind that level, and more densely so

at occipital border. Thorax dorsally mixed black and white as far back as tegulae,

with pair of indistinct black dots upon mesonotum anteriorly; small white tuft

upon middle of sides; remaining parts black. Legs hairy; small white subbasal spots

at outer faces of mid and hind tibiae, vestiges of white also at fore tibia. A thin

raised fringe of silky white at base of tergite 1, exceptionally long and tufty at sides

without forming condensed spots, the latter on 2—4 relatively small, subcircular,

placed wide apart in line, forming dense decumbent patches raised above level of

surface, but on 2 almost twice as large as on 3—4. Pygidial plate in side view

almost straight, subtriangular, length/breadth ratio 30: 20, raised margins straight

in dorsal view, tip not suddenly narrowed but almost parallel-sided, the rounded

apex with short, feebly raised median ridge; disk flat, impunctate, finely

reticulated, colour chestnut.

The diagnostic characters of italica, given in the key, can be supplemented as

follows:

Male. — Labrum squarish, disk concave, anterior border straight or slightly

convex, often with tiny midapical spicule. Fan-like silky white patch covering

middle of clypeus depressed or slightly raised, the hair tips usually exceeding

anterior border; long hairs at sides erect, black. Antennal scape fringed

roundabout with long raised white hairs (partly obscured or interspersed with

black only in dark populations). Similar long raised hairs on either side behind

wings not continuous across scutellum: median area of the latter always more or

less broadly black. Body pubescence throughout long and dense; black and white

partaking in design extremely variable, white hairs replaced by black in certain

populations. Outer face of hind tibia closely punctate, clothed more or less densely

with appressed hairs and thick setae, scantily interspersed with strong, acuminate,

spike-like denticles but often also fringed with long, erect, marginal bristles. Hind

basitarsus relatively short and broad, gently outcurved, surface at least slightly

LIEFTINCK: Palaearctic Melecta 317

Figs. 288—294. M. italica; 288, ventral view of tergite 7, ridges omitted (4, Philippeville, Algeria); 289,

sternite 7 and 8 (same specimen); 290, partial ventral and dorsal view of genital capsule (same speci-

men); 291, sternites 7 and 8 (4, Gafsa, Tunisia); 292, the same (Z, Tunisia); 293, ventral view of tergite

7, showing ridges (4, same specimen); 294, sternites 7 and 8 (4, Dahlia, Israel)

concave externally, the upper border nearly straight, usually beset with long raised

bristles, lower border convex; elongate-oval ventral pit near base small or

unapparent. Wings subhyaline, or fore wing membrane lightly infuscated toward

apex. Tergal spots greatly varying in size, occasionally vestigial, though never

entirely wanting on tergite 2. Ventral ridges at apex of sternite 6 low, hairy. Shape

of sternal plates 7 and 8 unstable, see figs. 275—294 and 359, bristly setae on apical

lobes of 7 characteristically more or less arranged in two groups, one on each side

of the median line. Gonocoxal angle never incurved, the diverging borders of

enclosure following a straight or feebly undulated (shallowly concave) course

toward distal edge of gonocoxite, the dorsoventral process of gonostylus almost or

fully twice as long as its breadth at base, subtriangular, varying in shape, but never

finger-like (fig. 276, of lectotype, taken from dry sclerite).

Female. — Previously undescribed. The most outstanding distinctives of this sex

are here recorded for the first time in the key.

318 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Like many other species, M. italica varies much in size and stature, certain males

and females listed from the Italian provinces of Emilia and Campania, and those

collected in Corsica and Mallorca (Balearics), are among the bulkiest and largest

of all populations examined, measuring up to 17 mm in length. These also have the

strongest and broadest antennae.

Outside Italy and the Tyrrhenian, viz. all populations of italica s.str., which

occur in the west, south and east of that area, look very different, the array of

white pubescence being on the average much more extensive in both sexes,

especially the white tergal maculations occupying more of the surface and are

more regularly arranged. Since intergradations do occur, I am reluctant to split

italica into two subspecies. But if it comes to that, meridionalis could be re-instated

as the first available name for a more richly white-marked race occurring in the

west, but ranging from southern France, Iberia and northwest Africa eastwards to

as far as Egypt and Israel-Jordan.

A few males of very small size, listed as from Cyprus and Rodos, are rather

puzzling (figs. 358—359). Though agreeing with the ‘‘meridionalis’’-type of italica in

respect of hairiness, with much more material these may ultimately prove to be

specifically distinct. They were collected in the same islands, together with some

equally small-sized individuals (both sexes) which I am unable to distinguish from

leucorhyncha (see under that species).

Melecta assimilis Radoszkowski

(figs. 295—301, pl. 8 fig. 40)

Melecta assimilis Radoszkowski, 1876, Horae Ent. Soc. Ross. 12: 122—123, pl. 3 fig. 6 (col. picture “9”

Egypte, wrongly sexed = 4).

Type material. — Egypt [collected by Comte Alexandre Branicki’s assistant Dr.

Dziedzicki, teste Rodoszkowski]: 1 ¢ (diss., figs. 295—300, pl. 8 fig. 40), labelled

“Egypt C: Bra” (capital print on green) “Type” (orange-red), “assimilis”

(Radoszkowski’s writing). Holotype Melecta assimilis Radoszk. (MNB).

27.3.1935 (written), Anastase Alfieri Collection 1965 (print), 921 (USNM).

This species is correctly described as a male, but in the legend of the very poor-

coloured picture the specimen is erroneously given as a female. The brief

description of the abdominal marks is liable to be misinterpreted: “… sur le

troisième de chaque côté deux taches rondes qui se touchent et dont la deuxième

est plus petite.” In fact the spot is merely somewhat indented by black behind, the

outermost portion being the smallest, — obviously an aberrance.

Male (unique). — General appearance, pl. 8 fig. 40. Additional characters:

Labrum slightly longer than its width at base (100 : 85.2), little narrowed toward

apex, coarsely punctate, anterior half with fine impunctate median carina.

Antennal scape a trifle longer than 3 + 4, fringed on either side with long white

hairs; 3 little longer than its apical width, exceeding length of 4 in the same degree

(10: 8), fig. 295. Punctation on mesonotum dense, punctures large and rather deep

LIEFTINCK: Palaearctic Melecta 319

Figs. 295—301. M. assimilis; 295, frontal view of antennal segments 2—S (g, holotype, Egypt); 296, ex-

ternal view of right hind tibio-basitarsus (same specimen); 297, dorsal and ventral view of tergite 7 (sa-

me specimen): 298, sternites 7 and 8 (same specimen); 299, partial dorsal and ventral aspects of genital

capsule (same specimen); 300, external view of left gonostylus (same specimen); 301, left lateral and

partial dorsal view of pygidial plate (9, Wadi Rischrash, Egypt). Scale line (298) | mm

on shiny ground, especially so where more widely spaced, at level of tegulae.

Scutellum strongly convex on each side of middle, transverse basal sulcus deep,

whole surface closely, coarsely and contiguously punctate; spines raised, bluntly

pricker-shaped, directed obliquely caudad, shorter than surrounding pubescence.

Hind tibia longer than basitarsus (100 : 77), outer face reticulate-punctate, rather

shiny, clothed with short black hairs and scattered spike-like setae. Felty pad at

outer face of mid tibia isolated, sharply defined, elongate-oval, finely black

roundabout, with few dark hairs posteriorly. Inner rami of mid and hind tarsal

claws slender, only little shorter than outer. Graduli of abdominal sternites beset

320 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

with longish black bristles, but all hairs much shorter and less closely set than in

italica. Tergal plate 7 with narrow, smooth and shiny, median area extending

nearly whole length, this space slightly concave but sparsely punctate at base only,

deeply sulcate between the two apical tubercles; colour brown, the posterior

border in ventral view yellowish (figs. 297). Apex of sternite 6 broadly rounded,

median depression elongate-triangular between low ridges, which are invisible in

profile but more hairy than the impressed area. Sternites 7—8, figs. 298. Genital

capsule, fig. 299: gonostylus short and broad, gradually narrowed toward apex,

which is broadly rounded; dorsobasal process distinct, subtriangular, one-fourth to

one-fifth as long as stylus, apical bristles very long, the longest exceeding tip of

stylus (fig. 300). Total length 14.0 mm approx., fore wing 11.2 mm.

Female (unique). — Head and thoracic sclerites rather deeply punctate on shiny

ground, punctures only little smaller than interspaces, except middorsally on

mesonotum where punctures are more spaced, those at sides being much smaller

and contiguous. Scutellar spines short and strong, bluntly pointed and shiny, raised

almost straight up, but shorter than surrounding pubescence. Abdomen short,

broadest at end of 1, as in male, deep black; tergites moderately lustrous, tergal

punctation and pubescence also similar, but all black hairs shorter than in male;

white lateral spots small, I—2 and 4 subcircular, 3 slightly more transverse, all

placed far apart, very compact, composed of depressed hairs slightly raised above

surface level, the hair tips not extending beyond hind margin of tergites; spots on

tergite | somewhat tufty, the broad space separating them clothed basally with few

long and thin, raised darkish hairs. Pygidial plate broad at base, then converging

with slightly convex, subacute and raised lateral carinae, the smooth median ridge

occupying about apical one-third of whole length of plate (fig. 301). Total length

13.5 mm approx., fore wing 11.0 mm.

The male of this remarkable species is very similar in many respects to italica

and was first thought to be only an aberrant individual of that species. It can be

distinguished therefrom by the thin fringes of much shorter and fewer raised hairs

covering the hind femora at all sides, the latter being replaced in all fresh

specimens of italica by a dense garment of much longer black pubescence covering

this and other parts of all legs.

Always excepting the sexual differences, the female is the only specimen which

corresponds closely with the type. As to its stature and markings, the resemblance

to Egyptian examples of italica is undeniable; but, in addition to the main key

characters, it can be distinguished from the latter by the darker wing membrane,

shorter third submarginal cell, smaller tergal spots, less shiny abdomen, and

shorter vestiture of all body parts; the more slender and swollen apical portion of

the pygidial plate of the female being an additional feature of assimilis.

Melecta curvispina Lieftinck

(figs. 302—309)

Melecta curvispina Lieftinck, 1958, Comment. Biol. Helsingfors 18.5: 22 & 25 (key SQ), 28—29, fig.

9—10, 16—17, 22—25, 35 & 38(3 9 struct., Tenerife & Gran Canaria).

LIEFTINCK: Palaearctic Melecta 321

Additional material. — Canary Is., Tenerife: 1 9, Tenerife, Dr. Verneau (MP); 1 3 2 9, Bajamar,

4.xii.1955, J. M. Fernandez (CB); 4 9, Las Canadas, 14.iv.1957 (9), El Rosaria, 20.iv.1960 (9), and

Cumbre de Ergo, 13.v.1960 (2 2), all O. Lundblad (NRS, ML); 1 9, Cumbre de Aguirre, 26.iv.1959,

Fernandez (ex coll. Verhoeff, ML); 9 3, Cruz da Afur, 5.iv.1904, E. Saunders coll. (BM); series ¢ 9

Tenerife, 1000—2100 m, v—vi.1964, K. M. Guichard (BM); series of 72 ¢ 9, Orotava, Agua Garcia,

Canadas, Mercedes, Villaflor and P. del Hidalgo, 25.iv—2.v.1971, H. Teunissen (CT, ML); 2 & 1 9,

Las Canadas, 6.iv.1972 (2 g) and Buenavista, 19.iv.1972 (9), K. W. R. Zwart (ML); 1 9, Arona,

Klimesch (CE); 4 g, Puerto de la Cruz, 23—27.iii.1976, S. Erlandsson, together with Anthophora a.

alluaudi J. P. (NRS, ML); 3 &, Vilaflor, 1300 m, 4.iv.1977, H. Wolf (CHW); 1 & 1 9, Bco. Infierno,

25.11.1972 (g), A. Machado, and Los Gigantos, 11.iii.1972 (9), B. Gustafsson (NRS). — La Palma:

series 4 9, v—vi.1964, K. M. Guichard (BM); 1 9, La Palma, Cumbre Nueva, W slope, “cleared pine

M. luctuosa, det. Friese (MNB); 1 ¢ (diss.), Kanar. Ins., La Gomera, Valley Gran Rey, 2—16.iv.1971, J.

Klimesch (CE). — Gran Canaria: 2 9, Grande Canarie, env. de Sta. Brigida, San José Montafiela,

Bartolomé de Tirajana, 16.iii.1968 (9), W. Backhuys (ML).

A very distinct species, well characterized by the curved processes arising from

the scutellar lobes. The type is from Tenerife.

302

303

305

Figs. 302—309. M. curvispina (Canary Is.); 302, frontal view of left antennal segments 3—4 (¢ 9, Tene-

rife); 303, left scutellar spines (same specimens): 304, apices of tergites 7 and 8 (3, same specimen);

305, apical border of sternite 6 (3, same specimen); 306, sternites 7 and 8 (Z, same specimen); 307, api-

ces of same; 308, oblique lateral view of gonostylus (3, same specimen); 309, dorsal view of pygidial

plate (9, Tenerife). After Lieftinck, 1958

322 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Male. — The characters summarized in my previous key (1958) are incomplete,

no mention having been made of one very important character, viz. the presence

of rhinaria on the antennal segments, which are well developed in the males of all

species known from the Canaries. In M. luctuosa these scent organs are absent, but

this species has not yet been found in the islands. With the amended note on the

antennal structure, the original descriptions and copied illustrations (figs.

302—309), it will be easy to separate curvispina froms its congeners. Apart from the

usual infraspecific variation in structure, no differences could be detected

between males from Tenerife and three other main islands of the group.

Female. — Two fairly distinct insular varieties (subspecies?) are recognizable:

(1) Tenerife: Clypeus, a broadly hairless anterior border excepted, invariably

clothed with raised black hairs; long dense pubescence fringing antennal scape

white anteriorly, black posteriorly; abdominal spots conspicuous, largest and

completely isolated on tergite 1. — (2) Gran Canaria: A brilliant fan-like patch of

decumbent silvery hairs upon middle of clypeus; hairs all around antennal scape

black; tergal spots similar to (1), but all comparatively smaller. These (unisexual!)

differences in hair colour and size of tergal spots between populations from

Tenerife (terr. typ.) and Gran Canaria, are worthy of note. By the absence of a

female from Gran Canaria, these features could not be taken into account in my

earlier description, but certainly deserve full attention. La Palma and Gomera are

new insular records for this species. Females from these islands are no more

available for comparison with the others.

Melecta canariensis Lieftinck

(figs. 310—312)

Melecta canariensis Lieftinck, 1958, Comment. Biol. Helsingfors 18.5: 26—28 (key 9), 30, fig. 11, 18 &

40 (9 struct., “Canary Is.”, MP).

Additional material. — Canary Is.: 1 9 (diss.), labelled ‘‘69/85” (written on white disk), small purple

square, “Canary Islands” (written on white rectangle), and my own identification “M. canariensis =

?curvispina Lieft., det. 1958” (BM).

This second female, though not in too good a condition, compares well

structurally, in colour and size, with the holotype of canariensis, whose precise

habitat is also unknown! Apart from the specific difference in the scutellar

processes, which are distinctly shorter and not markedly curved, it agrees best

with variety (2) of curvispina, since it possesses a silvery spot upon middle of

clypeus, while all hairs fringing the antennal scape are black. Pubescent pattern

and size of tergal spots as in populations from Gran Canaria. It is evident,

therefore, that this specimen was not collected on the island of Tenerife. With the

discovery of yet another species, M. prophanta spec. nov., in the Canaries, it seems

best to consider all taxa presently known from these islands distinct species. The

nearest ally of canariensis would seem to be M. leucorhyncha Gribodo, discussed

hereafter.

Male. — Unknown, but likely to be discovered in one of the islands, which will

be necessary to establish its status and affinities with more precision.

LIEFTINCK: Palaearctic Melecta 323

Melecta caroli Lieftinck

(figs. 313—321)

Melecta caroli Lieftinck, 1958, Comment. Biol. Helsingfors 18.5: 23 & 28 (key ¢ 9), 29—30, fig. 12—13,

19—20, 26—29, 36 & 41 (£Q struct., Lanzarote).

Melecta nigra Brullé, 1839, in Webb & Berthelot, Hist. nat. Iles Canaries 2 Ent: 89 (Q Canaries). —

Lieftinck, 1958, Comment. Biol. Helsingfors 8 (5): 30 (homonym).

Additional material. — Canary Is., Lanzarote: 4 9, Islas Canarias, Lanzarote, Puerto del Carmen,

W.N. Ellis (MA, ML). — Fuerteventura: 2 9, Betancuria, 2.v.1964 & above Ampuyenta, 29.iv.1964,

K. M. Guichard (BM); 1 9, Fuerteventura, Canar. Ins., Polatzek, ii.1894 N.1 (NMW); 2 ¢ 2 9 (diss.),

Fuerteventura, Betancuria (9), Vega Rio Palma (2 2) and La Oliva (9), 12—13.ii.1977, leg. M. Baez

(MLLT, ML); 2 2, Fuerteventura, La Oliva, 11.v.1974, J. M. Fernandez, and 29.v.1974, A. Machado

(NRS); 1 9, Canar. Ins., Fuerteventura, 13—26.iii.1926, Hering (MNB).

The rich supply of specimens presently available for study enabled me to

supplement the original description of this interesting little species (see also the

keys and figs. 313—321).

Both sexes. — Incomplete median carina of labrum slightly compressed and

upturned apically, occasionally extending upward toward base for about % whole

length of labrum. Antenna invariably 6-segmented.

Female. — Body totally black or almost so: occasionally a tiny white spot on

outer faces of mid and hind tibiae at some distance from base, and (still more

rarely) tufts of white in front of tegulae, behind wing bases and at outer sides of

tergite 1. No condensed black hair spots at sides of tergites 2—4.

Labrum short and squarish, broadest at base, anterior border slightly projecting,

the mid-apical crest ending in a pinched tubercle. Antenna slender, 3 much

shorter than scape, about one and one-third as long as next segments, but all

distinctly longer than broad; 2—5 subequal, 6 from one-half to two-thirds as long

as preceding ones. Head and thoracic sclerites closely punctate, punctures on

ocellocular area and disk of mesonotum deep, rather irregular, sometimes smaller

in part than smooth and shiny interspaces. Median mesonotal line very fine, not

impressed, ending at a level halfway length of tegulae, the latter superficially

punctate only near base. Scutellar tubercles suberect, bluntly pricker-shaped,

slightly divergent, shorter than surrounding pubescence. Inner (antecarinal) faces

of mid and hind femora impunctate for a !ong distance beyond base. Hind tibiae

densely hairy, outer faces dull, with numerous spicules evenly distributed. Inner

rami of mid and hind tarsal claws less than half as long as outer. Wings strongly

infuscated with slight bronze reflex, darkest along main veins and in cell centres;

third submarginal cell a little higher than long, outer angle not strongly convex;

nervellus well proximal to fork. Abdominal tergites finely, rather superficially

punctate, most closely on posterior segments, all punctures setiferous on smooth,

somewhat shiny ground. Pubescence moderately long and dense, as are the raised

subapical bristles on abdominal segments and legs, those fringing posterior faces

of mid and hind basitarsi distinctly longer than diameter of same. Pygidial plate

gradually downcurved, broad at base, length-breadth ratio about 100 : 70; sides

almost rectilinear, strongly converging as far as about % length from base, then

324 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Figs. 310-312. M. canariensis (Canary Is.); 310, frontal view of left antennal segments 3—4 (9 holotype);

311, left scutellar tubercle (same specimen); 312, dorsal view of pygidial plate (same specimen). — Figs.

313—321. M. caroli (Canary Is.); 313, frontal view of left antennal segments 3—4 (Lanzarote); 314, left

scutellar spines (same specimens); 315—316, contour of apex & tergite 7 (315) and apical border of

sternite 6 (316); 317, sternites 7 and 8 (4, same specimen); 318, apices of the same, more enlarged; 319,

oblique exterolateral view of right gonostylus (3, same specimen); 320, dorsal view of pygidial plate

(9, Lanzarote); 321, dorsal view of tergite 7 (4, Fuerteventura). All except ig. 321, after Lieftinck,

1958

pinched, the long slender apical portion almost parallel-sided (fig. 321).

Hab.: Apparently restricted to Lanzarote and Fuerteventura.

M. caroli is one of the five Melecta species presently known from these Atlantic

islands, and the only one showing pronounced melanism. It is quite distinct from

the other small-sized members, which are probably closely interrelated. At the

same time, however, caroli comes very near three other melanistic forms separated

from it geographically by a wide gap, all occurring in the mid-Mediterranean area.

These taxa are: gracilipes, from Menorca I. (Balearic Is.), and the much

obscurated geographical subspecies of two very distinct and widely distributed

species, viz. albifrons nigra and leucorhyncha taormina, occurring in most parts of

Italy as well as in the islands of Corsica, Sardinia, Sicily, and Malta. They share the

same tendency towards darkening of the vestiture, and accordingly look

remarkably similar, especially the two extremes amongst them, namely females of

taormina (Sicily) and caroli (Canaries), which are distinguishable only on close

scrutiny. The phenomenon is difficult to explain, but I venture to suppose that

climatic as well as edaphic influences have played an important role in the

development of subspecies showing a similar colour design.

LIEFTINCK: Palaearctic Melecta 325

Melecta leucorhyncha leucorhyncha Gribodo

(figs. 13—14, 322—332, 344357, map 3, p. 307)

Melecta luctuosa Scop. Var. II Leucorhyncha Gribodo, 1893, Bull. Soc. Ent. Ital. 25: 410—412 (pars:

“Piemonte: rarissima, e dall’Algeria: Boghari’’).

Melecta luctuosa Scop. var. leucorhynea (sic): Gribodo, 1924, Boll. Mus. Zool. Anat. comp. Univ. Torino

im April 1883, 9 auch von Ibiza am 20.April 1883”). Syn. nov.

Type and syntypic material. — Algeria: 1 9 (diss., fig. 325), “Algeria” (written,

white label), ‘‘Melecta luctuosa Scop. leucorhyncha Grib. 2g D. Gribodo”

(Gribodo’s handwriting, on light blue), lectotype M. leucorhyncha by present

selection (MCG). — Tunisia: 4 9 (diss.), with written black-rimmed labels

“Tunisi Belv[edere], 12.111.1882, G. e L. Doria” (1 9), “Tunisi dint. IV.1882, G. e

i Doria’ (2 9), and “Tunisia Cartagine, 1.111882; G. e L. Doria” (19);

Figs. 322—329. M. leucorhyncha leucorhyncha (N Africa); 322, sternites 7 and 8 (3, Chellala, Algeria);

323, sternites 7 and 8 (4, Colomb Bechar, Algeria); 324, dorsal view of pygidial plate (9, Oran, Alge-

ria); 325, the same of lectotype (9, Algeria); 326, the same (second 9 from Oran, Algeria); 327, ventral

view of tergite 7 (4, Bu-el-Gherab, Tripolitania); 328, sternites 7 and 8 (same specimen); 329, partial

ventral and dorsal view of genital capsule (same specimen)

326 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

syntypes, all under drawer label M. leucorhyncha Gribodo (MCG, 1 9 ML). —

Spain, Balearic Is: 1 & (diss., figs. 13, 330), “Mallorca, Friese” (print), “1891,

Type” (print on red), “Melecta” (print) “luctuosa v. ebusana Fr. Balear.” (Friese’s

writing), lectotype M. ebusana Friese, by present designation (MNB).

Further material. — Spain, Balearic Is.: 1 & (diss., figs. 331—332) 1 9, Mallorca, 20.iii.1959, O.

Mallorca, C’as Catala, 4.iv.1958, A. Compte Sart (ML). — N Africa (W to E): Morocco, 1 9 (diss.),

Middle Atlas, Azrou, 3000 ft., 6.iv.1935, D. Aubertin (BM); 2 9, Maroc, Moyen Atlas, Midelt, 1500 m,

30.v.1968, M. A. Lieftinck (ML); 2 9, Moyen Atlas, Ifrane, 1650 m, 26.v.1966, M. A. Lieftinck (ML); 4

Rozen (AMNH); 1 9, Marocco, Fritsch-Rein (?), von Heyden (SMF); 1 9, Marrakesh, 25.ii.1935, R.

& C. Koch (AMNH). — Algeria: 16 ¢ (2 diss, fig. 322) 2 9, Prov. d’Alger, Chellala, Jardin

Romanetti, Vauloger 138—97 (MP, ML); 1 & (diss.), Alger, Baraki, 10.iii.1910, Dr. J. Bequaert

(MNB); 4 G (2 diss., fig. 323), Algeria, Colomb Bechar, 4.iii.1944, K. M. Guichard, very small ex!

(BM); 1 9 (diss.), Algeria, Maison Carré, 27.iv.1943, K. M. Guichard, same locality as SQ M. italica

1 3, Algérie, coll. J. de Gaulle, M. luctuosa, det.?; Algérie, Tebassa, 2.v.1898, J. de Gaulle (1 9),

Biskra, coll. J. Perez 1915 (3 9) (all MP); 1 9, Algérie, Sétif, coll. de Saussure, M. luctuosa punctata

Marzo 1952 & Djelfa, 10 Aprile 1952 (IEB); 2 9, N Algeria, Biskra, 24.iii.1894, E. Saunders coll., and

Hamman Rhica, v.1911, Rothschild & Hartert (BM); 3 9, Algir, 593.108, 1881, M. /uctuosa, det. Friese

1893 (MBUD, ML). — Tunisia: 2 g, Gafsa, P. Chrétien 1910, no. 272 (MP); 2 9, Tunis, coll. J. Pérez

Santschi, M. armata var. mas (typewritten, NMB); | g (together with M. italica!), Tunis, Kairouan,

Gaulle (MP); 1 3 1 9, Médinine, furcata, det.? coll. J. Perez 1915 (4) and Medinine, M. luctuosa det.?

coll. J. Vachal 1911, no. 415(9); 1 g, Tunis, 3-5.12, Zobrys leg. (MNB); 2 9, Tunisie, Gafsa, A. Weis

M. luctuosa Scop. var. albovaria det.” (NMW); 1 3, Oase Gafsa, 28.11.1977, M. luctuosa, det.? (CMS). —

1953 (MA); 1 9, Cirenaica, Zavia, Mechili, Festa (MT); 1 9, Cirenaica, Tecnis, Festa, M. luctuosa

Scop. var. leucorhynca (sic) Grib. (Gribodo’s writing, (MT). — Egypt: 2 9 (diss.), Egypte, Naville

(MG); 1 9, Egypt, Hammam, 10.iii.1930, Min. Agric., coll. Andres-Priesner, Slg. Alfken (MNB); 1 9

(diss.), S. Sinai, ca. 1600 m, S. Atherina Area, 24—27.iv.1975, K. M. Guichard (BM). — Israel-Jordan:

Judaea Mts., Jerusalem, 3.iii.1940, and A 2099/Lower Jordan Valley, Jericho, 23.ii.1941, Y. Palmoni

(AID); 1 & (diss.), Israel, Arad, 10.ii.1966, and 2 g (diss.), Jerusalem, 29.ii.1940 & 2.iii.1940, M. luctuosa

doubtful status, are in (CBS). — Turkey (Asia minor): 4 & 1 9, Türkei, E. Uludere/Hakkari, 5.v.1977

& pass E Uludere Hakkri, 6.vi.1977, together with M. festiva m., K. Warncke (CKW, ML); 139 9 (all

M. luctuosa, det. Friese 1896 (MNB); 3 9 As. Turkei, Sille b. Konya, 17.vi.1968, J. Schmidt (CL) and

LIEFTINCK: Palaearctic Melecta 327

9—17.vi.1975, J. Heinrich (CJH); 8 9, As. Türkei, Gürün, 12—15.vi.1976, J. Heinrich (3 9) (CJH) and

same loc., 3—9.vi.1907, J. Gusenleitner (5 9) (CG); 3 9, As. minor, Konya, 1030 m, 11.vi.1966, K.

Kusdas (CJH), and same loc., 25.v.1965, M. Schwarz & 10—12.vi.1966, J. Schmidt (CMS); 2 9, As.

Turkei, Tarsus, 30.v.1965, M. Schwarz, and Perge nr. Antalya, 3.vi.1966, J. Schmidt (CMS); 1 9,

Akasaray, 8.vi.1964, J. Gusenleitner (CG); 1 9, Turkey, Havsa, 1500 ft., Samsun-Amasya Rd.,

19.v.1959, K. M. Guichard (BM); several 9 of rather doubtful status, collected in various localities all

over Turkey from April to June, are in (CW, CL, CJH, CMS, ML and MP). — Rodos I. (Greece): 1 ¢

(diss.), Ataviros, 18.v.1970 (1 3), and 24.iv.1976 (4 3, fig. 347), Profitis Ilias, 20—28.iv.1976 (7 &, 4

Agios Isodoros (4 3, figs. 345—346), M. Ataviro (2 g, figs. 348—349), Koskino (1 9, fig. 350), and

W. N. Ellis (MA, ML); 1 9 (aberrant), Mte Profeta Elia, 12.v.1935, O. Wettstein, M. luctuosa Scop.,

Ay. Neophytos, 27.iii.1971 (figs. 354—356), Nicosia, 500 ft., 5.iii.1971 (figs. 352—353), Antophonitis,

12.iii.1971 (fig 357), Kantara, 2000 ft, 11.iv.1971, and Vouni, 25.iii.1971, all K. M. Guichard (BM, ML);

1 g (diss.) 3 9, Cyprus, Yerasa and Paramytha, 6.iv.1978, and Yermasoyia, 3.iv.1978, H. Teunissen

Limassol, Yermasoyia, 17.iii.1979, L. A. Janzon & 1 9, Governor’s Beach, 16.iii.1979, same coll. (NRS,

ML); — Greece: 1 & (diss.), Peloponnesus, Chelmos, 1900 m, 2.vi.1962, Max. Schwarz (CMS); 1 3

(diss.), Parnass [= Mt. Parness 1200 m, 40 NW of Athens] (MCG); 1 9, Greece, Kifissis, 250 m,

iv—v.1977, K. M. Guichard (BM. — S USSR: Ukraine: 3 £ (all diss.), Chersonskaja dist., Novaja

Tjaginka, 19, 26 & 27.v.1954, one at flower of Caragana arborescens, two at Robinia pseudacacia, one

with M. luctuosa Scop., leg. & det. A. Z. Osychniuk (CO, ML).

Cherson, 14.v.1954 (1 ex.) and 19.v.1954 (4 ex.), A. Z. Osychniuk (CO, ML). The four females dated

19.v.1954, were collected the same day as a male M. luctuosa Scop., taken at nearby Novaja Tjaginka,

viz. the locality of M. leucorhyncha; probably both species (which are inseparable) are included in the

present series.

The revival of the name /eucorhyncha for a relatively small-sized Melecta, widely

and commonly distributed all over the Mediterranean region, is based upon two

considerations, namely (1) the conviction that it is specifically distinct from the

much earlier described italica Radoszkowski, and (2) that there is only a single —

though inappropriate — name available to denote it. As with his “‘/uctuosa Var. I.

meridionalis’’, Gribodo left his readers to decide which species should be assigned

to his “Var. II /eucorhyncha”, the brief Latin diagnosis giving no clue to that effect.

The name was given to bees from Piemonte (stated to be very rare at Torino) and

Algeria. There were, however, no examples from Piemonte to be found in his

collection, the only individuals under that varietal name being the five specimens

still before me. One is a female from Algeria while the remainder — like Gribodo’s

series of var. meridionalis — are labelled “Tunisi”, leg. Doria. A careful

investigation revealed meridionalis to be conspecific with italica Radoszkowski, see

under that species. Consequently, the choice of a type locality as well as the

selection of a lectotype, were also left to a future reviser. Since “Algeria

(Boghari)” is given as one of the localities for leucorhyncha, I have designated the

Algerian female lectotype, i.e. the first described sex and only specimen bearing a

328 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

label in Gribodo’s own handwriting (MCG). For that matter, we have seen that the

author himself got confused by arranging both sexes of his Tunesian series under

meridionalis, whereas four females passed for leucorhyncha,

Figs. 330—332. M. leucorhyncha leucorhyncha; 330, external view of right tibio-basitarsus (& lectotype

ebusana, Mallorca, Baleares); 331, dorsal view of tergite 7 (3, Mallorca); 332, sternites 7 and 8 (same

specimen) — Figs. 333—340. M. leucorhyncha taormina; 333, dorsal view of tergite 7 (¢, Bologna, Italy);

334, sternites 7 and 8 (same specimen); 335, dorsal view of tergite 7, sculpture omitted (4, Sassuolo,

Italy); 336, left hind tarsal claw (9, Sassuolo, Italy); 337, dorsal view of pygidial plate (same specimen);

338, sternites 7 and 8 (3, Corsica); 339—340, sternites 7 and 8 (2 3, Sicily). — Figs. 341—343, M. graci-

lipes (3 holotype, Menorca, Baleares); 341, external view of tibio-basitarsus; 342, ventral view of tergi-

te 7; 343, sternites 7 and 8

LIEFTINCK: Palaearctic Melecta 329

Figs. 344—357. M. leucorhyncha leucorhyncha; 344, genital capsule, left lateral and partial dorsal view,

showing bi-angulated gonocoxal angle (4, Profitis Ilias, Rodos); 345, dorsal view of tergite 7(¢, Agios

Isodoros, Rodos); 346, apex of sternite 8 (same specimen); 347, external view of right hind basitarsus

(4, Attaviros, Rodos); 348, dorsal view of tergite 7 (3, Attaviro, Rodos); 349, sternites 7 and 8 (same

specimen); 350, dorsal view of pygidial plate (9, Koskino, Rodos); 351, sternites 7 and 8 (4, Ayancik,

Anat. bor., Turkey); 352, sternites 7 and 8 (3, Nicosia, Cyprus); 353, right exterolateral view of gonos-

tylus (same specimen); 354, dorsal view of tergite 7 (4, Ay. Neophytos, Cyprus); 355, sternites 7 and 8

(same specimen); 356, right exterolateral view of gonostylus (same specimen); 357, sternites 7 and 8 (4,

Antiphonitis, Cyprus). — Figs. 358—359. M. italica; 358, dorsal view of tergite 7 (3, Lagoudera, Cy-

prus); 359, sternites 7 and 8 (same specimen)

330 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Pending an inquiry into reliable distinctive characters for both taxa, long series

of either sex had to be carefully compared, their habitations put on record one by

one, and the dates of capture specified as detailed as possible. As with italica, only

in that way it was possible to obtain a rough impression of the variation,

distribution and seasonal occurrence of both. We have seen that the two keep

company in many localities scattered all over their range. Added to that, structural

differences between them are slight, not easy to detect and evidently far from

constant. As to the vestiture, both exhibit considerable local differences, each

displaying its own individual departure from the familiar /uctuosa design, the

crucial point regarding this variability being that, in places where the two species

occur side by side, leucorhyncha may show a pattern analogous to that of italica, at

times tending towards obscuration (melanism), then again to a marked increase of

white pilosity. It must be admitted that in certain cases, where morphological

differences between the dark extremes of either are unapparent, no convincing

evidence could be provided that we are, indeed, dealing with a pair of sibling

species. In Sicily, for instance, marginal populations of each seem to meet,

become mixed and may even hybridize in certain localized spots. In any case some

females are so much alike as to become inseparable (see also sub italica).

M. 1. leucorhyncha Gribodo and luctuosa (Scop.)

It will be remembered that quite some controversial /uctuosa-like females

included in our locality lists had to be left unnamed, simply because reliable

recognition marks were not to be detected (see also under M. luctuosa). The

following observation may demonstrate the difficulty to distinguish between

females of leucorhyncha and others closely resembling them, especially /uctuosa. A

good example was found among Mr. A. Z. Osychniuk’s captures in the Ukraine

(USSR). He came across a mixed population of two near-alike Melecta occurring

at Novaja Tjaginka in the Chersonskaja district. On 19 May 1954, seven of these

papilionaceous shrub Caragana arborescens. Looking closely into their identity

later, it was easy enough to recognize two males of luctuosa, whereas the third

proved to be leucorhyncha. The four females collected in company with the latter

were so much alike in every respect that even at present I am not absolutely sure

which is which, or whether all belong to a single species. One week later (26 and 27

May), on visiting the same locality, two more males of leucorhyncha could be taken

at flowers of Robinia pseudacacia, but no more females were caught on that

occasion.

Melecta leucorhyncha taormina Strand, stat. nov.

(figs. 333—340 & pl. 8 fig. 41, map 3, p. 307)

Melecta luctuosa var. taormina Strand, 1919, Archiv f. Naturgeschr. 83 (11): 61—62 (9 Taormina-Lenti-

ni, Sicilien; incl. f. /entina nov. f. & f. tibiopunctata nov. f.).

Melecta nigra Lepeletier, 1841, Hist. nat. Ins. Hym. 2: 446 (9, ‘‘de Gênes en Italie, Musée du général

Dejean’’). Nom. preocc.: not M. nigra Spinola, 1806.

2 Melecta nigra: Dusmet y Alonso, 1905, Bol. Real Soc. esp. Hist. nat: 152, 153 (g Spain: Las Arenas,

Viscaya).

LIEFTINCK: Palaearctic Melecta 331

Type and syntypic material. — Italy: 1 9 (wing tips broken off), dark green

disk, “nigra” (old writing on vertical pin-label), lectotype (holotype?) Melecta nigra

Lepeletier (not Spinola), by present designation (MP). — Sicilia: 3 Q (diss.), all

with same printed label “Taormina-Lentini (Sizilien) v.1914 W. Trautman”; and

“Melecta luctuosa v. taormina m. Q” (Strand’s writing), “Strand det.” (print)

“Typus” (print on red), lectotype taormina by present selection; ‘‘Melecta luctuosa

v. taormina m. f. lentina m. Q” (Strand’s writing), “Type!” (id., written on red),

paralectotype taormina; and ““Melecta luctuosa Sc. v. taormina m. f. tibiopunctata”

(Strand’s writing), “Typus” (print on red), paralectotype taormina (all DEI).

Selinunt, 1.iv.1930, Segesta, 5.iv.1930, and Girgenti, 8.iv.1930, Troll/Hym. Inv. 31.iii Nos. 304, 305, 313,

M. luctuosa v. albovaria Er. form calabrina Rad. (Friese’s handwriting), det. Friese 1893 (2), M. luctuosa

M. italica (INER); 2 9, blue disk, Sicile, coll. J. Pérez 1915(MP, ML); | 9, Sicilia, Palermo, S. Martino

(MT). — Italia (continental, S to N): 1 9, Calabria, Sila Grande Silvana Mansio-M. d. Porcina,

ex. with “‘v. ruthenica”, | ex. with M. luctuosa Scop. v. ruthenica Rad., det. Alfken 1904 (all in Alfken’s

19, Abruzzo (Aquila), Gran Sasso d'Italia, östl. Arsita, 525 m, 23.vi.1962, leg. Reinig (SMF); 19,

Abruzzo, Mti Carbeolani, Castel vecchio, 1000 m, vi.1948, Bisleti leg. (CMC); 19 (diss.), Roma

(Lazio), M. Mario, 15.v.1946, Saracchi (CMC); | 9, Roma, Ostia Scavi, 10.v.1942, C.N.R., Ist. Naz.

Biol./Roma (INER); 2 9, Centocelle, 21.iv.1932 & Caffarella (Lazio), 29.iv.1935 (INER, ML); 1 9,

Capo Circeo (Lazio), Quarto caldo, 20.iv.1940, C.N.R. (INER); 2 9, Roma, 26.iv.1947 & Roma,

Torraccia, Montebello, 30.iv.1853, M. Comba leg. (CMC, ML); | 9, Macerata (Marche), vii (sic) 1897,

v. Bezzi, coll. P. Magretti (MCG); 1 9, Livorno (Toscana), Mann 1872, no. 70, M. luctuosa v. rutenica

figs. 333—334, Bologna, Ronzano, 9.v.1948, M. fasciculata Spin., det.? (IEB); 1 9, Emilia, Gaggio

M. luctuosa, det.? (3) and 28.vii (sic) 1890, sub M. ruthenica Rad., det.? (IEB); 1 g (diss.), Bologna

(Emilia-Romagna), Cafiualbo (?), di fiori, 21.iv.1889, coll. P. Magretti (MCG); 1 & 1 2 (diss, figs.

335—337) Emilia, Sassuola nr. Modena, 30.iv.1882, M. armata Pan., det.? (MT); 2 9, Modena (Emilia)

(MCG); 1 9, sine patria, 122 (written on square), det. Schmiedeknecht (S’s writing), coll. P. Magretti,

coll. Gribodo (print), “Je crois que c'est seulement une variété de M. luctuosa’’ (Schmiedeknecht’s

hand?) (MCG); 1 9, Genova (Liguria), v.1883, coll. Fea (MCG); | 9, Piemonte, under drawer label

drawer- and pin-labels in coll. Spinola (MT). — Corsica: 1 & (diss., fig. 338), Centr. Corsica, Col de

Bavella-Larone, 600—1240 m, 27.v.1965, ‘on dry hill slope together with Anthophora sicheli Rad.'), at

flowers of Astragalus sirinicus Ten. ssp. genargenteus (Moris) Brig.”, M. A. Lieftinck (ML); 1 9 (diss.),

') A species occurring also in Sicily.

332 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

NW Corsica, Fango-M. Estremo, 3—600 m, 6.vi.1965, M. A. Lieftinck (ML); 1 Q (diss.), Cors. Mann

luctuosa), Centr. Corsica, S. Pierre de Venaco, 4—800 m, 30.v.—2.vi.1964, M. A. Lieftinck (ML). —

1956 (IEB, ML).

Melecta gracilipes spec. nov.

(figs. 341—343)

Type material. — Balearic Is., Menorca I: 1 @ (diss., figs. 341—343), Mahon,

Cala de Figuera, sea level, 7.v.1958, “visiting flower of Echium spec. growing on

coastal promontory’’, M. A. Lieftinck. The specimen is the holotype (ML).

Male (unique, wings frayed). — Labrum square, coarsely densely punctate,

punctures of different sizes, those between large, subcircular, smooth and shiny

tubercles, small and contiguous, becoming larger, more impressed and partly

confluent toward apex; disk little concave, surface shiny; anterior border straight

with broadly rounded side-edges, not upturned but with minute, pinched

midapical tubercle. Mandibles normally unidentate, outer surface smooth,

sparsely minutely striato-punctate. Clypeus, frontal area and whole dorsal surface

of head strongly, almost contiguously punctate on shiny ground, punctures

circular, largest and most deeply impressed anterior to ocelli, leaving a small

impunctate areas just in front of median ocellus. Frontal carina impunctate,

sharply crested and highest anteriorly; temples and occipital area dull,

contiguously punctate. Antenna long and strong, scape closely finely punctate,

fringed at all sides with long raised hairs exceeding greatest diameter of the latter

and mostly white externally; 3 short, widest at apex (length-breadth ratio 100: 86),

only little longer than succeeding segments 4—12, which are squarish, 13 only a

trifle longer than broad; rhinaria deeply impressed, shaped similarly to those of

leucorhyncha from Mallorca (‘‘ebusana’’). Texture of thoracic sclerites as in the

latter: median mesonotal line extremely fine and partly obliterated (only apparent

at high magnification), the short parapsidal lines, however, quite distinct, raised,

impunctate and shiny; all sclerites strongly, closely punctate, the distance

separating circular punctures on mesonotum and scutellum hardly exceeding one

puncture width. Scutellar tubercles strong, conical, at least twice as long as broad

at base, directed obliquely upward and backward, distinctly diverging and slightly

downcurved. All lateral and posterior sclerites of thorax dull, extremely closely

punctate, including whole propodeum.

Legs noticeably more slender than in all leucorhyncha; hind femora, tibiae and

basitarsi distinctly more laterally compressed, all mid and hind tibiae relatively

longer (cf. figs. 330 and 341), e.g. hind tibia 1 : b = 100: 40, hind basitarsus 1: b =

100 : 31 (type “‘ebusana’’), as against 100: 32 and 100: 25 (type gracilipes). Mid

tibial pad pure white, much narrower than in the former (fig. 13), completely and

more broadly surrounded by black. Punctation of legs similar in the two, setiferous

punctures evenly distributed throughout, the hind femur lacking impunctate area

toward the end of its inner face, the median ridge less pronounced than in

Mallorcan leucorhyncha. Wings subhyaline, fore wing membrane only slightly

LIEFTINCK: Palaearctic Melecta 333

infuscated beyond cells; third submarginal comparatively long, exceeding its

height (115 : 100), the distal side moderately strongly angled, but more so than that

of the second cell. Shape and punctation of abdomen as in /eucorhyncha;

integument shiny, tergal punctures far apart, fine and of small size, those covering

the broad postgradular portion of tergites still finer, more widely spaced and

superficial. No indication of crowded punctures underlying compact hair spots on

any of the tergites, hairs replacing the white lateral spots on 2—4 not at all

depressed and hardly more condensed than those covering disk of tergites.

Pubescence covering all body parts moderately long, thinner and also more

disseminated than in either italica or leucorhyncha; predominantly black. Whitish

are: a broad, depressed, subquadrate patch of long semierect hairs upon middle of

clypeus; long raised hairs forming ill-defined collar on mesonotum anteriorly not

extending beyond level of front margin of tegulae; fluffy hairs at thoracic sides; a

few very long hairs behind bases of hind wing and at sides of tergite 1. Remaining

pubescence entirely black, the pile covering basal half of tergite 1 not at all dense

but all hairs of great length. Vestiture of legs much as in leucorhyncha, but hairs

longer and less closely set, those fringing hind femur posteriorly shorter than

greatest diameter of the latter; all black, with the exception of the tibial pad at

outer face of mid tibia.

Shape and texture of tergite 7 (fig. 342, ventral view, ridges and most of the hair

omitted!) hardly differing from my figure 331 of Balearic leucorhyncha: its dorsal

surface almost flat without any indication of lateral ridges; a broad, smooth and

rather shiny impunctate median area extends from base to apex, this strip broadest

at extreme base, a little constricted beyond halfway length, remainder of tergite

evenly closely punctate and hairy; tip narrow (0.2—0.3 mm), shallowly

emarginate, the lobes scarcely upturned, rounded; ventral ridges forming pair of

smooth, oblique tubercles, well visible in profile. Apex of sternite 6 slightly

projecting medially; a pair of low dark tubercles on either side of the usual, oval,

less sclerotized median area. Sternites 7—8 as in figs. 343. Genital capsule hardly

different in shape from leucorhyncha, the gonocoxal enclosure a little narrower,

feebly bi-angulate distally in dorsal aspect; gonostylus also similar, less than twice

as long as its width at base, tapering gradually to a broadly rounded apex;

dorsobasal process more slender and parallel-sided than in typical leucorhyncha

and the form of Mallorca, identical in shape to that seen in individuals from

Cyprus and Rodos (figs. 344 & 356).

Length 10.5 mm, fore wing 8.5 mm.

Female unknown.

This peculiar little species is probably most closely related to the polytypic

leucorhyncha, but can not be treated as a subspecies of the latter on account of its

strikingly more slenderly formed legs, the longer and more fluffy body

pubescence, and the reduced white markings. The above description is based

mainly on a comparison with ‘‘ebusana’’, from Mallorca, but the differences noted

apply equally well to nominotypical examples of leucorhyncha from North Africa,

which are quite similar (see under that species).

Remark. — The holotype of M. gracilipes was the only specimen of Melecta

334 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

observed during my stay in the island. It was flying in company with Anthophora

balearica (Friese) — originally described from Ibiza and Mallorca — on which it

might be parasitic.

APPENDIX

SPECIES INCERTAE SEDIS

Melecta nigripennis Lepeletier

Melecta nigripennis Lepeletier, 1841, Hist. Nat. Ins. Hym. 2: 445 (Q ‘‘Patrie inconnue. Musée du général

Dejean’’). Type lost (not in MP). Locality unknown.

The following is a translation of the French diagnosis.

Female. — Head wanting. Thorax black, its hairs white and black in places.

Abdomen black, all five segments carrying a triangular snow-white spot on either

side. Legs black, black-haired, except the fore coxae which are strongly tufted

with white. Fore wings wholly black, shading to purplish (‘‘violet’’); hind wings

much more transparent.

An enigmatic species, possibly identical with, and a synonym of, M. albifrons

nigra Spinola, 1806, though the description also points to M. leucorhyncha taormina

Strand, 1919, as characterized in the present work. Nigripennis being a nomen

oblitum, it should be discarded.

Male. — Unknown.

Melecta testaceipes Lepeletier

Melecta testaceipes Lepeleter, 1841, Hist. Nat. Ins. Hym. 2: 447 (9 “Patrie inconnue. Décrite de la col-

lection de M. Carcel’’). Type lost (not in MP). Locality unknown.

The translated French diagnosis runs as follows.

Female. — Entirely black, its pubescence black. Legs black, hairs black: all

tarsal segments including the claws, ferruginous (“d'un testacé ferrugineux’’).

Wings slightly enfumed.

Like the former, a species of unknown status, possibly also identical with one of

the species mentioned above.

Male. — Unknown.

Melecta octomaculata Radoszkowski

Melecta octomaculata Radoszkowski, 1876, Horae Soc. Ent. Ross. 12: 124 (2 — not 9! — Egypte).

This is another of Radoszkowski’s enigmatic species whose status could not be

explained. The holotype could nowhere be traced and is probably lost. Like the

type of italica Radoszkowski, described in the same paper, evidently a wrongly

sexed individual, possibly conspecific with M. transcaspica F. Morawitz, 1895? The

invalid name M. octomaculata Friese, 1925, based on a male from Turkey (Asia

minor), is synonymous with transcaspica (see under that species). Here follow

Radoszkowski’s original characterizations of octomaculata:

“Nigra; capite thoraceque pilis albis variegatis, dentibus scutellaribus subparvis,

LIEFTINCK: Palaearctic Melecta 335

abdomine opaco, segmento 1-0 dense albido-piloso, segmentis lateribus utrinque

macula nivea; tibiis omnibus albo-maculatis. Alis fumatis, hyalino maculatis. Long

15 mm.

Femelle. Noire. Chaperon garni de poils argentés. Le premier article des

antennes et le front faiblement garnis de poils gris. Mésothorax en dessus,

prothorax en dessus et dessous ainsi qu’une partie de la poitrine sous les ailes

garnis de poils hérissés, longs, gris blanchatres; les dents de l’écusson petites.

Abdomen opaque, garni de poils courts, noirs; premier segment garni de poils

longs blancs, vers chaque côté les poils plus épais et plus prononcés; chacun des

segments suivants portent sur leurs côtés une tache regulière allongee; la longueur

de ces taches va en diminuant vers l’anus. Les jambes des pieds antérieurs du côté

externe richement cilies de poils argentés; les jambes des pieds suivants couverts

du côté externe de poils très courts, couchés, argentes; les tarses de tous les pieds

couverts de poils pareils. Les ailes transparentes, leurs bouts et des taches dans les

cellules enfumés.”

This bee, along with the types of aegyptiaca and assimilis described in the same

publication, was collected by Comte Alexandre Branicki’s assistant, Dr.

Dziedzicki. We can only guess at its identity, so that it is best placed in the present

category qualified as a nomen oblitum.

Melecta eversmanni Radoszkowski

(fig. 112)

Melecta Eversmanni Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 180, 9 only!

(Q Orenbourg).

(different writing), “Type” (print on orange), “Melecta Eversmanni Radosz. Type,

rev. Dr. Enderlein”. Holotype, evidently recognized and designated as such by

Enderlein (MNB).

Like all other species described by Radoszkowski, the female of M. eversmanni

was unfortunately described in advance of the male, the genital organs of which

were figured (R's figs. 31 a—c, i). The male obviously came from a different

locality, Astrachan and Tachkend being mentioned after Orenbourg; moreover

the dissected male(s) could not be found any more in the Berlin museum and may

have become lost. So I feel justified to follow Enderlein’s unpublished view to

regard the present female as the type. Orenburg is situated due W of the southern

Yuzh Ural Mts.

The specimen differs in no way from a rather small-sized M. luctuosa and in all

probability is conspecific with that species. Its pygidial plate (fig. 112) is shaped

similarly, with somewhat raised lateral margins and a swollen, bluntly carinated

apex; surface dull, finely tessellate, with few punctures on the basal portion, dark

chestnut coloured.

Male. — Unknown.

336 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

Melecta ashabadensis Radoszkowski

(Figs 113)

Melecta ashabadensis Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3): 180 (9 —

excl. 3 ? — Ashabad). — Friese, 1895, Bienen Europa’s: 164 (original descr. quoted; for notes on

additional material, see M. fulgida spec. nov.!).

Type material. — USSR, Turkmeniya: | 9, Ashkhabad (or Ashkabad), but

labelled ““Astr.” (print on purple square)!), “Type” (print on red), ‘‘ashabadensis”

(Radoszkowski’s pencil writing), “Melecta ashabadensis Radosz. Type, rev. Dr.

Enderlein”. Holotype, evidently designated as such by Enderlein (MNB).

A small species. The type is a badly soiled specimen lacking its left antenna and

all tarsal claws as well, save those on right fore and left hind legs. The male of

ashabadensis could not be traced and is probably lost; there is nothing of any use in

the brief diagnosis which accompanies the poor sketches of its genital organs (loc.

Cit, fig. 32):

The following notes on the type may serve to a future recognition of this

nondescript taxon, provided that fresh topotypical examples of either sex may

come to hand.

Female (type). — Labrum a little longer than broad (14 : 10), narrowed toward

apex which, though rounded off, is slightly upturned and protuberant mesially;

surface coarsely punctate, clothed with the usual long raised black bristles.

Antenna slender; scape curved, not very hairy, all flagellar segments distinctly

longer than broad, 3 slightly longer than 4. White pubescence covering clypeus,

vertex and occiput white, becoming black laterally. Thoracic vestiture also white

anteriorly, scutellar area black with conspicuous tufts of white behind short,

pricker-shaped scutellar spines. Legs normal. Basal half of mid and hind tibiae

white externally, outer face of hind tibia dull, densely hairy with short spine-like

setae intermixed; tarsi all black; inner rami of mid and hind tarsal claws rather

broad, about half as long as outer. Wings smoky grey-brown, much darker than in

turkestanica (3), third submarginal cell rather long, scarcely higher than its median

length (10: 9), the distal angle not prominent, broadly rounded.

Abdomen short, cordate. All white lateral spots of large size, those on tergite 2

almost quadrangular, far apart, on 3 more prolonged inward and separated by a

distance about two-thirds their own breadth, on 4 nearer together; all spots

rounded off mesially. Pygidial plate chestnut coloured, slightly downcurved, finely

black-ridged laterally, apex a little broadened, black; surface somewhat shiny,

microscopically tessellate (fig. 113). Total length 9.0 mm, fore wing 7.5 mm.

Friese (loc. cit.), who received a male sub nom. ashabadensis from

Radoszkowski but failed to mention its locality, gave a brief description of it and

noted that, besides specimens from Kaukasus (Helenendorf) and Syria in the

Vienna museum, he had also seen a very dark male from Syria in the Budapest

museum. These examples were labelled as such by Friese but are not conspecific

with ashabadensis; all belong to a new species presently described as fulgida spec.

') Ashtrabad, to the west, is in northern Iran, nearer the Caspian Sea!

LIEFTINCK: Palaearctic Melecta 337

nov., which is altogether different from the type of ashabadensis (see p. 219).

Characterized by its small size, rather long third submarginal cell and relatively

large abdominal spots.

Male. — Not definitely known.

Melecta kashmirensis (Nurse)

Crocisa kashmirensis Nurse, 1903, Ann. Mag. Nat. Hist. (7) 11: 548—549 (9 “Kashmir, 5000—6000 ft.,

fairly common”.

Type and paratypic material. — India, Kashmir: | 9 (holotype), “Kashmir,

5—6000 ft., 5.01, Col. C. G. Nurse, B. M. 1920—72”, with holotype label 17B 1213

labels) (BM).

A West Himalayan species of doubtful identity.

Though very similar in general appearance and body size, these two small series,

all arranged under kashmirensis in the general collection, are probably not

conspecific, as shown by the differences. In the type and all others from Kashmir,

the white tergal spots are subrectangular, widely separated by black, on 1 and 2

broader than long, on 3 placed distinctly more inward than the preceding ones and

slightly oblique (the interspace still about three times wider than the diameter of

one spot), those on 4 being separated by a distance one and one-third their own

diameter. These Kashmir females are less hairy than those from Quetta. In the

former the body pubescence is a little shorter while the tergal spots are relatively

smaller, composed of more compactly set hairs; moreover, there are hardly any

raised hairs midbasally which connect the spots at sides of tergite 1, the latter

being also more widely separated than in the Quetta examples, thus differing from

luctuosa. In the Quetta females, on the other hand, thin collar-like hairs are present

at base of tergite 1, as in /uctuosa: in fact I am unable to distinguish the latter from

that species. However, even these luctuosa-like females are not quite

homogeneous, the specimen from Peshin, for example, has somewhat darker

wings than those from Quetta, while the outer faces of mid and hind tibiae (only

basal two-thirds white in the others) are entirely white-haired, as are the tarsi.

Structurally, the females of either locality are inseparable, the pygidial plates

having the same form. To sum up, in the absence of any male from the above

localities, the status of both taxa must unfortunately be left undecided.

Male. — Unknown.

Melecta candiae Strand

“Melecta sp. (Candiae Strand n. ad int.)" Strand, 1915, Archiv f. Naturgesch. 81A (4): 166 (9 Candia).

type”, in E. Strand’s handwriting (EIB). Holotype.

A very puzzling bee, though probably correctly identified as a species of

Melecta. The description is detailed enough, but the insect itself is unlike anything

338 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

known to me in this genus. The unique female, which lacks parts of its legs, is

obviously an immature, possibly freshly emerged, almost completely hairless bee

of moderate size. The body integument is dullish dark brown. A very peculiar

feature is found in the colour of the posterior borders of the abdominal tergites

2—4, which bear unusually broad light brown bands, the rest of the integument

being contrastingly coloured dark brown.

M. candiae is one of the numerous annoying, objectionably proposed names for

a unrecognizable Strandian insect. In the writer’s opinion, names like this (‘‘Sollte

diese Art neu sein, so möge sie den Namen Candiae bekommen...’’), though

unfortunately still considered available by the ICZN in case of an eventual

validation, are best rejected.

Male. — Unknown.

ADDITIONAL REFERENCES

Only pages dealing with Melecta are cited.

For a more complete bibliography, see Lieftinck (1972).

Alfken, J. D., 1914. Beitrag zur Kenntnis der Bienenfauna von Algerien. — Mem. Soc. Ent. Belg. 22:

235—236.

——, 1926. Beitrag zur Kenntnis der Bienenfauna von Aegypten. — Senckenbergiana 8: 120.

——., 1931. Zur Kenntnis einiger palaearktischer Bienen und Beschreibung einer neuen Art von Te-

neriffa. — Konowia 10: 164—165.

——., 1936. Apidae in Schwed.-Chines. wiss. Exped. Nordw. Prov. Chinas. — Arkiv f. Zool. 27A: 21,

SIR

——.,, 1937. Beitrag zur Kenntnis der Bienenfauna von Mittel-Italien. — Boll. Ist. Ent. Univ. Bologna

CEE

, 1940. Zur Kenntnis der Nomada pusilla Lep. — Sitzber. Ges. naturf. Freunde 15. Jan. 1940: 243.

Baez, M. & G. Ortega, 1978. Lista preliminar de los Hymenöpteros de las Islas Canarias. — Bol. Asoc.

esp. Entom. 2: 194, Salamanca.

Cockerell, T. D. A., 1931. Bees collected by the Rev. O. Piel in China. — Amer. Mus. Novit. 466: 6.

Day, M. C. & M.G. Fitton, 1977. Discovery in the Linnaean collection of type-material of insects des-

cribed by Johann Reinhold Forster, with notes on the Hymenoptera. — Biol. J. Linn. Soc. 9:

33— 34, 40.

Day, M. C., 1979. The species of Hymenoptera described by Linnaeus in the genera Sphex, Chrysis,

Vespa, Apis and Mutilla. — Biol. J. Linn. Soc. 12: 48.

Fahringer, J. & H. Friese, 1921. Eine Hymenopteren-Ausbeute aus dem Amanusgebirge (Kleinasien

und Nordsyrien, südl. Armenien). — Archiv f. Naturgesch. 87 A (3): 168, 176.

Frey-Gessner, E., 1907. Hym. Apidae |. — In Fauna insect. helvetiae, Schaffhausen: 377.

Friese, H., 1922. Eine Kriegsausbeute an Apiden (Bienen) aus Makedonien. — Zool. Jahrb. Abt. Syst.

etc. 46: 207.

——, 1925. Neue Formen von Schmarotzerbienen, besonders aus dem paläarktischen Gebiet. —

Konowia 4: 27—29.

Gribodo, G., 1893. Note imenotterologiche II. Nuovi generi e nuove specie di Imenotteri antofili ed os-

servazioni sopra alcune specie gia conosciute. — Bull. Soc. Ent. Ital. 25: 406—414.

Hurd, P. D., 1953. Notes on the Melectine bees of Mexico (Hymenoptera, Anthophoridae). — J. Kan-

sas Ent. Soc. 26: 35—37, map.

Kohl, F. F. & A. Handlirsch, 1889. Transcaspische Hymenopteren. — Verh. Sitzber. Zool. bot. Ges.

Wien 39: 273—274.

Kullenberg, B., 1961. Studies in Ophrys pollination. — Zool. Bidr. Uppsala 34: 1—340.

Kullenberg, B. & G. Bergström, 1976. Hymenoptera aculeata males as pollinators of Ophrys orchids. —

Zool. Scripta, Stockholm 5 (1): 15, 22, fig. 10.

Lieftinck, M. A., 1958. A preliminary account of the bees of the Canary Islands (Hym., Apoidea). — In

LIEFTINCK: Palaearctic Melecta 339

Entom. Results Finnish Exped. Canary Is. 1947—1951, no. 16. — Comment Biol., Helsingfors

18 (5): 20—30, pl. 1 figs. 9—11 & textfigs. 8—41.

——., 1972. Further studies on Old World melectine bees, with stray notes on their distribution and

host relationships (Hymenoptera, Anthophoridae). — Tijdschr. Ent. 115: 253—322, 2 pls., 57

figs., 1 tab. & 2 maps. (With full bibliography).

Michener, C. D. & Anne Fraser, 1978. A comparative anatomical study of mandibular structure in

bees. — Univ. Kansas Sci. Bull. 51: 463—482, figs.

Nurse, C. G., 1903. New species of Indian aculeate Hymenoptera. — Ann. Mag. Nat. Hist. (7) 11:

548— 549,

Panzer, G. W. F., 1799. Faunae Insect. Germ. init. 6 fasc. Heft 70, pl. 22.

Radoszkowski, O. I., 1876. Comte-Rendu des Hyménoptères receuillis en Egypte et Abyssinie en 1873,

par Général d’Artillerie O. Radoszkowsky. — Hor. Soc. Ent. Ross. 12: 96—97 (122—124), pls,

figs.

——., 1877. Matériaux pour servir à une faune hyménoptérologique de la Russie (Suite). — Hor. Soc.

Ent. Ross. (1876) 12: 333—335, pl. 2.

———, 1886. Faune Hyménoptérologique Transcaspienne. — Hor. Ent. Soc. Ross. 20: 18, pl. 3 fig. 15.

— — 1893. Revue des armures copulatrices des mâles des genres: Crocisa Jur., Melecta Lat., . . . etc.

— Bull. Soc. Imp. Nat. Moscou, n.s. 7: 163—188, pls.

Rossi, P., 1790. Fauna Etrusca...sistens Insecta... provinciis Florentina et Pisana praesertim collegit

P. Rossius, Liburni, 2: 110.

Rozen, J. G., 1969. The larvae of the Anthophoridae (Hymenoptera, Apoidea, Part 3. The Melectini,

Ericrocini, and Rhathymini [= Ctenioschelini]. — Amer. Mus. Novit. 2382: 1—24, 56 figs.

Schaeffer, J. C., 1766. Icones Insectorum... etc. Natürlich abgemahlte Abbildungen Regensburgi-

scher Insecten 1 (1) Regensburg 1766—79, tab. 22 fig. xiv.

Strand, E., 1919. Uber einige Apidae des Deutschen Entomologischen Museums. — Archiv f. Natur-

gesch. 83: 61—62.

Thorp, R. W., 1969. Ecology and behaviour of Melecta separata callura (Hymenoptera, Anthophoridae).

— Amer. Midland Nat. 82: 338— 345, 7 pls.

Tsuneki, K., 1973. Studies on Nomada of Japan (Hym., Apidae). — Etizenia, Occas. Publ. Biol. Lab.

Fukui Univ. 66: 1—83, figs.

340 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980

INDEX

Unless indicated otherwise all specific and subspecific names are of Melecta. Synonyms, homonyms

and otherwise invalid names are in italics.

Species incertae sedis are (bracketed). New taxa are marked with an *asterisk.

acervorum, Anthophora — 133, 200, 203 *fumipennis — 158, 178 (keys), 248

acutivalvis — 308, 314 fumipennis, Anthophora — 251

aegyptiaca — 169, 184 (keys), 244 funeraria, Eupavlovskia — 313

aestivalis, Anthophora — 203 gigantea — 193, 196

albifrons — 148 (key), 193 glasunowi, Paracrocisa — 261

albovaria — 149 (key), 196 *sracilipes — 176 (key), 332

*alcestis — 164 (key), 270 grandis — 165, 185 (keys), 286

*alecto — 164, 186 (keys), 273 *guichardi — 162, 179 (keys), 254

alluaudi, Anthophora —321 hispanica, Anthophora — 210, 286, 290

*amanda — 161, 182 (keys), 266 hispanica — 194

anglicana — 194 histrionicus, Thyreus — 194

*angustilabris — 157, 177 (keys), 251 *honesta — 161, 182 (keys), 264

armata — 193, 202 italica — 174, 191 (keys), 305, 309

(ashabadensis) — 192, 336 jakovlewi(i) — 151 (key), 215

assimilis — 171, 187 (keys), 318 jakowlevi — 215

aterrima — 204, 206 (kashmirensis) — 192, 337

baeri(i) — 171, 187 (keys), 282—283 kuschakewiczi, Paracrocisa — 261

bipunctata — 204, 206 lentina — 331

bituberculata — 306 leucorhyncha — 176, 192 (keys), 308, 325, 330

*brevipila — 156, 191 (keys), 235 lindbergi — 244

caesareae — 275, 282 luctuosa — 155 (key), 227, 330

calabrina — 204 mediterranea — 197, 313

canariensis — 192 (key), 322 *megaera — 175, 185 (keys), 243

(candiae) — 337 meridionalis — 308, 313

*candida — 170 (key), 268 minima — 197

canescens, Anthophora — 133 nigra — 149 (key), 204, 206

caroli — 175, 190 (keys), 323 (nigripennis) — 206, 334

chinensis — 150 (key), 218 nivosa — 157, 179 (keys), 257

corpulenta — 160, 181 (keys), 260 Nomada — 143, 213

crassicornis — 309, 315 notata — 205

crassipes, Anthophora — 133 novellai — 197

crinipes, Anthophora — 202, 219 (octomaculata) — 279, 334

Crocisa atra — 139, 204, 206 octomaculata — 275

curvispina — 173, 189 (keys), 320 orbatus, Thyreus — 194

diacantha, Pseudomelecta — 239 orientalis — 215

dispar, Anthophora — 133, 203, 230 oulskii, Andrena — 283

duodecimmaculata — 151 (key), 210 parietina, Anthophora — 202

duodecimmaculata, Nomada? — 139, 194, 212 personata, Anthophora — 192, 202

ebusana — 254, 325 plagiata, Anthophora — 202

eczmiadzini — 228, 270 plurinotata — 213, 215

edwardsii, Anthophora — 132 *prophanta — 170 (key), 290

egyptiaca — 244 pseudoarmata — 197

Eupavlovskia obscura simulatrix — 134 Pseudomelecta baeri — 139, 266, 282

(eversmanni) — 192, 228, 335 punctata — 193

*excelsa — 152 (key), 217 pygialis — 308

fasciculata — 204, 227, 233 quadripunctata — 309, 314

*festiva — 167, 186 (keys), 293 quadripustulata — 314

*fulgida — 158, 178 (keys), 219, 226 quatuordecimpunctata — 210, 213

fulvitarsis, Anthophora — 196, 203 retusa, Anthophora — 233

LIEFTINCK: Palaearctic Melecta 341

robusta, Anthophora — 289 *solivaga — 180 (key), 279

robusta — 260, 262 taormina — 177, 192 (keys), 330

rufipes — 204 (testaceipes) — 206, 334

rutenica — 153, 190 (keys), 233 Thyreus — 143

separata callura — 132 tibiopunctata — 331

sexta, Apis — 194 transcaspica — 168, 183 (keys), 275

sibirica — 173 (key), 238 Triepeolus — 138

sicheli, Anthophora — 331 *tuberculata — 134; 167, 188 (keys), 300

soederbomi — 210, 213 turkestanica — 172, 189 (keys), 239

342 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980 Presi

MBUD); fig. 3, M. duodecimmaculata duodecimmaculata (Rossi), & Athens, Greece (MH); fig. 4,

M. excelsa spec. nov., 4 holotype Jalalabad, Afghanistan (ML, ex MMB); fig. 5, M. fulgida spec. nov.,

d paratype Sandanski, Bulgaria (ML); fig. 6, same species, 3 Beirut, Lebanon (ML)

Pe LIEFTINCK: Palaearctic Melecta 343

= 3

Sy; a.

11 12 I |

Fig. 7, Melecta brevipila spec. nov, & holotype Vallis Kabak, Turkestan (MBUD); fig. 8, M. turkestanica

Radoszkowski, 3° lectotype, Boc-cy, Turkestan (MNB); fig. 9, same species, 4 ‘““Alai”,? Turkestan

(MP); fig. 10, same species, 9 topotypical (same label, MP); fig. 11, M. alcestis spec. nov., ¢ holotype,

“Jiz Rusko Toscoe”, S Russia (NMP); fig. 12, M. baeri (Radoszkowski), ¢ first described, Krasno-

wodsk, Turkestan (MBUD)

344 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980 PL. 3

17 18

Fig. 13, Melecta fumipennis spec. nov., 3 holotype, Wadi Gerani, Egypt (ML); fig. 14, M. angustilabris

spec. nov., ¢ holotype, Fayed, Egypt (ML); fig. 15, M. corpulenta Morawitz, ¢ Margelan, Turkestan

(ML, ex ZIL); fig. 16, M. honesta spec. nov., & holotype, Mut, Turkey (ML, ex CG); fig. 17, M. amanda

spec. nov, & holotype, Turkestan (MCG); fig. 18, M. candida spec. nov., ¢ holotype, Quetta,

Baluchistan (BM)

PL. 4 LIEFTINCK: Palaearctic Melecta 345

23 24

Fig. 19, Melecta aegyptiaca Radoszkowski, 3 Egypt (ML, ex SMF); fig. 20, same species, Z Caffarella,

Italy (INER); fig. 21, same species, 4 Sturovo, Slovakia CSR (CK); fig. 22, M. transcaspica Morawitz,

3 holotype, Chodshakala, Turkmenia (ZIL); fig. 23, same species, 9 Denizli, Turkey (SMF); fig. 24,

same species (holotype M. caesareae Friese), Q “Caesarea Palästina’, Israel (MNB)

346 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980 BS

Fig. 25, Melecta grandis Lepeletier, 9 lectotype, Oran, Algeria (MP); figs. 26—27, same specimen, late-

ral and frontal view; fig. 28, same species, 9 Algeria (BM); fig. 29, same species, ¢ Medinine, Tunisia

(MP); fig. 30, same species, & Sicily (NM W)

PL. 6 LIEFTINCK: Palaearctic Melecta 347

Fig. 31, Melecta festiva spec. nov., ¢ Caucasus, USSR (“eczmiadzini” sec. Friese, nec Radoszkowski)

(MNB); fig. 32, same species, 4 “Mesopotamia Castilia 1886” (MBUD); fig. 33, M. italica Radosz-

kowski, 3 (syntype M. grandis Lepeletier, 3 Oran, Algeria) (MP); fig. 34, same specimen, lateral view;

fig. 35, M. tuberculata spec. nov., 3 Vadillo de Castril, Spain (ML)

348 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 123, AFL. 6, 1980 Pi

Figs. 36—37, Melecta tuberculata spec. nov., 3 Cyprus, showing 8 orchid pollinia adhered to frontal

area of head, the same specimen photographed from two slightly different angles of view (BM)

PL. 8 LIEFTINCK: Palaearctic Melecta 349

40 41

Fig. 38, Melecta italica Radoszkowski, & Italia mer. (ML); fig. 39, same species, g Llobregat, Spain

(MP); fig. 40, M. assimilis Radoszkowski, 3 holotype, Egypt (MNB); fig. 41, M. leucorhyncha taormina

Strand, g Segesta, Sicilia

di È n ie a an i VOR

LI CARL) 1 TORI i;

Vit OMS ie Bik

M te ar, AU A

rhe | un

i ven 1: ij 0 Wi i

iw EN AN

EAN mR Ag |

vi } A i

OLI

4 iz = z N, A =

2 3 SK 5 2 |

: = =, = x 2 =

> 7 = > = ASE =

7 (77) 2 u AE 7) x

LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31YVY81

z 7 5 IE:

4 Sade 3 = a =

= Ce ARS 4 ~ ac =

1 < PH = < = <

5 x ‘45, = = = =

A dl: : = :

z D ji z i z oe

_ NOIINLILSNI _NVINOSHLIWS _ S3 IUVUSE LL LIBRARI ES_ SMITHSONIAN INSTIEG TIE

5 Ai 6 5 Se Ne

= > = œ = œ

> 0 5 2 5 a

= > N 3 N i > = p= =

= 6 N N = _ :

= WS > a >

mn N È 5 2 5

LIBRARIES SMITHSONIAN INSTITUTION

da x

NOILNLILSNI NVINOSHLINS S31#VH9171 LIBRARIES SMITHSONIAN INSTITUTIO!

NOILNLILSNI NVINOSHLINS SIIUVUGI

NVINOSHLINS

SMITHSONIAN

SM ITHSONIAN

NYINOSHLINS

LIBRARIES oil FISUNITAIN

| Zz u = un >

u u ui ;

= = = < À LA aa

= œ E œ rs €

z a z E 2

LIBRARI EMS nia TERRES NVINOSHLINS SIIUVUGI

6 = S E

= * » + r

>, > > >

nr x 5 =

= og = E

zm 6 m Di si

= u = , =

NOILNLILSNI NVINOSHLINS S314#VY911 LIBRARIES SMITHSONIAN

NVINOSHLINS ssatavadtt

NVINOSHLINS S31UVY 417

NVINOSHLIWS

N NS

SMITHSONIAN

LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31UVY8

5 di RT 5 wu 2 ON ul

ae, x LÀ / Z — oe st A N

3 <W 7, = < ZN“ <

È 4}, S E SS €

5 o 7 5 a = =

>= si = — > AE

NOILNLILSNI NVINOSHLINS S31YVY9171 LIBRARIES SMITHSONIAN_INSTITUTIO

= wo = o = Lt œ

5 = 5 20 5 Gly, =

= EE - 2 >. eR Se

vw m \ 2 m n° | m

= u = 7) = v

LI B RARI ES „SMITH ONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYVY8I

= Ar = ae | = Wi < a È Ws

= AUS EN IR 4 rho ETS ONU

z We + KA) = NY = Ag, (EMI I My

SMITHSONI

LIBRARIES

SIIUVUGEII

\RIES SMITHSONIAN

NVINOSHLIWS

LILSNI_ NVINOSHLINS

NOILNLILSNI

\RIES SMITHSONIAN

INSTITUTION

ARIES SMITHSONIAN

SMITHSONIAN

LIBRARIES

LILSNI_NVINOSHLIWS

— ry = R NN — _—

ae SNO = 5

È 4 2 N N S 4

= = N 2 =

in age ii ne

ARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IUVY 811

= n 5 Le ©

È = 2 à

A >= ISIN

< =| N <q

- È E BA È

È z 3 = Bee

LILSNI NVINOSHLIWS _ S23 IUVYE i LIBRARI ES_ SMITHSONIAN INSTITUTION

À ei = o ig, ©

N 5 0 = GL) 2

NY È = = Ly"

WY HE a -_ wi | ee,

NY 2 m 5 m

= u = v

INSTITUTION NOILNLILSNI NVINOSHLINS SIIUVYYAIT

Z x u z un È

< = i < =

= Ng F = =

E NS 2 = Z

EN > = 3

n°3 Iuvyg 17_LI BRARIES SMITHSONIAN INSTITUTION

w 2 + Z

= u = u

= = < =

œ “i ce =

= o aah ©

SJ z 3 =

INSTITUTION NOILNLILSNI NVINOSHLINS S31IYUVY911

= 5 = =

“0 = œ —

2 = 2 5

= = =? =

= F 2 E

| zov Z n 2

LILSNI NVINOSHLINS S 3 14vVH4a QU BRARI ES SMITHSONIAN _ INSTITUTION

= < we = Hi NS

7 ‘6 = = S

D : D : a a N

O 4 Ja oO 2 5

z = z E

hy ae Pe cie

INSTITUTION NOILNLILSNI NVINOSHLINS S31IUVY817

. + = N NS =

VA ZANE = = = N A

Zs = a Ge N =

Vy) a È = NME

i 2 5 = oe

S31YVYg911 LIBRARIES SMITHSONIAN

5 a E

= w È A, Ce

= > = 3 TA >

E 2 E Vf

2 n SION m

— [477 —_

INSTITUTION NOILNLILSNI NVINOSHLINS S314vVY917

È ae Le] z a

N = Wy = EE = Y

x = LY, iS ‘Uy,

ER NE Ag. & (AN) = Ty.

Hang SIIUVUAI

NVINOSHLIWS SIIUYVYEI1 LIBRARIES SMITHSONIAN

NVINOSHLI

INSTITUTION NOILNLILSNI

INSTITUTION NOILNLELSNI

CONIAN