European Journal of Taxonomy 28: 1-28

http://dx.doi.org/10.5852/ejt.2012.28

ISSN 2118-9773

WWW. europeanj oumaloftaxonomy. eu

2012 • Pauly A., Gibbs J. & Kuhlmann M.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

um;lsid;zoobank.org:pub:936AA3AA-lC2D-40B3-BE84-74AC42C5E82A

Capalictus, a new subgenus of Lasioglossum Curtis, 1833 from

South Africa, with description of three new species

(Hymenoptera, Apoidea, Halictidae)

Alain PAULY' , Jason GIBBS" & Michael KUHLMANN"

^ Royal Belgian Institute of Natural Seienees, Department of Entomology,

Rue Vautier 29, B-1000 Brussels, Belgium.

Corresponding author. Email: [email protected]

^ Cornell University, Department of Entomology, 3119 Comstoek Hall, Ithaea, NY, USA, 14853.

Email: [email protected]

^ The Natural History Museum, Department of Entomology, Cromwell Road, Eondon, SW7 5BD, UK.

Email: [email protected]

^ um:lsid:zoobank.org:author:0A734BB3-61Bl-489F-995F-FF6161A58C16

^ um:lsid: zoobank, org: author:B A42A49F-3EBC-4679-8F 03-A5 8E7981Q6B1

^ um:lsid:zoobank.org:author:B99AE0ED-FA89-4DFE-A658-lC8DF37F9FAB

Abstract. Capalictus, a new subgenus of Lasioglossum Curtis, 1833 (Hymenoptera, Apoidea, Halie-

tidae), endemie to the South Afriean Cape Provinee, is deseribed. The type speeies is Halictus mosselinus

Coekerell, 1945. Evylaeus (Sellalictus) jynbosensis (Pauly et al, 2008) is a new junior synonym of

L. (C.) mosselinum. Three new speeies are deseribed: Lasioglossum (Capalictus) hantamense sp. nov.,

L. (C.) tigrinum sp. nov. and L. (C.) timmermanni sp. nov. DNA sequenee data from three nuelear genes

support morphologieally-determined speeies limits. Capalictus is a basal elade of the Hemihalictus

series of Lasioglossum.

Key words, taxonomy, Anthophila, new speeies.

Pauly A., Gibbs J. & Kuhlmann M. 2012. Capalictus, a new subgenus of Lasioglossum Curtis, 1833 from South

Africa, with description of three new species (Hymenoptera, Apoidea, Halictidae). European Journal of Taxonomy

28: 1-28. http://dx.doi.org/10.5852/eit.2012.28

Introduction

We deseribe a new subgenus of Lasioglossum, endemie to the Cape Provinee of South Afriea. This new

subgenus was first deseribed as a new speeies-group of Sellalictus Pauly, 1980 by Pauly et al. (2008),

based on a single new speeies, ''Evylaeus (Sellalictus) jynbosensis Pauly, Timmermann & Kuhlmann,

2008”. Diseovery of at least three new speeies belonging to the same speeies-group as L. fynbosense

(eomb. nov.) in eombination with reeent moleeular phylogenetie results (Gibbs et al. 2012) ineline

us to deseribe this group rather as a distinet subgenus. A reeent moleeular phylogeny of halietid bees

demonstrated with strong support, that this group eomposes a basal elade of the Hemihalictus series

European Journal of Taxonomy 2^\ 1-28 (2012)

(Gibbs et al. 2012). This position on the phylogenetic tree strongly supports subgeneric status for these

species.

Material and method

Terminology

Terminology for morphological characters follows Engel (2001) and Michener (2007). Terminology

for the ‘propodeum’ and ‘metapostnotum’ follows Gibbs (2011). We used the glossary of Harris (1979)

for description of the surface sculpture (except “shagreened” used here in the sense of “dull due to

the presence of microsculpture”). Puncture density is given in terms of relationship between puncture

diameter (d) and the spaces between them (i), such as i>d. Hair length is given in relative units based

on the median ocellar diameter (OD). Metasomal sterna and terga, and flagellomeres are abbreviated

S, T, and F, respectively, followed by the appropriate number. The following abbreviations are used

throughout: upper interocular distance (UOD), lower interocular distance (LOD), ratio of mesoscutellum

and metapostnotum lengths (MMR).

Measurements

Measurements were taken on enlarged pictures or directly using an ocular micrometer.

Collections

Material is preserved in the following collections:

BMNH = The Natural History Museum, London, United Kingdom

CUIC = Cornell University Insect Collection, Ithaca, USA

RBINS = Royal Belgian Institute of Natural Sciences, Brussels, Belgium

SANC = South African National Collection of Insects, Pretoria, South Africa

No collection numbers are allocated to specimens.

DNA analysis

DNA was extracted and amplifred following protocols explained elsewhere (Danforth 1999; Danforth et

al. 2003, 2004) from frve specimens belonging to the new subgenus described below as part of ongoing

studies of the phylogenetic relationships of halictid bees (Gibbs et al. 2012; Gibbs unpublished data.).

Fragments of three nuclear genes were sequenced in both directions (elongation factor-1 alpha: ef-la,

wingless: wnt-1, and long-wavelength rhodopsin: opsin) using Big Dye Terminator for the sequencing

reaction. Sequencing was performed at the Cornell University Life Sciences Core Laboratories

Center using Applied Biosystems Automated 3730 DNA analyzers. Sequences were assembled using

Sequencher (GeneCodes).

Results

Classis Hexapoda Blainville, 1816

Ordo Hymenoptera Linnaeus, 1758

Superfamilia Apoidea Latreille, 1802

Epifamilia Anthophila Latreille, 1804

Familia Halictidae Thomson, 1869

Subfamilia Halictinae Thomson, 1869

Tribus Halictini Thomson, 1869

GQxms Lasioglossum Curtis, 1833

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Lasioglossum {Capalictus) subgen. nov.

um:lsid:zoobank.org:act:D6B54F16-5465-4163-BCCE-C89B76E04C3C

Type species

Halictus mosselinus Cockerell, 1945.

Description

Capalictus is distinct from other African subgenera of the Hemihalictus series by the combination of

following characters: small size (length 6-7.5 mm). Body black, non-metallic, metasoma red in some

species (this red colouration is unique in African subgenera of the Hemihalictus series, with the exception

of the kleptoparasitic L. ereptor). Metasomal terga without patches of tomentum. Mesoscutum smooth-

shiny, without tessellate surface. Eower paraocular area shiny and sparsely punctate. Propodeum

posterior surface weakly carinate in females, not carinate in males. The carina in females not extending

medially across upper margin (Fig. IF). Clypeus and mandibles of males black. Inner metatibial spur in

females with about five short teeth, the one in males without teeth. Gonocoxite of males without retrorse

lobe, gonostyli pointed, half as long as gonocoxite (Fig. 15). Apex of marginal cell pointed, ending on

wing margin (Fig. 1C, E).

Differential diagnosis

The new South-African subgenus Capalictus belongs to the Hemihalictus series {sensu Michener 2007)

of the genus Lasioglossum Curtis, 1833, which is characterized by weak cross-submarginal veins Irs-

m and 2rs-m of forewing (Fig. IC-E). Capalictus is morphologically close to the Holarctic subgenera

Evylaeus Robertson, 1902 and Dialictus Robertson, 1902, and to the African subgenera Sellalictus

Pauly, 1980 and Afrodialictus Pauly, 1984 (see classification of Pauly 1999). From Sellalictus it

differs by the absence of a patch of tomentum on the metasomal terga (males of Sellalictus bear a

very characteristic large patch of tomentum on base of tergum 2 while females have small baso-lateral

spots of tomentum on T2-T4). Capalictus also differs by the apex of the marginal cell close to wing

margin (Fig. IC-E) (in Sellalictus the apex of the marginal cell is minutely separated from the wing

margin and minutely appendiculate as in Fig. ID). Yxom Afrodialictus, the new subgenus differs by the

lack of microtessellate texture on the body (microtessellate surface sculpture is characteristic on head,

mesoscutum and propodeum of Afrodialictus', in Capalictus, mesoscutum is polished and shiny), lower

part of paraocular area shiny and with some punctation (Fig. 1 A) (with Afrodialictus the lower parts of

the paraocular area has a different dull, frosted-like and completely impunctate surface as in Fig. IB)

and posterior face of propodeum with a very weak carina in the female (Fig. IF) (carina entirely lacking

in Afrodialictus). Capalictus can be separated from the African subgenus Mediocralictus Pauly, 1984 by

the inner metatibial spur of the males without teeth (pectinate in males of Mediocralictus). Females of

Mediocralictus can be separated by the peculiar shape of the propodeal carina (Pauly et al. 2001: 121,

fig. 36). Capalictus can be separated from most Palaearctic species of the Hemihalictus series by the

gonocoxites of males without retrorse lobe. Palaearctic Evylaeus lacking a retrorse lobe can be easily

distinguished from Capalictus by the carinate propodeum (male Capalictus lack distinct propodeal

carinae) and metasomal sterna with very sparse, and short setae (<OD) (sternal hairs sometimes long

(2-2.5 OD) and plumose in Capalictus).

Lasioglossum {Capalictus) mosselinum (Cockerell, 1945)

Figs 2, 3, lOA-B, 12A, 13A, 14A-B, 15A, 16A, E, 17A

Halictus mosselinus Cockerell, 1945: 280-281.

Patellapis (Chaetalictus) mosselinaFmily, 1999: 150, 174.

Evylaeus {Sellalictus) Jynbosensis Pauly, Timmermann & Kuhlmann, 2008: 85-91. Syn. nov.

Lasioglossum mosselinum - Timmermann & Kuhlmann 2009: 180.

European Journal of Taxonomy 2^\ 1-28 ( 2012 )

Type material

Halictus mosselinum: Holotype SOUTH AFRICA, Cape Province, Mossel Bay, Sep. 1933 (white

label), S. Africa, R.E. Turner, Brit. Mus. 1933-552 (white label). Type (red-white label), B.M. TYPE

HYM. 17a. 1109 (white label), Halictus mosselinus Ckll Type (handwritten white label) (BMNH),

examined in January 2012 .

Fig. 1. —Lasioglossum (Capalictus) mosselinum (Cockerell, 1945). A. Head showing lower paraocular

area. C, E. Wing venation showing apex of marginal cell and weak veins of submarginal cells. C. $.

E. (?. F. $, propodeum and metapostnotum (dls = dorso- lateral slopes). — Lasioglossum (Afrodia-

lictus) sp. B. Head showing lower paraocular area. — Lasioglossum (Sellalictus) sp. D. $, wing venation

showing apex of marginal cell and weak veins of submarginal cells. Scale line = 0,5 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Evylaeus {Sellalictus) Jynbosensis: Holotype SOUTH AFRICA, 15 km NW Nieuwoudtville, Farm

Engelsepunt, Fynbos, Pf El, 3I°I4’3I“ S - I8°59’08” E, 830 m, 28 Aug. 2003, leg. K. Timmermann

(SANC). Paratypes: 31 34 33, together with holotype, different dates at same locality (BMNH,

SANC, RBINS).

Additional material

17 33 , 1 ?, SOUTH AFRICA, WCP, 16 km E. Clanwilliam, Pakhuis Pass, 700 m, 32°08.rS -

18°59.7’E, 8 Sep. 2001, coll. CUIC.

Fig. 2. Lasioglossum {Capalictus) mosselinum (Cockerell, 1945), $ holotype. A. Total view. B. Head.

C. Mesosoma. D. Propodeum. E. First tergum. F. Metasoma. Scale line = 0,5 mm.

European Journal of Taxonomy 2^\ 1-28 (2012)

Diagnosis

Male

Body length 7.5 mm. Wing length 6 mm. Legs very long, espeeially tarsi (length of posterior tarsi equal

to length of mesosoma), totally blaek to blaekish brown exeept foretarsi yellowish brown (Figs 12A,

13A, 14A-B) (all tarsi yellowish in L. hantamense andZ. tigrinum). Metapostnotum with longitudinal

ridges. Dorsolateral slopes of propodeum shining, punetate. Punetation of mesoseutum sparse between

Fig. 3. Lasioglossum (Capalictus) mosselinum (Coekerell, 1945), f. A. Head. B. Mesoseutum and

vertex. C. Propodeum. D. First tergum. E. Metasoma, dorsal view. F. Metasoma, ventral view. Seale

line = 0,5 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

parapsidal lines (i = 3-4d) (Fig. 2C) (denser in L. timmermanni). Metasoma blackish, sometimes partly

orange red (Fig. 3E). Gonostylus with long apical setae (longer than basal width of gonostylus).

Female

Body length 7 mm. Wing length 6 mm. Metasoma red, base of T1 and T5-T6 blackish (Figs 2E-F, 16E)

(metasoma largely black in L. hantamense and L. tigrinum; T5 red in L. timmermanni). Metapostnotum

with longitudinal ridges, dorsolateral slopes of propodeum shining (Fig. 2D) (slopes shagreened in

L. timmermanni). Punctation of mesoscutum sparse between parapsidal lines (i = 2d) (Fig. 2C).

Lasioglossum (Capalictus) hantamense sp. nov.

um:lsid:zoobanlc.org:act:8743F44B-A7E0-4D61-ACDF-2812BC6FA43D

Figs 4, 5, llA-B, 12C, 13C, 14E-F, 15C, 16C, 17B

Diagnosis

Male

Close to L. mosselinum but all tarsi pale yellow and not so long (Fig. 12C) (mid and hind tarsi brown

in L. mosselinum and L. timmermanni). Metapostnotum with rugae extending 2/3 distance to posterior

margin (extending 4/5 distance inL. tigrinum).

Female

Differs from L. mosselinum and L. timmermanni by black metasoma (Figs 5E, 11 A) and dull shagreened

dorso-lateral slopes of propodeum (Fig. 5C). T1 nearly impunctate, punctures very fine, indistinct

(numerous distinct punctures in L. tigrinum).

Etymology

The specific epithet refers to the Hantam Mountains near Calvinia where the species was first discovered.

Type material

Holotype

7 Sep. 2010, leg. M. Kuhlmann (SANC).

Paratypes

4 and 6 ??, idem, 30 Aug. 2011, 28 (?(? and 10 ?? (Coll. Kuhlmann, BMNH, RBINS, SANC,

CUIC).

Type locality

SOUTH AFRICA, Plateau Hantam Mts., near antenna, 9 km N Calvinia, 31°22’29''S 19°47’03''E,

1570 m.

Description

Male

Body length. 6.5-7 mm.

Forewing eength. 5.5 mm.

CoEOURATiON. Head and mesosoma black. Metasomal terga black with apical rim translucent straw-

coloured. Clypeus and mandibles completely black. Flagellum with ventral surface black. Wing

membrane subhyaline, venation, pterostigma and tegula brown. Eegs black except femoral apices and

tibial bases brown and all tarsi pale yellow.

European Journal of Taxonomy 2^\ 1-28 (2012)

Pubescence. Face below antennal soekets with dense appressed greyish white setae. Long ereet setae on

seape, vertex and gena (2-2.5 OD). Metanotum, side of mesosoma, propodeum on lateral and posterior

surfaees with sparse, short, greyish white and plumose setae (2-2.5 OD). Metasomal terga without

patehes of tomentum but with sparse and short setae on lateral parts of terga. Metasomal sterna nearly

glabrous exeept some long setae laterally.

Fig. 4. Lasioglossum (Capalictus) hantamense sp. nov., f. A. Head. B. Mesoseutum and seutellum.

C. Propodeum and metanotum. D. First tergum. E. Metasoma, dorsal view. F. Metasoma, ventral view.

Seale line = 0,5 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Fig. 5. Lasioglossum {Capalictus) hantamense sp. nov., $. A. Head. B. Mesoscutum and scutellum.

C. Propodeum and metanotum. D. First tergum. E. Metasoma. Scale line = 0,5 mm.

European Journal of Taxonomy 2^\ 1-28 (2012)

Surface sculpture. Clypeus and supraclypeal area finely and densely punetate. Frons with minute

eontiguous punetures (i <d) of velvet-like appearanee. Upper paraoeular area shagreened and irregularly

mieropunetate. Vertex rugulose to shagreened. Gena dull andruguloso-striate. Mesoseutum smooth with

punetures separated by puneture width (i = 1-1.5 d), punetures denser laterally of parapsidal lines (i =

0.5d). Mesoseutellum and metanotum more finely punetate (i = 0.5-Id). Preepistemum, hypoepimeral

area and metepistemum with minute eontiguous punetures. Mesepistemum with larger and more

widely spaeed punetures, spaees between punetures shining (i = 0.5d). Metapostnotum rugae relatively

weak, extending no more than 2/3 distanee to posterior margin. Dorso-lateral slopes of propodeum

dull shagreened. Lateral surfaee of propodeum dull and minutely roughened, indistinetly punetate. T1

moderately punetate (i = l-1.5d), impressed apieal margin with few with punetation similar to Tl, dise

of T4 and T5 with sparse piliferous punetures.

Structure. Head nearly as long as wide (length/width ratio = 0.94). Eyes eonverging below (UOD/LOD

ratio = 1.62). Flagellomeres long (F2-F10 length/diameter ratio 1.8-1.5). Vertex short. Gena narrower

than eye. Propodeum without earina, dorsal surfaee relatively long (MMR =1.8). Legs moderately long

(length of posterior tarsus equal to 0,7 length of mesosoma) (Figs 13C, 14E-F).

Terminalia. Gonostylus pointed apieally, laterally eovered with short setae (less than 1/4 basal width of

gonostylus) (Fig. 15C).

Female

Body length. 6-6.5 mm.

Forewing length. 5 mm.

Colouration. Head, mesosoma, metasoma and legs blaek. Metasomal terga blaek with apieal rim

translueent straw-eoloured. Wing membranes subhyaline, venation, pterostigma and tegula brown.

Pubescence. Greyish white and sparse. Metasomal terga without patehes of tomentum, apieal rim of

T2-T4 with sparse fringes.

Surface sculpture. Clypeus shiny with large and sparse punetures. Supraelypeal area shiny with fine

punetures. Frons dull and finely rugulose, with velvet-like appearanee. Paraoeular area and oeelloeular

area finely and densely punetate, semi-dull. Lower paraoeular area shining and sparsely punetate. Vertex

shagreened. Gena finely and densely strigose-punetate. Mesoseutum smooth and shining with sparse

punetures (i = l-2d), punetures denser anteriorly and laterally of parapsidal lines (i = d). Mesoseutellum

more finely and densely punetate, shiny medially. Metanotum dull matt with minute eontiguous punetures.

Preepistemum, mesepistemum and metepistemum dull matt, minutely roughened. Hypoepimeral area

minutely and densely punetate.

Metapostnotum. It is with weak anastomosing mgae extending no more than 2/3 distanee to posterior

margin. Lateral surfaee of propodeum minutely roughened and shagreened. Tl smooth and nearly

impunetate on dise, apieal margin finely punetate (i = 1.5d). T2 punetate on basal and apieal margins,

impunetate in middle of dise. T3-T4 with very sparse punetation.

Structure. Head nearly as long as wide (length/width ratio = 0.91). Eyes slightly eonvergent below

(UOD/LOD ratio = 1.09). Vertex short. Gena as wide as eye. Inner metatibial spur with five short teeth.

Metapostnotum moderately long (MMR = 1.25). Propodeum with very weak lateral and oblique earinae

eurved to the middle of the posterior surfaee.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Lasioglossum {Capalictus) tigrinum sp. nov.

um:lsid:zoobank.org:act:6924FBAB-4F53-4C34-95BA-306F59D2328F

Figs 6, 7, IIC-D, 12D, 13D, G-H, 15D, 17C

Diagnosis

S- Close to L. hantamense but all tibiae (except a dark central maculation on tibiae II and III), tarsi, apical

margins of terga and sterna yellowish orange (Fig. 12D). Mid and hind tarsi black in L. mosselinum and

L. timmermanni punctate in middle of disc as on apical margin (Fig. 7D), punctation of mesoscutum

denser (Fig. 7B). Differs from L. mosselinum andL. timmermanni by the largely black metasoma.

Etymology

The name refers to the distinctive yellow-orange banding of the male metasoma which is striped like a

tiger.

Type material

Holotype

S, 30 Aug. 2011, leg. M. Kuhlmann (SANC).

Paratypes

1 (J, 1 $ (Coll. M. Kuhlmann).

Type locality

SOUTH AFRICA, Plateau Hantam Mts., near antenna, 9 km N Calvinia, 31°22’29''S 19°47’03''E,

1570 m.

Description

Male

Body length. 7.5 mm.

Forewing eength. 6 mm.

CoEOURATiON. Head and mesosoma black. Terga black with the whole apical impressed margin amber

translucent. Metasomal sterna yellowish orange. All tarsi and tibiae (except a central dark maculation

on tibiae II and III) yellowish orange. Clypeus, mandible and ventral surface of flagellum black. Wing

membrane subhyaline, venation, pterostigma and tegula dark brown

Pubescence. Face below antennal sockets with dense appressed greyish white setae. Long erect setae on

scape, vertex and gena (2-2.5 OD). Metanotum, sides of mesosoma, propodeum on lateral and posterior

surface with sparse, short, greyish white and plumose setae (2-2.5 OD). Metasomal terga with sparse

and short setae on lateral parts of terga. Metasomal sterna nearly glabrous except some short and sparse

setae.

Surface scuepture. Clypeus and supraclypeal area finely and densely punctate. Frons with minute

contiguous punctures (i < d) of velvet-like appearance. Upper paraocular area shagreened and irregularly

micropunctate. Vertex rugulose to shagreened. Gena dull andruguloso-striate. Mesoscutum smooth with

punctures separated by one puncture width (i = d), punctures denser anteriorly and laterally of parapsidal

line(i = 0.5d) (Fig. 8b). Preepistemum, hypoepimeral area, metepistemum as well as mesepistemum

with minute, contiguous punctures. Metapostnotum rugae relatively weak, extending 4/5 distance to

posterior margin. Dorso-lateral slope of propodeum dull shagreened. Lateral surface of propodeum dull

and minutely roughened, indistinctly punctate. T1 moderately-densely punctate (i = 0.5-ld), impressed

European Journal of Taxonomy 2^\ 1-28 (2012)

apical margin with few minute punetures. T1 anterior surfaee smooth, without striations. Dises of T2 and

T3 with punetation similar to Tl, dises of T4 and T5 with sparser punetures.

Structure. Head nearly as long as wide (length/width ratio = 0.97). Eyes eonverging below (UOD/LOD

ratio = 1.70). Flagellomeres long (F2-F10 length/diameter ratio 1.6-1.4). Vertex short. Gena narrower

Fig. 6. Lasioglossum (Capalictus) tigrinum sp. nov., f. A. Head and forelegs. B. Mesoseutum and

seutellum. C. Propodeum and metanotum. D. Metasoma, dorsal view. E. First tergum. F. Metasoma

ventral view. Seale line = 0,5 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

than eye. Propodeum without carina, dorsal surface relatively long (MMR = 1.66). Legs moderately

long (length of posterior tarsi equal to 0,85 length of mesosoma) (Figs 13D, 14G-H).

Terminalia. Gonostylus pointed apically, laterally with some long setae (longer or subequal to basal

width of gonostylus) (Fig. 15D).

Fig. 7. Lasioglossum {Capalictus) tigrinum sp. nov., $. A. Head. B. Mesoscutum and scutellum. C. Pro¬

podeum and metanotum. D. First tergum. E. Metasoma. Scale line = 0,5 mm.

European Journal of Taxonomy 2^\ 1-28 (2012)

Female

Body length. 7 mm .

Forewing eength. 5 mm.

CoEOURATiON. Head, mesosoma and metasoma blaek. Metasomal terga blaek with apieal rim translueent

straw-eoloured. Wing membrane subhyaline, venation and pterostigma brown, tegula blaekish brown.

Pubescence. Greyish white and sparse. Apieal rim of T2-T4 with sparse fringes.

Surface scuepture. Clypeus shiny with large and sparse punetures. Supraelypeal area shiny with fine

punetures. Frons dull and finely rugulose, with velvet-like appearanee. Upper paraoeular area and

oeelloeular area finely and densely punetate, semi-dull. Lower paraoeular area shining and sparsely

punetate. Vertex shagreened. Gena finely and densely strigose-punetate. Mesoseutum smooth and shining

with sparse punetures (i = l-2d), punetures denser anteriorly and laterally of parapsidal line (i = d)

with minute eontiguous punetures. Preepistemum, and mesepistemum dull matt, minutely and densely

roughened. Hypoepimeral area and metepistemum minutely and densely punetate. Metapostnotum with

weak anastomosing rugae extending no more than 2/3 distanee to posterior margin. Lateral surfaee of

propodeum minutely roughened and shagreened. T1 smooth and distinetly punetate in middle of dise

as well as on apieal margin (i = L5d), smooth and impunetate anteriorly and laterally. T2 punetate in

middle of dise as on apieal margin. T3-T4 with sparse punetures.

Structure. Head nearly as long as wide (length/width ratio = 0.90). Eyes slightly eonvergent below

(UOD/LOD ratio = 1.13). Vertex short. Gena as wide as eye. Inner metatibial spur with five short teeth.

Metapostnotum moderately long (MMR = 1.56). Propodeum with very weak lateral and oblique earinae

eurved to the middle of the posterior surfaee.

Lasioglossum (Capalictus) timmermanni sp. nov.

um:lsid:zoobank.org:aet:BD39F9EB-D335-4B6F-BC5E-096313677BlF

Figs 8, 9, lOC-D, 12B, 13B, 14C-D, 15B, 16B, 17D

Diagnosis

Male

Close to L. mosselinum but punetation of mesoseutum mueh denser between parapsidal lines (i =

1-1.5d) (Fig. 8B) (sparse in L. mosselinum, 1 = 3-4d). Eegs eompletely blaek, tarsi shorter than those of

L. mosselinum (Figs 12B, 13B, 14C-D) (all tarsi pale yellow to yellow-orange in L. hantamense and

L. tigrinum, fore tarsus yellow in L. mosselinum).

Female

Close to L. mosselinum but metasoma with more extensive red eolouration on last tergum (Fig. 16F),

base of first tergum often blaek (metasomal terga blaek with pale apieal margins in L. hantamense and

L. tigrinum). Rugae of metapostnotum are finer and shorter than in L. mosselinum, dorso-lateral slopes

of propodeum with mierotessellate dull surfaee, not shiny as in L. mosselinum (Fig. 9C-D).

Etymology

The speeies is named for Kim Timmermann, formerly Munster (Germany), who eolleeted many of the

speeimens.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Type material

Holotype

S, 4 Sep. 2011, leg. Erhardt (SANG).

Paratypes

1 (?, 25 Aug. 2010, leg. M. Kuhlmann (BMNH). 4 ??, 17 Aug. 2011, coll. BMNH, RBINS. 10 ??

collected at Farm Engelsepunt, 16 km NW Nieuwoudtville, Fynbos, PfEl, 31°14’3E'S 18°59’08''E,

830 m, leg. K. Timmermaim (BMNH, RBINS): 1 ?, 2-26 Jul. 2003; 1 ?, 6 Aug. 2003; 2 ??, 27 Aug.

2003; 6 28 Aug. 2003, PfEl.

Fig. 8. Lasioglossum {Capalictus) timmermanni sp. nov., S- A. Head. B. Mesosoma and vertex.

C. Propodeum and metanotum. D. First tergum. E. Metasoma, dorsal view. F. Metasoma, ventral view.

Scale line = 0,5 mm.

European Journal of Taxonomy 2^\ 1-28 (2012)

Type locality

SOUTHAFRICA, FarmAvontuur, Fynbos, 12kmNWNieuwoudtville, 31°16’18''S 19°02’55''E, 770m.

Fig. 9. Lasioglossum (Capalictus) timmermanni sp. nov., A. Head. B. Mesoscutum and scutellum.

C. Propodeum. D. Metanotum. E. Metasoma. F. First tergum. Seale line = 0,5 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Description

Male

Body length. 7.5 mm.

Forewing eength. 6 mm.

CoEOURATiON. Head, mesosoma, terga, sterna, clypeus, mandible, antenna and legs black. Wing membrane

subhyaline, venation, pterostigma and tegula black.

Pubescence. Face below antennal sockets with dense appressed greyish white setae. Long erect setae

on scape, vertex and gena (2-2.5 OD). Metanotum, pleuron, propodeum on lateral and posterior surface

with sparse, short, greyish white and plumose setae (2-2,5 OD). Metasomal terga without patches of

tomentum but with sparse and short setae on lateral. Metasomal sterna nearly glabrous except some long

and sparse setae on lateral sides.

Surface scuepture. Clypeus and supraclypeal area finely and densely punctate. Frons with minute

contiguous punctures (i<d) of velvet-like appearance. Upper paraocular area shagreened and irregularly

micropunctate. Vertex and ocellocular area rugulose to shagreened. Gena dull and ruguloso-striate.

Mesoscutum smooth with punctures separated by more than a puncture width (i = l-1.5d), punctures

denser laterally of parapsidal lines (i = 0.5d) and anteriorly. Mesoscutellum more finely punctate

(i = 0.5-Id). Metanotum with contiguous fine punctures. Preepistemum, hypoepimeral area and

metepistemum with contiguous minute punctures. Mesepistemum with larger alveolate and contiguous

punctures. Metapostnotum rugae relatively weak and anastomosing, extending more than 4/5 distance

to posterior margin. Dorsolateral slopes of propodeum dull shagreened. Lateral surface of propodeum

dull with distinct dense punctures. T1 moderately punctate (i = l-1.5d), impressed apical margin without

punctures. T1 anterior surface smooth and without striations. Disc of T2 and T3 with punctation similar

to Tl, disc of T4 and T5 with sparse piliferous punctures.

Structure. Head rounded, nearly as long as wide (length/width ratio = 0.91). Eyes converging below

(UOD/LOD ratio = 1.69). Flagellomeres long (F2-F10 length/diameter ratio 1.5-1.7). Vertex short. Gena

narrower than eye. Propodeum without carina, metapostnotum relatively long (MMR = 1.42). Legs

moderately long (length of posterior tarsi equal to 0,8 length of mesosoma) (Figs 13B, 14C-D).

Terminaeia. Gonostylus pointed apically, with relatively long apical setae (subequal to basal width of

gonostylus) (Fig. 15B).

Female

Body eength. 7 mm.

Forewing eength. 6 mm.

CoEOURATiON. Head, mesosoma and legs black, metasoma red orange except Tl anterior surface black and

lateral parts of T2-T4 blackish. Wing membranes subhyaline, venation, pterostigma and tegula brown.

Pubescence. Greyish white and sparse. Metasomal terga without patches of tomentum.

Surface scuepture. Clypeus shiny with large and sparse punctures. Supraclypeal area shiny with fine

punctures. Frons dull and finely rugulose, with velvet-like appearance. Paraocular area and ocellocular

area finely and densely punctate, semi-dull. Lower paraocular area shining and sparsely punctate. Vertex

shagreened. Gena finely and densely strigose-punctate. Mesoscutum smooth and shining with sparse

punctures (i = 2-3d), punctures denser anteriorly and laterally to parapsidal line (i = d). Mesoscutellum

European Journal of Taxonomy 2^\ 1-28 (2012)

more finely and densely punetate, shiny medially Metanotum dull matt with minute eontiguous punetures.

Preepistemum, and mesepistemum dull matt, minutely and densely roughened. Hypoepimeral area and

metepistemum minutely and densely punetate. Metapostnotum with weak anastomosing rugae extending

no more than 2/3 distanee to posterior margin. Lateral surfaee of propodeum minutely roughened and

shagreened. T1 smooth and nearly impunetate on dise, apieal margin finely punetate (i = 1.5d). T2

punetate on basis and apieal margin, not in the middle. T3-T4 with very sparse punetures.

Sculpture. Head nearly as long as wide (length/width ratio = 0.89). Eyes slightly eonvergent below

(UOD/LOD ratio = 1.09). Vertex short. Gena as wide as eye. Inner metatibial spur with five short teeth.

Metapostnotum moderately long (MMR = 1.48).

Propodeum. It is with very weak lateral and oblique earina eurved to the middle of the posterior surfaee.

Additional material

The following speeimens are identified as Capalictus, but not deseribed as new beeause only a single sex

is known. They differ slightly from the speeies above by the punetation of mesoseutum and metasomal

terga. The information is provided to eneourage and faeilitate future study of the subgenus.

Lasioglossum {Capalictus) sp. 1

Material

1 S, Farm Papkuilsfontein, 20 km S Nieuwoudtville, Fynbos, 31°33’16''S 19°08’3L'E, 680m, 15 Aug.

2010, leg. M. Kuhlmann

This male speeimen is elose to L. timmermanni sp. nov., but differs by foretibia orange on inner side and

impressed apieal margin of T1 punetate. The female is unknown.

Lasioglossum {Capalictus) sp. 2

Material

All 5?: 2 5? (CUIC) (metasoma blaek), SOUTH AFRICA, Western Cape Provinee (WCP), Kunje

Farm, 760 m, 28 km SSE Citrusdal, 32°40.35’S 19°12.67’E, 23-25 Sep. 2001; 2 ? ? (CUIC) (metasoma

red), WCP, Kunje Farm, 760 m, 28 km SSE Citrusdal, 32°40.35’S 19°12.67’E, 23-25 Sep. 2001; 4

(CUIC) (2 females Voueher 01-54) (metasoma blaek), WCP, 21 km N Hermanus, 157 m, 34° 17.82’S

19°07.71’E, 28 Sep. 2001; 1 $ (Voueher 01-45) (CUIC) (metasoma red), WCP, Cape Agulhas,

34°49.92’S 20°00.14’E, 27 Sep. 2001; 1 $ (CUIC) (metasoma blaek), WCP, Betty’s Bay, botanieal

garden, 27 m, 34°21.14’S 18°55.62’E, 28 Sep. 2001.

Speeimens from Hermanus (4 $ $ ineluding Voueher 01-54) are similar in size and eolour to L. tigrinum

sp. nov., but their tergum 1 is nearly impunetate on the dise, punetures of the mesoseutum are larger and

sparser, punetures on the faee are stronger.

Speeimens with blaek metasoma from Kunje Farm (2 $ $) and Betty’s Bay (1 $) are similar in size and

eolour to L. hantamense sp. nov. but punetures of the mesoseutum are stronger.

The speeimen from Cape Agulhas (1 $, Voueher 01-45) is similar to L. timmermanni sp. nov. by its

eolouration (metasoma red, ineluding last terga) and shagreened propodeum.

Speeimens from Kunje Farm with red metasoma (2 $ $) are similar to L. timmermanni sp. nov. by red

metasoma and shagreened propodeum, but differs by blaek last terga. They are smaller in size than the

speeimen from Cape Agulhas.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Fig. 10. Dorsal habitus. — A-B. Lasioglossum mosselinum (Cockerell, 1945). A. $. B. ( 5 '. — C-D. Lasio-

glossum timmermanni sp. nov. C. $. D. S- Scale line = 1 mm.

European Journal of Taxonomy 28: 1-28 (2012)

Fig. 11. Dorsal habitus. — A-B. Lasioglossum (Capalictus) hantamense sp. nov.. A. $. B. S-

— C-D. Lasioglossum (Capalictus) tigrinum sp. nov. C. $. D. S- Scale line = 1 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Fig. 12. Lateral habitus, S- — A. Lasioglossum (Capalictus) mosselinum (Cockerell, 1945).

— B. Lasioglossum (Capalictus) timmermanni sp. nov. — C. Lasioglossum (Capalictus) hantamensQ

sp. nov.. — D. Lasioglossum (Capalictus) tigrinum sp. nov. Scale line = 1 mm.

Fig. 13. S foretarsi. — A. Lasioglossum (Capalictus) mosselinum (Cockerell, 1945). — B. Lasio¬

glossum (Capalictus) timmermanni sp. nov. — C. Lasioglossum (Capalictus) hantamense sp. nov.

— D. Lasioglossum (Capalictus) tigrinum sp. nov. Scale line = 0,5 mm.

European Journal of Taxonomy 2^\ 1-28 (2012)

Key to species

Males

1. Tibiae of middle and posterior legs yellowish orange (exeept for a eentral dark maeulation)

(Figs 12D, 13D, 14G-H); apieal impressed area of terga largely amber translueent (Fig. 6D)

. L. {Capalictus) tigrinum sp. nov.

- Tibiae of middle and posterior legs dark; apieal impressed area of terga blaek, rarely amber, apieal

rim slightly straw-eoloured . (2)

2. All tarsi pale yellow (Figs 12C, 13C, 14E-F); gonostylus with short setae, length less than % basal

gonostylus width (Fig. 15C). L {Capalictus) hantamense sp. nov.

- Tarsi of middle and posterior legs dark (Fig. 14A-D); gonostylus with long setae, length subequal to

or longer than basal gonostylus width (Fig. 15A-B).(3)

3. Foretarsi blaek (Fig. 13B); punetation of mesoseutum denser (i = 1-1.5d) (Fig. 8B); dorso-lateral

slope of propodeum with mierotesselate dull surfaee (Fig. 8e).

. L. {Capalictus) timmermanni sp. nov.

- Foretarsi yellowish (Fig. 13A); punetation of mesoseutum sparser (i = 3^d) (Fig. 3B); dorso-lateral

slope of propodeum shining (Fig. 3C) . L. {Capalictus) mosselinum (Coekerell, 1945)

Fig. 14. S hind legs (left) and mid legs (right). — A-B. Lasioglossum {Capalictus) mosselinum (Coeke¬

rell, 1945). — C-D. Lasioglossum {Capalictus) timmermanni sp. nov. — E-F. Lasioglossum {Capalictus)

hantamense sp. nov. — G-H. Lasioglossum {Capalictus) tigrinum sp. nov. Seale line = 0,5 mm.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Females

1. Metasoma largely red (Fig. lOA, C).(2)

- Metasoma black, apical margins pale (Fig. 11 A, C).(3)

2. Terga 5-6 largely red (Fig. 16F); dorso-lateral slopes of propodeum shagreened (Fig. 9C, D);

ridges of metapostnotum short and anastomosing (Fig. 9C, D; also Fig. 7 in Pauly et al. 2008)

. L. (Capalictus) timmermanni sp. nov.

- Terga 5-6 blackish (Fig. 16E); dorso-lateral slopes of propodeum shining (Fig. 2D); ridges

of metapostnotum more linear (Fig. 2D, also Fig. 6 in Pauly et al. 2008).

. L. (Capalictus) mosselinum (Cockerell, 1945)

3. Smaller body size (length 6-6.5 mm); T1 nearly impunctate on disc, punctures very weak and

superficial, apical margin with fine punctures (Fig. 5D, E). L. (Capalictus) hantamense sp. nov.

- Earger body size (length 7 mm); T1 with distinct punctuation on disc as well as on apical margin

(Fig. 7D, E). L. (Capalictus) tigrinum sp. nov.

Fig. 15. 3 genitalia, dorsal view. — A. Lasioglossum (Capalictus) mosselinum (Cockerell, 1945).

— B. Lasioglossum (Capalictus) timmermanni sp. nov. — C. Lasioglossum (Capalictus) hantamense

sp. nov. — D. Lasioglossum (Capalictus) tigrinum sp. nov. Scale line = 0,25 mm.

European Journal of Taxonomy 2^\ 1-28 (2012)

Fig. 16. A-D. Lateral view of mesosoma. E-F. Last terga. — A. Lasioglossum {Capalictus)

mosselinum (Coekerell, 1945). —B. Lasioglossum (Capalictus) timmermanni sp. nov. — C. Lasioglossum

(Capalictus) hantamense sp. nov. — D. Lasioglossum (Capalictus) tigrinum sp. nov. — E. Lasioglossum

(Capalictus) mosselinum (Coekerell, 1945). — F. Lasioglossum (Capalictus) timmermanni sp. nov.

Seale line = 0,5 mm .

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Fig. 17. Distribution maps of Lasioglossum {Capalictus) spp. in southern Africa. — A. Lasioglossum

{Capalictus) mosselinum (Cockerell, 1945). — B. Lasioglossum {Capalictus) hantamense sp. nov.

— C. Lasioglossum {Capalictus) tigrinum sp. nov. — D. Lasioglossum {Capalictus) timmermanni

sp. nov. — E. Unidentified specimens oiLasioglossum {Capalictus), $ $.

European Journal of Taxonomy 2^\ 1-28 (2012)

Molecular results

DNA sequence data from five specimens of Lasioglossum (Capalictus) (1 specimen of L. hantamense

sp. nov., 2 ofL. tigrinum sp. nov., and 2 ofL. afif. tigrinum (n°01-54) includes putatively fixed characters

suggestive of species-level differentiation. A total of 1934 bp was sequenced for all five specimens

(EFl-a: 951 bp, wnt-1: 457 bp, and opsin: 525 bp). Lasioglossum hantamense sp. nov. differed from

L. tigrinum sp. nov. by 7 nucleotide substitutions (3 in EF-la, 1 in wnt-1, and 3 in opsin) and from

L. aff tigrinum by 9 nucleotide substitutions (5 in EF-la and 4 in opsin). Lasioglossum tigrinum

sp. nov. differed from L. aff tigrinum by 10 nucleotide substitutions (4 in EF-la, 1 in wnt-1, and 5 in

opsin). Only one variable nucleotide position was found within L. aff tigrinum (in EF-la) and 2 were

found in L. tigrinum sp. nov. (1 each in EF-la and opsin). The levels of genetic differentiation in EF-la

are consistent with those discovered between other closely related bee species (Danforth et al. 1999;

Kuhlmann et al. 2007).

Discussion

Phylogenetic studies of Lasioglossum are ongoing, but recent molecular study clearly supports the

position of Capalictus as a basal lineage of the Lasioglossum {Hemihalictus series) (Gibbs

et al. 2012) (Fig. 18). Nodal support values for the relevant nodes of the topology are very high (100)

when the data are analysed with either Bayesian methods (measured in posterior probabilities) (Gibbs

et al. 2012) or parsimony (measured in GC values; Goloboff et al. 2003) (J. Gibbs unpublished results).

The subgenus Capalictus seems to be largely restricted to the winter-rainfall region of South Africa that

is known as a centre of bee endemism and species diversity of global importance (Kuhlmann 2009).

In this region some other bee genera are known that represent basal lineages within their families,

subfamilies or tribes. These are Fidelia, Fideliopsis, Afroheriades, Aspidosmia (Megachilidae) (Fitman

et al. 2011), and Haplomelitta (Melittidae) (Michez et al. 2009). Additional research in this centre of

bee and fioral diversity is of special interest for the understanding of bee phylogeny and evolution

(Kuhlmann 2009).

100

Lasioglossum series

Capalictus

DIallctus

Evylaeus

Hemihalictus

Sellalictus

Sphecodogastra

Sudila

Fig. 18. Summary of phylogenetic relationships of Capalictus relative to remaining Lasioglossum,

based on Gibbs et al. 2012. Numbers represent posterior probability support values.

PAULY A., GIBBS J. & KUHLMANN M., Capalictus subgen. nov. from South Africa

Acknowledgements

MK is much indebted to Onno Huyser, Manager of the Table Mountain Fund, and Noel Oettle, Managing

Director of Avontuur Sustainable Agriculture, for giving him access to the farm Avontuur and their

permission to study the bees on the property. Northern Cape Nature Conservation Service is gratefully

acknowledged for giving their permission to collect bees. Basie Nel of the farm Renosterhoek has kindly

given MK access to the plateau of the Hantam Mountain and Hergen Erhardt, Edewecht (Germany),

supported MK by collecting bees in the wider Nieuwoudtville area at various occasions.

Molecular sequencing was supported by NSF grants in systematic biology (DEB-0814544, DEB-

0742998) to Bryan Danforth (Cornell University).

References

Cockerell T.D.A. 1945. Descriptions and records of bees. CXCVL Annals and Magazine of Natural

History 11), 12: 273-282.

Danforth B.N. 1999. Phylogeny of the bee genus Lasioglossum (Hymenoptera: Halictidae) based on

mitochondrial COI sequence data. Systematic Entomology 24:377-393. http://dx.doi.org/10.1046/

i.1365-3113.1999.00087.x

Danforth B.N., Conway E. & Ji S.Q. 2003. Phylogeny of eusocial Lasioglossum reveals multiple losses

of eusociality within a primitively eusocial clade of bees (Hymenoptera: Halictidae). Systematic Biology

52: 23-36. http://dx.doi.org/10.1080/10635150390132687

Danforth B.N., Brady S.G., Sipes S.D. & Pearson A. 2004. Single copy nuclear genes recover Cretaceous

age divergences in bees. Systematic Biology 53: 309-326. http://dx.doi.org/10.1080/10635150490423737

Engel MS. 2001. A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera).

Bulletin of the American Museum of Natural History 259: 1-192.

Gibbs J. 2011. Revision of the xxiQt?i\\\c Lasioglossum (Dialictus) ofEastem North America (Hymenoptera:

Halictidae: Halictini). Zootaxa 3073: 1-216.

Gibbs J., Brady S.G., Kanda K. & Danforth B.N. 2012. Phylogeny of halictine bees supports a shared

origin of eusociality for Halictus and Lasioglossum (Apoidae: Anthophila: Halictidae). Molecular

Phylogenetics and Evolution 65: 926-939. http://dx.doi.Org/10.1016/i.vmpev.2012.08.013

Goloboff P.A., Farris J.S., Kallersjo M., Oxelman B., Ramirez M.J. & Szumi C.A. 2003.

Improvements to resampling measures of group support. Cladistics 19: 324-332. http://dx.doi.

org/10.1111/i. 1096-0031.2003 .tb00376.x

Harris R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology 28: 1-31.

Kuhlmann M. 2009. Patterns of diversity, endemism and distribution of bees (Insecta: Hymenoptera:

Anthophila) in southern Africa. South African Journal of Botany 75: 726-738.

Kuhlmann M., Else G.R., Dawson A., & Quicke D.E.J. 2007. Molecular, biogeographical and pheno-

logical evidence for the existence of three western European sibling species in the Colletes succinctus

group (Hymenoptera: Apidae). Organisms, Diversity and Evolution 7: 155-165. http://dx.doi.

org/10.1016/i.ode.2006.04.001

Eitman J.R, Danforth B.N., Eardley C.D. & Praz C.J. 2011. Why do leafcutter bees cut leaves? New

insights into the early evolution of bees. Proceedings of the Royal Society B 278: 3593-3600. http://

dx.doi.org/10.1098/rspb.2011.0365

Michener C.D. 2007. The Bees of the World [2nd Edition]. Johns Hopkins University Press, Baltimore,

Maryland.

European Journal of Taxonomy 2^\ 1-28 ( 2012 )

Michez D., Patiny S. & Danforth B.N. 2009. Phylogeny of the bee family Melittidae (Hymenoptera:

Anthophila) based on eombined moleeular and morphologieal data. Systematic Entomology 34: 574-

597. http://dx.doi.Org/10.llll/i.1365-3113.2009.00479.x

Pauly A. 1999. Classifieation des Halietini de la Region Alfotropieale (Hymenoptera ApoideaHalietidae).

Bulletin de ITnstitut royal des Sciences naturelles de Belgique, Entomologie 69: 137-196.

Pauly A., Brooks R.W., Nilsson L.A., Pesenko Y.A., Eardley C.D., Terzo M., Griswold T., Sehwarz M.,

Patiny S., Munzinger J. & Barbier Y, 2001. Hymenoptera Apoidea de Madagascar et des lies voisines.

Annales Seienees zoologiques 286, Musee royal de PAfrique eentrale, Tervuren.

Pauly A., Timmermann K. & Kuhlmann M. 2008. Deseription of a new interesting speeies from

South Afriea, Evylaeus (Sellalictus) Jynbosensis n.sp. (Hymenoptera Apoidea Halietidae). Journal of

Afrotropical Zoology 4: 85-91.

T imm erma nn K. & Kuhlmann M. 2009. Taxonomie revision of the Afriean bee subgenera Patellapis,

Chaetalictus md Lomatalictus (Hymenoptera: Halietidae, genus Patellapis Friese 1909). Zootaxa 2099:

1-188.

Manuscript received: 29 May 2012

Manuscript accepted: 29 October 2012

Published on: 12 November 2012

Topic editor: Koen Martens

In eomplianee with the ICZN, printed versions of all papers are deposited in the libraries of the institutes

that are members of the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National

Botanie Garden of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium;

Natural History Museum, London, United Kingdom; Royal Belgian Institute of Natural Seienees,

Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.