Tijdschrift voor Entomologie

A journal of systematic and evolutionary entomology since 1858

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E. J. van Nieukerken (elected 1986) and J. van Tol (1985)

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Published with index of volume 137 (1994).

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Ontwerpers B.V., Aad Derwort, ’s-Gravenhage

A. J. DE BOER

MCZ

FIBRARY

Institute for Systematics and Population Biology Zon Museum), University of Amsterdam

THE TAXONOMY AND BIOGEOGRAPHY RRVAE

CICADA GENUS PAPUAPSALTRIA GEN; Ni VERSITY

(HOMOPTERA, TIBICINIDAE)

fi )

Boer, A.J. de, 1995. The taxonomy and biogeography of the cicada genus Papuapsaltria gen. n.

(Homoptera, Tibicinidae). — Tijdschrift voor Entomologie 138: 1-44, figs. 1-190, table 1 [ISSN

0040-7496]. Published 15 June 1995.

The genus Papuapsaltria is erected for 18 species, distributed in New Guinea and several near-

by islands. Three species (Thaumastopsaltria nana (Jacobi, 1903), Baeturia phyllophora Blöte,

1960, and Baeturia ustulata Blöte, 1960) are transferred to Papuapsaltria and redescribed, while

15 species (P. angulata, P. baasi, P. bidigitula, P. brassi, P. dioedes, P. dolabrata, P. goniodes, P

lachlani, P. novariae, P. plicata, P. stoliodes, P. spinigera, P. toxopei, P. ungula, and P. woodlar-

kensis) are described as new. Baeturia famulus Myers is brought into the synonymy of P. nana

(Jacobi). The phylogeny of Papuapsaltria is discussed and some remarks are made on its phylo-

genetic relationships with some other New Guinean and Australian genera. A very short prox-

imal spine on the fore femur is regarded an autapomorphy for the genus. Several monophylet-

ic subgroups are recognised. Two species (P. woodlarkensis and P. phyllophora), missing the

apomorphy of the genus and not clearly belonging to any of the subgroups, are preliminarily in-

cluded. A key to the males and maps of distribution are presented.

A. J. de Boer, Instituut voor Systematiek en Populatie Biologie (Zoölogisch Museum),

University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands.

Key words. — Papuapsaltria, taxonomy, new genus, new species, biogeography, New Guinea.

The great majority of the tibicinid cicadas of New

Guinea belong to the ‘Baeturia and related genera

complex’ as defined earlier (De Boer 1990) by an S-

curved aedeagus with winged lateral crests. The genus

Papuapsaltria is erected for 18 species, clearly belong-

ing to that complex, that could not comfortably be

accommodated in any of the existing genera. The spe-

cies of Papuapsaltria are supposed to be closely relat-

ed, which is corroborated by the distributions of a

number of presumed apomorphous characters. Sev-

eral monophyletic subgroups can be recognized,

based upon apparently sound apomorphies. An un-

ambiguous apomorphy for Papuapsaltria as a whole

could not be found, however. Nevertheless, one char-

acter, a very short proximal spine on the fore femur, is

shared by most of its species. Furthermore, this char-

acter is unique for Papuapsaltria.

The phylogenetic position of P. phyllophora (Blöte,

1960) and P. woodlarkensis sp. n. is very uncertain.

These species are included in the genus, since they are

more similar to the species of Papuapsaltria than to

any other species of the ‘Baeturia and related genera

complex’.

The phylogenetic relationships of Papuapsaltria are

discussed; similarities in male operculum suggest a

possible sister group relationship with the genus

Guineapsaltria De Boer.

Papuapsaltria is distributed in New Guinea, in-

cluding the nearby islands of Yapen, Normanby, and

Waigeu, and has an endemic species on Woodlark is-

land. One species, otherwise found in western New

Guinea and on Yapen Island, is also recorded from

Wetar island near Timor, but not known from any

other islands of the Banda area. Papuapsaltria has

most of its species in Papua New Guinea and seems to

be absent from Cendrawasih and the most southern

parts of central New Guinea.

MATERIAL AND METHODS

The material examined for this study is deposited

in the following collections:

AMNH American Museum of Natural History, New

York

AMSA Australian Museum, Sydney

BMNH Natural History Museum [formerly: British

Museum (Natural History)], London

BPBM _ Bernice P. Bishop Museum, Honolulu

CAS California Academy of Sciences, San Francis-

Institut Royale des Sciences Naturelles de

Belgique, Bruxelles

ISNB

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Baeturia

Gymnotympana

Scottotympana

Venustria

Chlorocysta

Glaucopsaltria

Owra

Guineapsaltria

Papuapsaltria

one Aedeastria

5 Mirabilopsaltria

Thaumastopsaltria

Cystopsaltria

‚Cystosoma

Prasiini

Fig. 1. Phylogenetic relationships of Papuapsaltria. Num-

bers refer to characters discussed in the text.

MCSN Museo Civico di Storia Naturale ‘G. Doria’,

Genova

Moul Personal collection Mr M.S. Moulds, Sydney

MVMA Museum of Victoria, Melbourne

NCSU North Carolina State University Insect Col-

lection, Raleigh

NMWC National Museum of Wales, Cardiff

RMNH Nationaal Natuurhistorisch Museum (for-

merly: Rijksmuseum van Natuurlijke Histo-

rie), Leiden

SMFD Natur Museum und Forschungs Institut

‘Senkenberg Frankfurt am Main

SMNS Staatliches Museum für Naturkunde, Stutt-

gart

SMTD Staatliches Museum für Tierkunde, Dresden

TMB _ Természettudomány Múzeum, Budapest

ZILS Zoologiska Institutionen Zoologiska Museet,

Lund

Instituut voor Systematiek en Populatie Bio-

logie (Zoölogisch Museum), Amsterdam

ZMAN

The following sources have been used for tracing

localities: ‘Atlas van tropisch Nederland’ (1938), “The

Times Atlas of the World Comprehensive Edition’

(1968), ‘Papua New Guinea a travel survival kit

(Wheeler 1988) and a ‘List of New Guinea localities’

published by the Bernice P. Bishop Museum (1966).

To examine the male genitalia, the pygofer was

pulled out, after overnight softening, with a sharp

needle inserted between the pygofer and the 8th ab-

dominal segment. The aedeagus was pulled out at the

same time, by inserting the needle between the clasp-

ers. Some of the terms used in the descriptions are ex-

plained in figs. 12 and 17. Body and tegmen lengths of

all specimens were measured, other measurements are

based on a maximum of ten specimens, when avail-

able.

PHYLOGENY

Papuapsaltria is a fairly heterogeneous genus, which

forms part of a larger monophyletic group, defined ear-

lier as the “Baeturia and related genera complex’ (De

Boer 1990). A phylogenetic analysis of this complex,

using the computer program PAUP (Swofford 1993) is

in preparation. The oriental Prasiini as defined by De

Jong (1985), which form the most likely sister group of

that complex, will be included as outgroup in this ana-

lysis. Preliminary results of the computer analysis indi-

cate that the species of Papuapsaltria are closely related,

that the genus is probably monophyletic and that it

forms the sister group (or groups) of the monophyletic

genus Guineapsaltria (De Boer 1993a).

The cladograms presented here, showing the phy-

logenetic position of Papuapsaltria within the

‘Baeturia and related genera complex’ (fig. 1) and a

tentative reconstruction of the relationships of the

species of the genus (fig. 2), comply to a large extent

with the afore mentioned PAUP analysis. Minor differ-

ences are due to the influence of several strongly ho-

moplasious characters, as a result of which the most

parsimonious trees do not recognize some of the ob-

vious synapomorphies as such.

The phylogenetic position of Papuapsaltria

The presumed apomorphous characters indicated

angulata

6 goniodes

4 5 brassi

7 lachlani

plicata -

Zi 11 1213 stoliodes

9 10 nana

14 ungula

8 -bidigitula

spinigera

baasi

dolabrata

15 novariae

ustulata

dioedes

toxopei

.phyllophora

woodlarkensis

Fig. 2. Phylogenetic reconstruction of Papuapsaltria. Num-

bers refer to characters discussed in the text.

by numbers in the cladogram of fig. 1 are discussed in

short. Many of these characters have been discussed

earlier (De Boer 1990; 1991; 1992b; 1993a).

1. — Aedeagus S-curved and provided with wing-

shaped lateral crests. This is the supposed apomorphy

of the ‘ Beaturia and related genera complex’ (De Boer

1990). The Prasiini have a C-curved aedeagus with-

out lateral crests, which is regarded as plesiomor-

phous.

2. — Surfaces of head and pronotum fairly smooth;

pronotum generally without a distinct medial furrow.

This could be two separate characters. Other genera

of the complex and the Prasiini have a more wrinkled

head and pronotum, while the pronotum generally

has a distinct medial furrow, which is regarded as ple-

siomorphous. This character is not absolutely dis-

criminating; anomalies sporadically occur in both

subgroups.

3. — A fairly broad hyaline border along the hind mar-

gin of the tegmen. Other genera of the complex and

the Prasiini have a very narrow border along the hind

margin of the tegmen, which is regarded as plesio-

morphous. This character too, is not absolutely dis-

criminating. Furthermore, in Baeturia Stal, 1866, this

border is quite narrow again, somewhat intermediate

relative to the plesiomorphous state.

4. — A large male operculum; the medial margin of

the operculum lies medially of the meracanthus. The

phylogenetic relationships between the four genera

that have this character are discussed in more detail

elsewhere (De Boer 1995a). Other genera of the com-

plex and the Prasiini have a generally smaller male op-

erculum, of which the medial margin lies laterally of

the meracanthus. Such small opercula are regarded as

plesiomorphous. The larger opercula of P. plicata and

P. stoliodes (see below) form two exceptions (see figs

59 & 73) which must be explained by paralellism.

5. — Tegmina with more than 8 apical areas and a

more or less continuous band of subapical areas (see

plate 22 figs. 2, 3, 6, and 7 in Moulds 1990).

6. — A very small male operculum with a rounded dis-

tal part. The distal part is often shorter than the basal

part of the operculum. Though this character is not

found in all species, it might indicate a sister group re-

lationship between Papuapsaltria and Guineapsaltria.

Such a relationship is in compliance with the most

parsimonious computer analyses.

7. — A narrow tymbal cavity. This character is not

present in all species of the group it is supposed to de-

fine.

8. — The absence of a distinct ridge along the tergite

margin bordering the tymbal cavity. This character is

also not present in all species of the group it is sup-

posed to define.

9. — A very long first apical area of the tegmen (see

figs. 4-5 De Boer 1992b). This character is shared by

DE BOER: The genus Papuapsaltria

most species of Thaumastopsaltria Kirkaldy, 1900,

and some species of Mirabilopsaltria De Boer, 1995,

and is supposed to be synapomorphous for these two

genera together, or for Thaumastopsaltria and a part

of Mirabilopsaltria. In the latter case Mirabilopsaltria

cannot be regarded as monophyletic (see De Boer

1995b).

10. — Apically pointed tegmina.

11. — Tegmina with reticulate venation (see plate 23

figs. 1-3 Moulds 1990).

The phylogenetic position of Aedeastria is ambigu-

ous, as expressed by the question marks in fig. 1.

Aedeastria, Mirabilopsaltria, and Thaumastopsaltria

share a very similarly shaped angular male operculum

and generally very distinct diverging fissures on the

vertex. The distinct diverging fissures are possibly

synapomorphous for these three genera together.

However, nearly all species of Mirabilopsaltria,

Cystosoma, Cystopsaltria, and Thaumastopsaltria share

a strongly bent proximal spine on the fore femur, a

character that is also widely distributed in the Prasiini

and presumably plesiomorphous. This spine is gener-

ally more erect in Aedeastria and other genera of the

complex, which could be a synapomorphy.

Ingroup phylogeny

A tentative cladogram of Papuapsaltria is given in

fig. 2. Only characters that are supposed to be apo-

morphous are discussed here; these are indicated by

numbers in the cladogram. The cladogram presented

here differs in details from the most parsimonious re-

constructions currently calculated by PAUP; differen-

ces are caused by a more parsimonious distribution of

homoplasious characters. Such homoplasious charac-

ters, in which the species of Papuapsaltria differ from

each other, and which are mentioned in the species

descriptions, (e.g., postclypeus shape, number of

tymbal ridges, size of auditory capsules, and presence

or absence of apical lobes on the aedeagus) are not

discussed, since they are presumably of a limited phy-

logenetic importance. These characters will be inclu-

ded in a final data matrix, which forms the basis for

the current computer analysis. The most parsimo-

nious solutions will be published later.

1.- A reduced, knobby, proximal spine on the fore

femur (figs. 8-9). This character is regarded as a prob-

able apomorphy for the genus Papuapsaltria as a

whole, though possibly excluding P. phyllophora and

P. woodlarkensis (see remark below). However, even

when these two species are excluded from the genus,

this character is not found in all species. P. bidigitula,

P. nana, and to a lesser extent P. baasi, P. dolabrata,

and P. ustulata, have a fairly long and generally point-

ed proximal spine on the fore femur. In these latter

species the proximal spine is always distinctly shorter

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

ere Zu en ; ; ; i Fig. 3. Localities of Papua-

RES i ebidigitula || psaltria bidigitula, P. nana,

6 nana P. plicata, P. spinigera, P.

D à 40) @plicata stoliodes, and P. ungula.

6 A spinigera

FAR sstoliodes © x

“a N vungula FAC

BN, ce

than the distance to the middle spine, however, and it

possibly represents just another state of the very same

character. Only P. phyllophora and P. woodlarkensis

have a distinctly longer spine; longer than the distance

to the middle spine. This latter character is also found

in related genera, and therefore presumably plesiomor-

phous. A reduced spine is unique for Papuapsaltria.

2.— A strongly convex distal margin of the pygofer,

forming a sometimes weakly incurved distal lobe. The

character is less clear in P. angulata and considered

lost in P. bidigitula and P. nana. A similar, but small-

er and more distinctly incurved, lobe was described

for Scottotympana biardae De Boer, 1991 and S.

huibregtsae De Boer, 1991, which must be explained

by parallelism. The pygofer of P. phyllophora also has

a distinctly convex, though much longer, distal mar-

gin, and it is not clear whether this fact indicates a

close relationship between P. phyllophora and the

group of 8 species defined by the here discussed apo-

morphy. The distal margin of the pygofer is only

weakly convex the remaining species of

Papuapsaltria and in most of the related genera,

which is supposedly the plesiomorphous state.

3. — A squarish clasper, without distinct dorsal

crest, but with an angular distodorsal corner. This

character is not recognized in P. nana and P. ungula,

presumably caused by the development of a globular-

ly rounded dorsodistal protrusion on the clasper

(character 14).

4, — A distinct and inwards curving rectangular

corner below the lateral protuberance of the pygofer.

Due to this incurving corner the ventral part of the

pygofer opening is somewhat horseshoe-shaped, with

concave margins (fig. 22). The character is unique for

four species, but not found in all specimens of one of

these four (P. angulata).

r — : a L- I = === = — == I

abc mdioedes

RSS e phyllophora no

- Sr MOR v toxopei

Len, >: x ae ||

E nc XE

TT dA v = N 4

Di CSI ) ° Q x a Po - Hu

B n Woon 7 4

L 4 B 0° N \

SP È UU] n °

er ©

Ha off { N © © ge

a LA ~ x o ant En |

DS da ze. ne N Fig. 4. Localities of Papua-

ae È, na psaltria dioedes, P. phyllo-

iL 4 on, ee ne phora, P. toxopei, and P.

= I LE

1322 136° 140°

woodlarkensis.

5. — A square-shaped laminiform clasper, with a

single, sharply pointed and outcurving, distoventral

protrusion. The character is unique for four species.

6. — A nearly triangular distal part of the male op-

erculum, with a very long and straight medial margin

(figs. 17, 29). This operculum shape presumably in-

dicates a sister group relationship between P. angula-

ta and P. goniodes. Similarly shaped opercula occur in

P. dioedes (fig. 163), Guineapsaltria chinai (Blöte,

1960), Chlorocysta vitripennis Westwood, 1851 and

Cystosoma saundersii Westwood, 1842, which must

be explained by parallelisms.

7. — A very long and stout aedeagus. Several other

species of this genus (P. angulata, P. baasi, P. dolabra-

ta, P. novariae, and P. spinigera) have a fairly long ae-

deagus as well, which stands erect between the clasp-

ers and reaches with its curved apical part well beyond

the pygofer and anal valves. Similarly exposed aedea-

gi were also found in several species of

Gymnotympana. The aedeagi of P. brassi and P. lach-

lani are extremely long, however, and presumably in-

dicate the monophyletic origin of these two species.

8.- A thorn-shaped lateral protrusion on the clasp-

er. The bluntly rounded lateral swelling in the clasper

of P. nana is supposed to be homologous with this

protrusion, while a slight swelling on the clasper of P.

woodlarkensis might also be homologous, and thus in-

dicate a relationship of that species to the species with

this apomorphy. The more lobate lateral protrusions

described for the claspers of some species of the genus

Thaumastopsaltria (see De Boer 1992b) are consid-

ered to be non-homologous.

9. — A flattened and distinctly posteriorly project-

ing protuberance on the lateral lobes of the pygofer.

The character is presumed to be lost in P. nana,

though that species still has a fairly stout protu-

berance, compared to other species of the genus.

10. — The medial part of the clasper forming an al-

most transparent membrane, arching from the apex

to the base of the clasper. This character is presumed

lost by a reversal in P. plicata.

11.—A fairly large male operculum, with the medi-

al margin of its distal part reaching medially of the

meracanthus. Opercula that extend to medially of the

meracanthus are synapomorphous for the genera

Baeturia, Gymnotympana, Scottotympana, and

Venustria (see above), but in these genera the distal

part of the operculum is generally larger and of a dif-

ferent shape than in the two species of Papuapsaltria

which share this apomorphy. Similarities with the

four genera mentioned above must be explained by

parallelism.

12. — A very stout and bicuspidate caudodorsal

beak, separated from the pygofer by a distinct fold at

its base (figs. 58, 69). A bicuspidate, though differ-

ently shaped, caudodorsal beak was found in two spe-

DE BOER: The genus Papuapsaltria

cies groups of Baeturia: the B. nasuta group and the

B. viridis group (De Boer 1982; 1992a), which must

be explained by parallelism or convergence. The fold

at the base of the beak is unique for these two species

of Papuapsaltria.

13. — An aedeagus with a pair of distinct and angu-

lar dorsal ridges. Less well-developed dorsal ridges, ei-

ther single or paired, occur in many other species of

Papuapsaltria and in many related genera.

14. — A globularly rounded dorsodistal protrusion

on the clasper (figs. 77, 89). A similar protrusion was

found in Aedeastria hastulata De Boer, 1993, and sev-

eral species of Gymnotympana, which must be ex-

plained by parallelisms.

15. Claspers of which the dorsal crests closely en-

circle the aedeagus as a short shaft. The claspers of the

four species with this character are very similar in

other respects, mainly differing in the angle between

their apical and basal parts. The character is unique

for these four species.

Remark: The phylogenetic positions of P. phyllo-

phora and P. woodlarkensis are very uncertain. The

shape of the head and the operculum of these species

suggest that they either belong to Papuapsaltria or to

Guineapsaltria. In spite of the fact that these two spe-

cies have a fairly long proximal spine on the fore fe-

mur, they are preliminary included in Papuapsaltria,

since their ocelli are more closely together than is

common in Guineapsaltria and they do not share the

distinct subapical lobe on the aedeagus, which is re-

garded synapomorphous for that genus (De Boer

1993a). The inclusion of these two species in

Papuapsaltria is supported by the preliminary results

of the computer analysis; in several of the most parsi-

monious reconstructions these species appear at vari-

ous places between the other species of the genus.

Discussion of other characters of possible

phylogenetic relevance

Several other characters, not included in the phylo-

genetic discussion, are discussed below, since they are

supposed to be indicative of a close relationship be-

tween species of Papuapsaltria:

Cicadas generally have eight apical areas in the teg-

men, but several species of Papuapsaltria have a differ-

ent number of apical areas, a number which often

even varies between the right and left tegmen of indi-

viduals. P. novariae has 9 apical areas, this number

varies between 8-9 in P. nana, between 8-10 in P.

lachlani and P. plicata, and between 9-12 in P. gonr

odes. P. stoliodes has generally 8-9 apical areas in the

tegmen, but 11-12 in one specimen, and one speci-

men of P. baasi has 7 apical areas in the right tegmen.

Such variations only sporadically occur in related

genera and the fact that it occurs relatively often in

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 5-11. — 5, head in dorsal view P. goniodes, holotype; 6, head in dorsal view P. nana, Popondetta; 7, female genital seg-

ment in lateral view P. goniodes, Daulo; 8, fore femur P. goniodes, holotype; 9, detail fore femur P. goniodes, holotype; 10, fore

femur P. woodlarkensis, holotype; 11, head in lateral view P. goniodes, holotype.

Papuapsaltria might indicate a close relationship

between its species.

Tegmina with a more strongly variable number of ap-

ical areas (10-15), are characteristic for Thaumasto-

psaltria (De Boer 1992b), a monophyletic group of four

Gymnotympana species (De Boer 1995a), a presumed

monophyletic group consisting of Chlorocysta, Glauco-

psaltria, and Owra (for figs. see Moulds 1990), and for

two species of Mirabilopsaltria (De Boer 1995b).

Apical and ulnar areas are generally long and slen-

der in Papuapsaltria (figs. 21, 65), but distinctly

shorter in P. baasi, P. dolabrata, P. novariae, P. phyl-

lophora and P. ustulata (fig. 137). The tegmina of

these five species resemble those of Guineapsaltria.

P. dioedes and P. woodlarkensis have very similar

opercula, which might indicate a close relationship

between these species; the lateral margin of the distal

part runs almost straight to the distolateral corner of

the basal part (figs. 163, 172).

Apart from P. plicata, P. phyllophora, P. stoliodes, P.

toxopei, P. woodlarkensis, and some specimens of P.

bidigitula the species of Papuapsaltria have a very

slender caudodorsal beak (compare figs. 12 and 53),

often with concave margins in dorsal view (cf. fig.

44). Such an extremely slender caudodorsal beak on-

ly sporadically occurs in related genera.

BIOGEOGRAPHY

Papuapsaltria is widely distributed throughout the

greater part of New Guinea and further recorded

from the islands of Normanby, Roon, Waigeu,

Woodlark, and Yapen (figs. 3-4, 32, 129). The genus

appears to be absent from Cendrawasih (the

Vogelkop peninsula), the southern parts of central

New Guinea, and the central mountain ranges of

western New Guinea. A record of P. bidigitula from

Wetar island is considered doubtful, since that species

is not known from any of the other Banda islands.

The cicada genera that occur on New Guinea are

not evenly distributed over the island. The various

genera are generally concentrated with most, and of-

ten endemic, species in different parts of the island.

Most of these parts, thus recognised as areas of en-

demism, coincide with fragments of a historic volcan-

ic island arc, known as the Outer Melanesian Arc.

Over the last 25 million years or so these fragments

have collided at various times and localitions with the

northern craton of the Australian continent (for geo-

logical information see Pigram & Davies 1987).

These facts suggest that the various New Guinean

genera evolved on different island arc fragments,

prior to the collision of these fragments with

Australia. It was inferred (De Boer 1994; 1995a), that

the genera Diceropyga, Gymnotympana, and Thauma-

stopsaltria probably evolved on the East Papua

Composite terrane, which collided about 15 My ago

with New Guinea and now forms the greater part of

the Papuan peninsula (Pigram & Davies 1987).

Similarly Rhadinopyga and possibly also Aedeastria

might originate from any of two microcontinents

now forming Cendrawasih, these genera concentrate

in western New Guinea (De Boer 1992b; 19936;

1994). The occurrences of such genera outside their

presumed area of origin is supposed to have resulted

from dispersal since the time of collision of these are-

as. Such a hypothesis is corroborated by the fact that

those species that do occur outside the areas of pre-

sumed origin generally have a relatively wide distribu-

tion.

The distribution of the genus Papuapsaltria does

not readily suggest an origin on any of the arc-frag-

ments or microcontinents that collided with

Australia. Certainly, most species of Papuapsaltria are

found in Papua New Guinea. Of its 18 species, eleven

occur in Papua New Guinea, and nine are endemic

there. Five of these endemics are restricted to the

Papuan peninsula. These numbers of endemics seem

to indicate that Papuapsaltria, or at least part of that

genus, also originates from the East Papua Composite

terrane. Several species may have dispersed from the

Papuan peninsula to the mountains of western Papua

New Guinea. It is remarkable that many species of

Papuapsaltria were collected from montane altitudes

(table 1), so that such a dispersal, from one montane

area to another, might have been easily achieved.

However, Papuapsaltria has no less than five en-

demic species in northwestern New Guinea, four of

which presumably form a monophyletic group.

Furthermore, disregarding the record from Wetar, P.

bidigitula could also be an endemic of northwestern

New Guinea. The distribution of these endemics can-

not easily be explained by a dispersal from the Papuan

peninsula and a different area of origin, on another

fragment of the historic island arc, must certainly be

considered for these species. The endemics of north-

DE BOER: The genus Papuapsaltria

Table 1. Altitudinal ranges of the species of Papuapsaltria

P. angulata 700-2070 m

P. baasi 100-750 m

P. bidigitula 0-1350 m

P. brassi 1650-1950 m

P. dioedes 2280-2950 m

P. dolabrata 0-2500 ft

P. goniodes 700-3000 m

P. lachlani 400-1400 m

P. nana 0-150 m

P. novariae 1200-2530 m

P. phyllophora 0-1800 m

P. plicata 2070-2800 m

P. spinigera 700-1550 m

P. stoliodes 1100-2000 m

P. toxopei 1800 m

P. ungula 450 m

P. ustulata 1300-1500 m

P. woodlarkensis 100 m

ern New Guinea, or at least some of them, could orig-

inate either from the Gauttier or Torricelli terranes,

or from both. These terranes formed a, probably

more or less continuous, part of the Outer

Melanesian Arc and collided at about 10 My ago with

New Guinea (Pigram & Davies 1987).

As explained above, the monophyly of Papua-

psaltria is far from certain. The apparent dual origin

of the genus, on different parts of the Outer

Melanesian Arc, is another factor shedding doubt on

its assumed monophyly.

TAXONOMY

Genus Papuapsaltria gen. n.

Type species. — P. angulata sp. n.

Description

The species of Papuapsaltria are mostly fairly small.

Males generally have a body length shorter than 2 cm

and, more often, do not exceed 1.5 cm. Only P. dioe-

des is distinctly larger, with a body length of 20.0-

26.5 mm. Body generally ochraceous to reddish

brown and often with traces of green, but without

special colour markings, though abdomen sometimes

with reddish segmental hind margins. Many of the

species possibly entirely green when alive. Females of

most species, on average, slightly smaller than males

but with more robust head and thorax and slightly

longer tegmina. Tegmina of males 1.0-1.3 X as long

as body length, of females 1.5-1.6 X. Male abdomen

distinctly longer (1.2-2.2 X) than head and thorax, of

females 1.0-1.3 X. Head fairly broad and short (figs.

5-6), only slightly narrower than anterior part of pro-

notum, 2.3-2.9 X as wide as long and 1.2-1.5X as

wide as distance between eyes. Postclypeus 2.0-3.3 X

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

as wide as long, oblong in dorsal view and weakly

protruding, and not swollen ventrally; anterior mar-

gin (lateral view) straight or weakly concave (fig. 11).

Sides of postclypeus fairly smooth, but often with

some weak furrows and with about 6 rows of short

parallel ridges in a narrow band along the lorum.

Vertex fairly broad and smooth, though with distinct

medial fissure. Diverging fissures from center of head

to corners of postclypeus generally weakly developed,

almost obsolete. Vertex 1.5-2.2X as wide as long;

1.3-2.1 X as wide as postclypeus and 1.9-2.5X as

wide as eye. Ocelli small and fairly close together.

Distance between lateral ocelli 1.2-2.6 X the width of

frontal ocellus and about as long as (0.8-1.2X) dis-

tance between lateral ocellus and eye. Pronotum 2.2-

2.9 X as wide as long and smoothly vaulted, without

distinct medial furrow. Pronotal collar laterally angu-

larly amplified, and slightly curving down at its ante-

rior margin. Tegmina and wings hyaline, venation

ochraceous. Tegmina generally with 8, but fairly of-

ten more (9-12) apical areas, though rarely with sub-

apical areas. Tegmen with distinct hyaline costal area

and generally a broad hyaline border along hind mar-

gin. Wings with G apical areas, a distinctly broader

hyaline border, and large anal fields. Legs ochraceous

and unmarked. Fore femur long and slender, with

row of 3 erect spines. Proximal spine in most species

very short and apically rounded, only slightly longer

than middle spine and generally distinctly shorter

than distance to middle spine (figs. 8-9). Tymbal

with 5-9 parallel sclerotized ridges. Short intercalary

ridges can often be discerned, forming a darkened

band at half-width across the tymbal. Opercula gene-

rally very small. Basal part of operculum slightly

vaulted with two rounded elevations, and upcurved at

distolateral corner into distinct crest rounding that

corner. Lateral part of that crest often short and

knobby, distal part generally fairly long and gradually

amplifying into distal part of operculum (cf. fig. 17),

though in some species shorter and abruptly ampli-

fying into distal part (cf. fig. 29). Distal part of oper-

culum angularly oblong, in most species shorter than

basal part. Male operculum not, or only partly, cover-

ing tymbal cavity in ventral view, often leaving

greater part of folded membrane exposed. Its lateral

margin straight and slightly directed mesiad.

Operculum shorter than meracanthus and generally

not extending medially beyond meracanthus. P. pli-

cata and P. stoliodes with distinctly larger opercula.

Female operculum shorter, but generally of same sha-

pe as that of male. Male abdomen very delicate, in the

larger sized species distinctly inflated, in small sized

species not notably inflated and then of almost the

same shape as that of female. First tergite in male fairly

short medially, sometimes hidden under metanotum.

Medial part of second tergite less than 1.5 X as long

as lateral parts, proximal margin of second tergite

weakly convex, sometimes slightly concave medially

(figs. 68, 177) and almost straight between auditory

capsules and sternite 2, forming a distinct ridge along

tymbal cavity (cf. fig. 17). Lateral parts of 2nd tergite

weakly swollen at anterior margins and almost ad-

jacent to tymbals, leaving a narrow gap between tym-

bal and 2nd tergite. First sternite swollen and bluntly

rounded posteriorly. Female abdomen more robust

than that of male, with slender pygofer. Ovipositor

sheaths reaching beyond apex of bluntly rounded

caudodorsal beak. Male pygofer globularly rounded,

convexly rounded dorsally, and often with distinctly

convex distal margins. Caudodorsal beak short and

straight, not curved over basal part of claspers or anal

valves (much larger and strongly bent in P. plicata

and P. stoliodes). Claspers fairly short and directed

downwards, fused, or nearly fused, at base, forming a

ring-shaped collar around base of anal valves. Clasper

generally sharply pointed at apex, and sometimes

with sharply pointed lateral protrusion. Apical part of

clasper generally with slender and sharply edged clas-

per hollow. Aedeagus S-curved with winged lateral

crests, often with 1 or 2 dorsal ridges, and sometimes

with small subapical lobe (aedeagus of P. phyllophora

strongly deviating). Aedeagal pore round.

Gender: Feminine.

Key to the males

1. Clasper distinctly bicuspidate, with sharply poin-

ted apical protrusion and distinct, pointed or

rounded, lateral protrusion. (cf. fig. 56) .......... 2

— Clasper not bicuspidate, with only one pointed

and, generally apical protrusioni ttt. i,

2. Opercula reaching medially of meracanthus (figs.

59, 73). Pygofer with sharp fold at base of caudo-

dorsal beak (figs. 54, 66). Caudodorsal beak bi-

cuspidate (figs. 58, 69); broad in lateral view.

Aedeagus with short and broad lateral crests and

distineedorsalmdeesi(ties102 WA ne 3

Opercula not reaching medially of meracanthus

(cf. fig. 82). Pygofer without sharp fold at base of

caudodorsal beak. Caudodorsal beak not bicuspi-

date; slender in lateral view (cf. figs. 78-79).

Aedeagus with long and slender lateral crests and

either without, or with weak, dorsal ridges ...... 4

3. Operculum reaching beyond apex of meracan-

thus (fig. 73). Claspers very broad towards apex,

ending in two short thorn-shaped protuberances;

medial part of clasper not membranous (fig. 72).

Auditory capsules weakly swollen. Body length

under Sme ee P. plicata

— Operculum not reaching beyond apex of mera-

canthus (fig. 59). Claspers with very long and

slender apical protrusion and shorter lateral pro-

trusion; medial part of clasper membranous (fig.

56). Auditory capsules distinctly swollen. Body

lenethrovent Simi P. stoliodes

Clasper with globularly swollen dorsodistal pro-

trusion and a very long spiny apical protrusion;

medial part of clasper membranous (figs. 77, 89

arrow). Tegmen with fairly narrow hyaline bor-

deualoneihindimargn an wenn. anna. 5

Clasper without globularly swollen dorsodistal

protrusion and a fairly short apical protrusion;

medial part of clasper not membranous (figs.

100, 104). Tegmen with broad hyaline border

along hind margin

Caudodorsal beak very long and slender in lateral

view, more than 3X as long as broad (fig. 78).

Distal margin of pygofer strongly convex. Lateral

lobe of pygofer with long, slender, and distinctly

posteriorly projecting protuberance. Clasper with

sharply pointed lateral spine (fig. 77). Folded

membrane visible in ventral view. Tymbal with 8

ridges. Auditory capsules distinctly swollen.

Anterior margin of postelypeus almost continu-

ous with anterior margins of vertex lobes. Body

leneth under Sy nn... P. ungula

Caudodorsal beak fairly short, about 2X as long

as broad (fig. 84). Distal margin of pygofer weak-

ly convex. Lateral lobe of pygofer with broadly

rounded, hardly protruding protuberance. Clas-

per with bluntly rounded lateral protrusion (fig.

89). Folded membrane not visible in ventral

view. Tymbal with 5 ridges. Auditory capsules

hardly swollen. Anterior margin of postelypeus

angularly curving back at lateral corners, forming

a right angle with anterior margins of vertex

lobes. Body length over 15 mm ............ P. nana

Clasper base forming broad ring-shaped collar

with two finger-shaped dorsomedial protrusions

around base of anal valves (fig. 100). Lateral pro-

trusion of clasper shorter, or as long as, apical

protrusion. Tymbal with 6 ridges. Folded mem-

brane visible in ventral view. Caudodorsal beak

pointed at apex. Lateral protuberance on pygofer

lobe broad; pygofer forming a distinct angular

corner under lateral protuberance (fig. 93) .........

Bn eee P. bidigitula

Clasper base not forming a distinct collar around

base of anal valves (fig. 104). Lateral protrusion of

clasper distinctly longer than apical protrusion.

Tymbal with 8 ridges. Folded membrane not visible

in ventral view. Caudodorsal beak rounded at apex.

Lateral protuberance on pygofer lobe narrow; py-

gofer forming a very small angular corner under lat-

eral protuberance (fig. 105) ................ P. spinigera

Claspers parallel, only weakly diverging near api-

ces (cf. fig. 162)

Claspers strongly diverging towards apices (cf.

E aa 12

10.

11.

DE BOER: The genus Papuapsaltria

Tymbal with 6 ridges. Caudodorsal beak slender

in lateral view (cf. fig. 157)

Tymbal with 7 or 9 ridges. Caudodorsal beak

fairly broad in lateral view (cf. fig. 183) ......... 11

Body length over 19.5 mm. Distal part of opercu-

lum fairly long, longer than basal part, and with

long medial margin (fig. 163). Pygofer with nar-

row protuberance on lateral lobe (fig. 157).

Auditory capsules hardly swollen. Aedeagus not

incised at apex (fig. 158). Tegmina with long and

slender apical and ulnar areas and a broad hyaline

border along hind margin (cf. fig. 21)

rdt RAR NOR er P. dioedes

Body length under 16.5 mm. Distal part of oper-

culum shorter than basal part, and with short me-

dial margin (fig. 136). Pygofer with broad protu-

berance on lateral lobe (cf. fig. 134). Auditory

capsules distinctly swollen. Aedeagus weakly in-

cised at apex (cf fig. 131). Tegmina with short ap-

ical and ulnar areas and a fairly narrow hyaline

border along hind margin (cf. fig 137) .......... 10

Tegmina with 9 apical areas. Sternites 1 and 2

not adjacent (fig. 136). Dorsal margin of clasper

in lateral view forming a right angle with ring-

shaped elaspenbasem nenn een P. novariae

Tegmina with 8 apical areas. Sternites 1 and 2 al-

most adjacent (fig. 145). Dorsal margin of clasp-

er in lateral view almost continuous with ring-

shapediclasperjbase nur Anna. na P. dolabrata

Body length under 15 mm. Tegmina with short

apical and ulnar areas (cf. fig. 137), costa distinct-

ly swollen at pentagonal first ulnar area (fig. 176).

Anterior margin of postclypeus almost continu-

ous with anterior margins of vertex lobes.

Proximal spine of fore femur fairly long, as long

as, or longer than distance to middle spine.

Tymbal with 9 ridges. Folded membrane

between abdomen and operculum not visible in

ventral view. Aedeagus very broad in lateral view

and pick-shaped, with long and slender down-

wards directed apical part, square-shaped mem-

branous dorsal part, and short and broad lateral

crests (fig. 186). Clasper with straight and round-

ed apical part, and forming incurving dorsodistal

protuberance over aedeagus (fig. 185)

nee ae pee RE ee, era ii eae P. phyllophora

Body length over 19 mm. Tegmina with long and

slender apical and ulnar areas (cf. fig. 21), costa

not swollen at quadrangular first ulnar area (cf.

fig. 175). Anterior margin of postclypeus angu-

larly curving back at lateral corners, forming a

right angle with anterior margins of vertex lobes.

Proximal spine of fore femur very short, distinct-

ly shorter than distance to middle spine. Tymbal

with 7 ridges. Folded membrane between abdo-

men and operculum visible in ventral view.

12%

15°

14.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Aedeagus slender S-curved with long and slender

lateral crests. Clasper with recurving and pointed

apical part, not forming dorsodistal protuberance

ver ae eus (it 52) P. toxopei

Distal margin of pygofer weakly convex between

base of caudodorsal beak and lateral protu-

berance. Angular corner under lateral protu-

berance of pygofer not curved inwards or absent,

or strongly folded inwards and then thorn-

shaped; ventral half of pygofer opening generally

not horseshoe-shaped (cf. fig. 167) ................ 13

Distal margin of pygofer strongly convex between

base of caudodorsal beak and lateral protu-

berance. Angular corner under lateral protu-

berance of pygofer strongly curved inwards; ven-

tral half of pygofer opening somewhat

horseshoesshapedh(fie 22. 16

Body length over 17 mm. Folded membrane

between operculum and abdomen visible in ven-

tral view. Tegmina with long and slender apical

and ulnar areas, and a broad hyaline border along

hindimarpin (ffe Re 14

Body length under 15 mm. Folded membrane

between operculum and abdomen not visible in

ventral view. Tegmina with short apical and ulnar

areas, and a fairly narrow hyaline border along

hindkmarein (erstem a) hen 15

Clasper squarish, laminiform, with thorn-shaped

outcurving protrusion on ventral margin (fig. 15).

Pygofer with distinct rectangular corner under lat-

eral lobe (fig. 12). Distal part of operculum almost

triangular, with very long medial margin; lateral

margin concave to distolateral corner of operculum

base (fig. 17). Proximal spine of fore femur short,

much shorter than distance to middle spine.

Auditory capsules distinctly swollen. Caudodorsal

beak slender in lateral view (fig. 12) … P. angulata

Clasper hook-shaped, with sharply pointed

downwards directed apical part (fig. 171).

Pygofer with thorn-shaped, inwards curving, pro-

tuberance under lateral lobe (fig. 167). Distal

part of operculum oblong, with short medial

margin; lateral margin almost straight to disto-

15?

lateral corner of operculum base (fig. 172).

Proximal spine of fore femur fairly long, longer

than distance to middle spine. Auditory capsules

weakly swollen. Caudodorsal beak broad in later-

AMIENS RICO) Bn P. woodlarkensis

Pygofer with distinct rectangular corner under

lateral lobe. Tymbal with 7-8 ridges. Anterior

margin of postclypeus almost continuous with

anterior margins of vertex lobes. Sternites 1 and 2

adjacent. Aedeagus with small subapical lobe (fig.

Kae ee P. ustulata

Pygofer without rectangular corner under lateral

lobe. Tymbal with 6 ridges. Anterior margin of

postclypeus angularly curving back at lateral cor-

ners, forming a right angle with anterior margins

of vertex lobes. Sternites 1 and 2 not adjacent.

Aedeagus without subapical lobe (fig. 124) .........

ee, inner lues P. baasi

16. Body length over 18 mm. Distal part of opercu-

lum fairly long, longer than basal part. Sternites 1

and 2 widely separated. Aedeagus reaching not,

or only just, beyond apex of anal valves (fig. 24)

P. goniodes

Body length under 16 mm. Distal part of opercu-

lum shorter than basal part. Sternites 1 and 2 al-

most adjacent. Aedeagus reaching far beyond

apextof/analiyalyesi(efafien 34) ar ee. 17

17. Pygofer lobe with broad laminiform lateral protu-

berance (fig. 34). Clasper with concave distal

margin (fig.42). Claspers strongly diverging to-

wards apices (fig. 38). Clsaper base forming a low

collar around base of anal valves … … P. lachlani

Pygofer lobe with narrow lobate lateral protu-

berance (fig. 45). Clasper with convex distal mar-

gin (fig. 47). Claspers weakly diverging towards

apices (fig. 46). Clasper base forming a broad col-

lar around base of anal valves … … … …. P. brassi

Papuapsaltria angulata sp. n.

(figs. 12-20, 32)

Type material. — Holotype d: ‘N Guinea: NE

Bulldog Rd. 2070 m 60 km S. Wau [print]; ‘22-

Figs. 12-20. Papuapsaltria angulata sp. n. — 12, pygofer in lateral view, paratype; 13, male caudodorsal beak in dorsal view,

paratype; 14, pygofer from aslant, paratype; 15, claspers, paratype; 16, female operculum, Nami Creek; 17, male operculum,

paratype; 18, detail aedeagus apex, holotype; 19, aedeagus in lateral view, holotype; 20, female caudodorsal beak in dorsal

view, Nami Creek.

Lettering: bp = basal part of operculum; c = crest around distolateral corner of basal part of operculum; cb = caudodorsal beak;

di = distal margin of pygofer; dm = distomedial margin of operculum; do = dorsal margin of pygofer; dp = distal part of op-

erculum; fm = folded membrane; lm = lateral margin of operculum; m = medial margin of operculum; me = meracanthus; pr

= protuberance on lateral lobe of pygofer; rc = rectangular corner below lateral protuberance of pygofer; ve = ventral margin

of pygofer.

Figs. 22-31. Papuapsaltria goniodes sp. n. — 22, pygofer from behind, Ayura; 23, pygofer from aslant, Ayura; 24, pygofer in

lateral view, Daulo, arrow indicating distal lobe; 25, male caudodorsal beak in dorsal view, Daulo; 26, claspers, Daulo; 27, ae-

deagal apex, holotype; 28, aedeagus in lateral view, holotype; 29, male operculum, Daulo; 30, female caudodorsal beak in dor-

sal view, Daulo; 31, female operculum, Daulo.

Lettering: st 1 = sternite 1; st 2 = sternite 2.

DE Boer: The genus Papuapsaltria

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

31.v.1969° [print]; ‘J. Sedlacek Collector Bishop

Mus.’ [print], BPBM. — Paratypes: PAPUA NEW GUINEA:

same data as holotype, 19, BPBM; Nami Ck., Wau,

1700-1800 m, 17.ix.1965, J. Sedlacek, 16, ZMAN;

Nami Ck., 6 km W. Wau, 1700 m, 12.vi.1962, J.

Sedlacek, 12, BPBM.

P. angulata is the type species of Papuapsaltria.

Males are easily recognized by the broad and rectan-

gular clasper, with slender, sharply pointed, and later-

ally bending ventral protrusion.

Description

Body light brown and unmarked. Females slightly

larger than males, with distinctly longer tegmina.

Tegmina of males 1.1 X as long as body length, of fe-

males 1.4. Male abdomen 1.7-1.8X as long as

head and thorax, of females 1.2 x.

Head: Ochraceous brown, without setae. Postcly-

peus angularly protruding, anterior margin angularly

bent medially, and almost continuous with anterior

margins of vertex lobes. Postclypeus not swollen; an-

terior margin (lateral view) weakly concave. Distance

between lateral ocelli 0.9-1.0 as long as distance

between lateral ocellus and eye, and 1.9-2.3 X as wide

as frontal ocellus.

Tegmina: Hyaline, with 8 long and slender apical

areas, a broad hyaline costal area and a very broad

hyaline border along hind margin.

Legs: Fore femur with three very short, about

equally long, erect spines. Proximal spine finger-

shaped and distinctly shorter than distance to middle

spine.

Tymbal: Five parallel transverse sclerotized ridges

spanning the tymbal from dorsal to ventral margin

and a 6th, most proximal, ridge almost reaching ven-

tral margin. Intercalary ridges can hardly be dis-

cerned.

Opercula: Male operculum (fig. 17) not covering

tymbal cavity in ventral view, leaving a wide gap

between operculum and abdomen; folded membrane

clearly visible in ventral view. Distal part of opercu-

lum longer than basal part and almost triangle-

shaped, medially elongate. Lateral margin bending

Fig. 21. Papuapsaltria goni-

odes sp. n. Left tegmen,

male Mt. Otto.

convexly into straight distal margin and concavely

into crest of basal part. The lateral part of this crest

short and knobby. Distomedial corner rectangular.

Medial margin long and straight. Meracanthus long-

er than operculum, but not reaching to abdomen.

Female operculum (fig. 16) much shorter than that of

male, with oblong shaped distal part.

Abdomen: Male abdomen inflated. First tergite

short, partly hidden under metanotum. Medial part

of 2nd tergite less than 2X as long as lateral part.

Anterior margin of 2nd tergite distinctly concave me-

dially. First and 2nd sternite adjacent. Auditory cap-

sules weakly developed, not visible in dorsal view.

Female abdomen very slender. Female caudodorsal

beak (fig. 20) slender, pointed at apex. Ovipositor

sheaths just reaching to apex of caudodorsal beak.

Male genitalia: Pygofer in lateral view as in fig. 12.

Dorsal margin weakly convex, continuous with short

and very slender caudodorsal beak. Distal margin

weakly convex between caudodorsal beak and lateral

protuberance, forming a weakly developed distal

lobe. Caudodorsal beak in dorsal view (fig. 13) short

and slender, triangle-shaped and rounded at apex.

Lateral lobe of pygofer curving outwards towards ven-

tral margin, forming a weakly developed, but broad

and rectangular laminiform protuberance. Pygofer

forming a very distinct rectangular and slightly in-

curving corner just below this protuberance (figs. 12,

14). Ventral margin of pygofer angularly convex.

Ventral half of pygofer opening in holotype horse-

shoe-shaped (seen from behind cf. fig. 22), with

broad, rounded, basal margin and concave lateral

margins, incurving towards rectangular corners, but

narrower, V-shaped with converging margins, in

paratype. Clasper (fig. 15) laminiform, with almost

rectangular dorsodistal corner and straight distal mar-

gin, and with a sharply pointed thorn-shaped protru-

sion at half-length of ventral margin. Protrusion of

clasper bending strongly outwards and reaching later-

ally beyond pygofer margin. Rectangular dorsodistal

corner of clasper bending mesiad, distally of aedeagus

and supporting aedeagus in upright position.

Claspers strongly diverging towards ventrodistal cor-

ners. Clasper base forming a smoothly rounded,

though discontinuous ring around base of anal valves;

with dorsal parts of clasper bases curving mesiad,

proximally of aedeagus, and almost touching each

other. Aedeagus standing erect between claspers, fair-

ly close to anal valves, but with apical part curving

away, reaching beyond distal margins of claspers.

Aedeagus (fig. 19) long and slender, weakly S-curved,

with long slender lateral crests and a pair of weakly

developed dorsal crests, restricted to its distal half.

Aedeagus strongly curved near apex. Aedeagal pore

round (fig. 18).

Measurements: Body length d: 17.5 & 18.6 mm,

2 19.5 mm; tegmen length d: 21.2 mm, 9: 28.1

mm; head length d: 1.2 mm, ?: 1.2 & 1.5 mm; pro-

notum length d: 1.7 mm, 9: 2.0 mm; mesonotum

length d: 3.5 mm, 9: 5.7 mm; head width d: 3.1 &

3.3 mm, 2: 3.9 mm; width of pronotal collar d : 4.3

& 4.8 mm, 9: 6.2 mm.

Distribution (fig. 32). — Central Papua New

Guinea, the most western part of the Papuan penin-

sula.

Etymology. — Angulatus (Latin) means angular and

refers to the angular claspers.

Papuapsaltria goniodes sp. n.

(figs. 5, 7-9, 11, 21-32)

Type material. — Holotype d : ‘NEW GUINEA (NE)

Mt. Otto, 2200 m, June 24, 1955’ [print]; J.L.

Gressitt collector’ [print], BPBM. — Paratypes: PAPUA

NEW GUINEA: Aiyura, nr Kainantu, 1700-2200 m,

6.1.1965, J.L. Gressitt, 1d, ZMAN; Daulo Pass, 2400

m, 15.v.1963, J. Sedlacek, 16, BPBM; Daulo Pass,

Asaro-Chiambu Div., 3000 m, 13.vi.1955, J.L.

Gressitt, 1d, BPBM; same data but 8500 ft, J.J.M.

Scent-Ivany, 19, MVMA; Mt. Hagen, 1959, E.

Reiner, 14, SMED; Mt. Piora, 6°45’ S 146°00’ E,

2100 m, 12.vi.1966, G.A. Samuelson, 19, BPBM.

P. goniodes is slightly larger than P. angulata. These

two species share a similar, almost triangularly elon-

gate operculum but the male genitalia differ consider-

ably. P. goniodes can be recognized by its variable teg-

men venation and its deviating shape of the head,

with the lateral ocelli somewhat wider apart than in

other species of Papuapsaltria.

Description

Body light brown to dark reddish brown, unmar-

ked. Females slightly smaller than males, but with

more robust head and thorax, and longer tegmina.

Tegmina of males 1.2-1.3 X as long as body length,

of females 1.4-1.5X. Male abdomen 1.5-1.9X as

long as head and thorax, of females 1.2-1.3 X.

Head (fig. 5): Reddish brown, without setae.

Postclypeus much broader than in P. angulata and

De BOER: The genus Papuapsaltria

bluntly rounded, anterior margin weakly convex, al-

most continuous with anterior margins of vertex

lobes. Postclypeus not swollen; anterior margin (later-

al view) straight or weakly concave (fig. 11). Distance

between lateral ocelli distinctly longer (1.1-1.2) than

distance between lateral ocellus and eye, and 2.0-

2.6 X the width of frontal ocellus.

Tegmina: Hyaline. Venation variable (fig. 21), of

ten differing between left and right tegmen of indi-

viduals, with 9-12 apical areas and, rarely, a subapical

area between ulnar and apical areas. Only 9-10 apical

areas reach the ulnar or basal areas, others are formed

by a more distal splitting of veins. Tegmen generally

with a very broad hyaline costal area and a very broad

hyaline border along hind margin.

Legs: Fore femur with three very short, about

equally long, erect spines. Proximal spine finger-

shaped and distinctly shorter than distance to middle

spine (figs. 8-9).

Tymbal: Five parallel transverse sclerotized ridges

spanning the tymbal from dorsal to ventral margin, a

6th ridge almost reaching ventral margin and a 7th,

most proximal, ridge only partly separated from prox-

imal tymbal margin, reaching to about half the tym-

bal width. Six short intercalary ridges clearly visible.

Opercula: Male operculum (fig. 29) not covering

tymbal cavity in ventral view, leaving a wide gap

between operculum and abdomen; folded membrane

sangulata 0°

vbrassi

e goniodes

Alachlani

4°

8 (e)

122

140° 144° 148°

Fig. 32. Localities of Papuapsaltria angulata, P. brassi, P.

goniodes, and P. lachlani.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

clearly visible in ventral view. Distal part of opercu-

lum longer than basal part and medially elongate, al-

most triangle-shaped, as in P. angulata, but wich lat-

eral margin often bending more angularly from crest

of basal part. Lateral margin convexly bent into al-

most straight distal margin. Distomedial corner of

operculum almost rectangular, medial margin long

and straight. Meracanthus slightly longer than oper-

culum, but not reaching abdomen. Female opercu-

lum (fig. 31) much shorter than that of males. Distal

part shorter than basal part, oblong. Distal margin

straight, in one specimen slightly convex at half-

length, distomedial corner rectangular, medial mar-

gin short and straight.

Abdomen: Male abdomen inflated. First tergite

short, partly hidden under metanotum. Medial part of

2nd tergite less than 1.5X as long as lateral part.

Anterior margin of 2nd tergite very weakly concave

medially. First and 2nd sternite generally adjacent, but

sometimes separated. Auditory capsules well devel-

oped, protruding and visible in dorsal view. Female ab-

domen slender. Female caudodorsal beak (fig. 30) slen-

der and sharply pointed at apex. Ovipositor sheaths not

reaching to apex of caudodorsal beak.

Male genitalia: Pygofer in lateral view as in fig. 24.

Dorsal margin weakly convex, continuous with short

caudodorsal beak. Distal margin distinctly convex

between caudodorsal beak and lateral protuberance,

forming a weakly protruding lobe on pygofer (fig. 24

arrow). Caudodorsal beak fairly slender in lateral

view, but broader at its base than in P. angulata, tri-

angular with concave margins and pointed at apex

(fig. 25). Lateral lobe of pygofer with narrow, round-

ed, and outcurving lobate protuberance. Pygofer

forming a distinct rectangular and slightly incurving

corner just below this protuberance (figs. 23-24).

Ventral margin of pygofer angularly convex. Ventral

half of pygofer opening horseshoe-shaped (fig. 22),

with broad, rounded basal margin and concave later-

al margins; in curving towards rectangular corners.

Clasper (fig. 26) very broad and short, square-shaped

in lateral view. Rectangular distodorsal corner of

clasper bending mesiad, distally of aedeagus and sup-

porting aedeagus in upright position. Distal margin

of clasper straight, but slightly recurving to sharply

pointed, downwards directed, thorn-shaped apex.

Claspers slightly diverging towards apices. Clasper

base forming a low ring-shaped collar around base of

anal valves. This collar is medially indented around

aedeagus, and less distinct than in foregoing species.

Aedeagus standing erect between claspers, close to

anal valves, but with apical part curving away, reach-

ing well beyond distal margins of claspers. Aedeagus

(fig. 28) weakly S-curved, most strongly curved and

distinctly narrowing near apex, its most apical part al-

most rectangularly bent. Aedeagus with long and dis-

tinct lateral crests, a rounded middorsal ridge and a

pair of weakly developed dorsal crests near apical

bending point. Apex of aedeagus with slight medial

incision (fig. 27).

Measurements: Body length d: 18.9-20.5 mm

(x 19.9 mm), 2: 17.8 & 18.0 mm; tegmen length d:

23.0-25.3 mm (x 24.0 mm), 2: 26.0 & 27.1 mm;

head length d: 1.2-1.5 mm, 9: 1.5 mm; pronotum

length d: 1.9-2.0 mm, 9: 2.1 & 2.2 mm; mesono-

tum length d: 4.0-4.7 mm, 9: 4.7 & 4.8 mm; head

width d: 3.6-3.9 mm, 9: 4.0 mm; width of pronotal

collar &: 4.7-5.5 mm, 9: 5.6 & 5.7 mm.

Distribution (fig. 32). — P. goniodes is distributed in

central northern Papua New Guinea, just west of the

Huon peninsula.

Etymology. — Goniodes (Greek) means angular

and refers to the angular claspers.

Papuapsaltria lachlani sp. n.

(figs. 32-42)

Type material. — Holotype d : ‘PAPUA NEW GUINEA

Wau 28 Dec 1970 R.B. Lachlan [print]; AMSA. —

Paratypes: IRIAN JAYA: Ifar, Cyclops Mts., 400-800

m, 7-9.ix.1962, J. Sedlacek, 16, BPBM; PAPUA NEW

GUINEA: Kiambavi vill., Saidor, Finisterre Range, 1-

28.viii.1958. W.W. Brandt, 1d, ZMAN; Wau, Big

Wau Creek, 1200 m, Thomas W. Davies, 19, CAS;

Wau, Morobe Distr., 20.x.1969, James E. Tobler,

19, cas; Wau, Morobe Distr., Mt. Missim, 2080 m,

17.111.1966, Gressitt, 16, BPBM.

P. lachlani closely resembles P. goniodes in shape of

male genitalia, but is distinctly smaller and has almost

oblong-shaped opercula. Some specimens have an

aberrant tegmen venation.

Description

Body red-brown to greenish. Females slightly smal-

Figs. 33-42. Papuapsaltria lachlani sp. n. — 33, pygofer from aslant, holotype; 34, pygofer in lateral view, holotype; 35, male

caudodorsal beak in dorsal view, holotype; 36, pygofer in lateral view, Ifar; 37, aedeagal apex, holotype; 38, claspers from be-

hind, holotype; 39, male operculum, holotype; 40, female operculum, Big Wau Creek; 41, female caudodorsal beak in dor-

sal view, Big Wau Creek; 42, claspers, holotype.

Figs. 43-52. Papuapsaltria brassi sp. n. — 43, pygofer from aslant, paratype; 44, male caudodorsal beak in dorsal view, holo-

type; 45, pygofer in lateral view, paratype; 46, claspers from behind, holotype; 47, claspers, holotype; 48, aedeagus in lateral

view, paratype; 49, aedeagal apex, paratype; 50, male operculum, holotype; 51, female operculum, Purosa Camp; 52, female

caudodorsal beak in dorsal view, Purosa Camp.

DE BOER: The genus Papuapsaltria

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

ler than males, but with more robust head and thorax.

Tegmina of males 1.1-1.3X as long as body length,

of females 1.3-1.4X. Male abdomen 1.4-1.7X as

long as head and thorax, of females 0.9-1.2 x.

Head: Reddish brown, often with some short red-

brown setae on postclypeus and anterior parts of ver-

tex. Diverging fissures on vertex fairly distinct in

holotype, almost obsolete in paratypes. Postclypeus of

holotype and females oblong and angularly protrud-

ing beyond vertex lobes, with almost straight anterior

margin, angularly bending back at lateral corners and

forming an almost right angle with anterior margins

of vertex lobes. Postclypeus of other material with

weakly convex anterior margin, almost continuous

with anterior margins of vertex lobes. Postclypeus

slightly swollen; anterior margin (lateral view) weakly

convex. Distance between lateral ocelli in males as

long as, in females slightly longer than distance

between lateral ocellus and eye, and 2.0-2.3X the

width of frontal ocellus.

Thorax: Pronotum with weak medial furrow.

Tegmina: Hyaline, venation sometimes variable,

forming 8-10 long and slender apical areas and a very

distinct hyaline costal area. Tegmen with a broad

hyaline border along hind margin.

Legs: Fore femur with three very short, about

equally long, erect spines. Proximal spine often ex-

tremely short and finger-shaped, though fairly long

and pointed in specimen from Saidor, and always

shorter than distance to middle spine.

Tymbal: Six parallel transverse sclerotized ridges

spanning the tymbal from dorsal to ventral margin

and a 7th, most proximal, ridge almost reaching ven-

tral margin. Six short intercalary ridges can hardly be

discerned.

Opercula: Male operculum (fig. 39) very short, not

covering tymbal cavity in ventral view; leaving a wide

gap between operculum and abdomen; folded mem-

brane partly visible in ventral view. Distal part of op-

erculum distinctly shorter than basal part and angu-

lar, almost oblong-shaped. Lateral margin almost

straight, bending gradually into crest of basal part and

angularly into long and straight distal margin. Distal

margin forming a sharp angle with short and straight

medial margin. Meracanthus reaching well beyond

operculum, but not reaching abdomen. Female oper-

culum (fig. 40) as in male, but shorter.

Abdomen: Male abdomen not inflated. First ter-

gite short, partly, or completely, hidden under meta-

notum. Medial part of 2nd tergite less than 1.5 X as

long as lateral part. Anterior margin of 2nd tergite

weakly convex medially. First and 2nd sternite not

adjacent. Auditory capsules swollen and clearly visible

in dorsal view. Female caudodorsal beak (fig. 41) tri-

angular, almost pointed at apex. Ovipositor sheaths

reaching just beyond apex of caudodorsal beak.

Male genitalia: Very similar to P. goniodes. Pygofer

in lateral view as in fig. 34. Dorsal margin weakly

convex, continuous with short and slender caudodor-

sal beak. Distal margin strongly convex between cau-

dodorsal beak and lateral protuberance, forming an

almost rectangular and incurving lobe. Caudodorsal

beak in dorsal view broad, almost rectangularly pro-

truding in Saidor specimen, but narrower and apical-

ly rounded in others (fig. 35). Lateral lobe of pygofer

with distinct, angularly rounded, lobate, outcurving

laminiform protuberance. Pygofer forming a distinet

rectangular and slightly incurving corner just below

this protuberance (fig. 33). Ventral margin of pygofer

straight. Ventral half of pygofer opening horseshoe-

shaped (cf. fig. 22), with broad, rounded basal margin

and concave lateral margins; incurving towards rec-

tangular corners. Claspers (fig. 42) as in P. goniodes,

very broad and short, square-shaped in lateral view.

Dorsal margin of clasper straight or weakly convex.

Rectangular distodorsal corner of clasper bending

mesiad, distally of aedeagus and supporting aedeagus

in upright position. Distal margin of clasper straight,

but slightly concave to sharply pointed, downwards

directed, thorn-shaped apex. Claspers strongly di-

verging towards apices (fig. 38). Clasper base forming

a low ring-shaped collar around base of anal valves.

This collar medially indented around aedeagus.

Aedeagus standing erect between claspers, close to

anal valves, with its apical part curving away, reaching

well beyond distal margins of claspers. Aedeagus of

holotype very long, reaching far beyond apex of cau-

dodorsal beak (fig. 34), in other specimens much

shorter, curved directly above dorsal margins of clasp-

ers (fig. 36). Aedeagus weakly S-curved, angularly but

slightly bent at about 3/4 its length (fig. 33) and

strongly curved near apex, with short straight subapi-

cal part. Lateral crests long and distinct. Apex of ae-

deagus slightly dilated, with round pore (fig. 37).

Measurements: Body length d: 13.7-14.0 mm, 9:

13.3 & 13.7 mm; tegmen length d: 16.6-20.1 mm,

2:18.2 & 18.4 mm; head length d: 1.3-1.4 mm, 9:

1.4 mm; pronotum length d: 1.4-1.7 mm, 9: 1.6 &

1.7 mm; mesonotum length d: 2.3-3.1 mm, 9: 3.1

mm; head width &: 3.1-3.3 mm, 9: 3.3 mm; width

of pronotal collar d: 3.7-4.1 mm, 2: 4.2 & 4.3 mm.

Distribution (fig. 32). — P. lachlani is widely dis-

tributed in northern New Guinea and the western

part of the Papuan peninsula.

Etymology. — This species is named in honour of

Mr R.B. Lachlan who collected the holotype.

Papuapsaltria brassi sp. n.

(figs. 32, 43-52)

Type material. — Holotype d: ‘No 10, Purosa

Camp, Okapa area, 1950 m, ix-25-1959° [print];

‘Eastern Highlands District L.J. Brass coll’ [print];

‘sixth Archbold Exped. to Papua New Guinea’ [print];

‘AMNH New York’ [print], AMNH. — Paratypes: PAPUA

NEW GUINEA: same data as holotype but 29.ix.1959,

16,19, AMNH; Okapa, 13 km SE, 1650-1870 m, 26.-

viii.1964, J. & M. Sedlacek, 1, BPBM; Okapa, 1900

m, 3.vi.1967, G.A. Samuelson, 1 9, BPBM.

P. brassi strongly resembles P. lachlani, but is

slightly larger and has slightly shorter and less strong-

ly diverging claspers. P. brassi is most easily separated

from P. lachlani by a more oval-shaped male opercu-

lum, a narrower gap between operculum and abdo-

men and smaller protrusions on the lateral lobes of

pygofer.

Description

Body yellow-brown or olive green. Females larger

than males. Tegmina of males 1.3 X as long as body

length, of females 1.4X. Male abdomen 1.4-1.6X as

long as head and thorax, of females 1.2-1.3 x.

Head: Reddish brown, with some short red-brown

setae on postclypeus and vertex. Postclypeus hardly

protruding, anterior margin weakly convex, almost

continuous with anterior margins of vertex lobes, and

not swollen; anterior margin (lateral view) weakly

concave. Distance between lateral ocelli 1.0-1.2 X as

long as distance between lateral ocellus and eye and

1.9-2.3 X the width of frontal ocellus.

Legs: Fore femur with row of three erect spines.

Most proximal spine very short, hardly longer than

middle spine and much shorter than distance to mid-

dle spine.

Tegmina: Hyaline, with 8 apical areas, a narrow

but distinct costal area and a fairly narrow hyaline

border along hind margin.

Tymbal: Six parallel transverse sclerotized ridges

spanning the tymbal from dorsal to ventral margin

and a 7th, most proximal, ridge almost reaching ven-

tral margin. Six short intercalary ridges can hardly be

discerned.

Opercula: Male operculum (fig. 50) very short, not

covering tymbal cavity in ventral view, but leaving

only a narrow gap between operculum and abdomen;

folded membrane not visible in ventral view. Distal

part of operculum distinctly shorter than basal part

and oval-shaped. Lateral margin very short, bending

gradually into crest of basal part and convexly curving

into long and weakly convex distal margin.

Distomedial corner broadly rounded, medial margin

weakly convex. Meracanthus reaching well beyond

operculum and beyond anterior margin of abdominal

segment 2. Female operculum (fig. 51) much shorter

than that of males. Distal part much shorter than ba-

sal part, nearly oblong, with weakly convex distal

DE BOER: The genus Papuapsaltria

margin. Distolateral corner rounded, distomedial

corner almost rectangular. Medial margin short and

straight.

Abdomen: Male abdomen not inflated. First ter-

gite short, either partly, or completely, hidden under

metanotum. Medial part of 2nd tergite hardly longer

than lateral parts. Anterior margin of 2nd tergite

weakly convex medially. First and 2nd sternite not

adjacent. Auditory capsules swollen and clearly visible

in dorsal view. Female abdomen of same shape as in

male. Female caudodorsal beak in dorsal view (fig.

52) slender triangular, narrowly rounded at apex.

Ovipositor sheaths just reaching beyond apex of cau-

dodorsal beak.

Male genitalia: Pygofer in lateral view (fig. 45)

closely resembling that of P. lachlani. Caudodorsal

beak very slender and straight, dorsally not continu-

ously rounded with convex dorsal margin of pygofer.

Distal margin of pygofer strongly convex between

caudodorsal beak and lateral protuberance, forming a

fairly distinct inwards curving lobe. Caudodorsal

beak narrowly triangular with concave margins, and

apically rounded (fig. 44). Lateral lobe of pygofer

with distinct, narrowly rounded lobate outcurving

protuberance. Pygofer forming a rectangular and

slightly incurving corner just below this protu-

berance, which is smaller than in P. /achlani and

hardly visible in lateral view (figs. 43, 45). Ventral

margin of pygofer straight. Ventral half of pygofer

opening horseshoe-shaped (cf. fig. 22), with broad,

rounded basal margin and concave lateral margins;

incurving towards rectangular corners. Claspers (fig.

47) as in P. lachlani, but reaching less far beyond py-

gofer margin. Dorsal margin of clasper straight.

Rectangular distodorsal corner of clasper bending

mesiad, distally of aedeagus and supporting aedeagus

in upright position. Distal margin of clasper weakly

convex towards thorn-shaped apex. Claspers weakly

diverging towards apices (fig. 46). Clasper base form-

ing a very distinct ring-shaped collar around base of

anal valves. This collar is medially indented around

aedeagus. Aedeagus standing erect between claspers,

close to anal valves, but with apical part curving away,

reaching well beyond distal margins of claspers.

Aedeagus (fig. 48) weakly S-curved, with short, but

distinct, lateral crests restricted to its proximal half,

and a distinct and slender dorsal crest. Aedeagal pore

almost round (fig. 49).

Measurements: Body length d : 14.6-15.5 mm, ©:

15.6 & 16.1 mm; tegmen length d: 19.1-19.6 mm,

9: 22.3 & 22.4 mm; head length d: 1.1-1.2 mm, 9:

1.3 mm; pronotum length d: 1.6 mm, 9: 1.8 mm;

mesonotum length d: 3.1-3.5 mm, 9: 3.5 & 4.2

mm; head width d: 3.1-3.2 mm, ®: 3.3 & 3.4 mm;

width of pronotal collar d: 3.9-4.3 mm, 9: 4.2 &

4.9 mm.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Distribution (fig. 32). — All specimens come from

Okapa, in the Fastern Highlands of Papua New

Guinea.

Etomology. — This species is named in honour of

Mr L.J. Brass who collected the holotype, and many

other specimens on which the current revision of the

‘Baeturia and related genera complex’ is based.

Papuapsaltria stoliodes sp. n.

(figs. 3, 53-64)

Type material. — Holotype d : ‘NEW GUINEA: PAPUA

/ Owen Stanley Range / Goilala: Bome, 1950 m /

iv.16-30.1958° [print]; “W.W. Brandt / Collector /

BISHOP’ [print], BPBM. — Paratype: PAPUA NEW GUIN-

EA: Wau, 1100-1300 m, 2.1.1966, L. & M. Sedlacek,

1d, BPBM.

Other material: PAPUA NEW GUINEA: Mt. Kaindi,

30.iv.1962. J.L. Gressitt, 19, BPBM; Edi Creek, 2000 m,

31.v.1962, J. Sedlacek, 18, BPBM; Mt Missim, 2100 m,

21.11.1968, J.H. Sedlacek, 19, BPBM; Wau, 9-12.1.1962, J.,

J.H., & M. Sedlacek, G. Monteith & native collectors, 19,

BPBM.

Males of P. stoliodes are easily recognized by the

folded pygofer and the long and slender clasper, with

a thorn-shaped lateral protuberance. Three females

from the same area presumably belong to this species

but are not included in the type series.

Description

Body light brown. Females smaller than males, but

with more robust head and thorax, and longer tegmi-

na. Tegmina of males 1.2-1.3X as long as body

length, of female 1.6 X. Male abdomen 1.5-1.9X as

long as head and thorax, of females 1.1-1.3 X.

Head: Postclypeus angularly protruding, smoothly

rounded anteriorly, its anterior margin weakly con-

vex, almost continuous with anterior margins of ver-

tex lobes. Postclypeus not swollen; anterior margin

(lateral view) concave (fig. 53). Distance between lat-

eral ocelli 0.8-0.9 X as long as distance between later-

al ocellus and eye and 1.5-2.2 X the width of frontal

ocellus.

Legs: Fore femur (fig. 57) with row of three erect

spines. Most proximal spine very short, hardly longer

than middle spine and much shorter than distance to

middle spine.

Tegmina: Hyaline, venation in holotype with 11-

12, in other material with 8, long and slender apical

areas. Tegmen with a distinct costal area and a broad

hyaline border along hind margin.

Tymbals: Six slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin,

and a 7th, most proximal, ridge spanning only about

3/4 of tymbal width. Lateral band of seven intercalary

ridges weakly developed.

Opercula: Male operculum (fig. 59) quite large,

covering greater part of tymbal cavity; folded mem-

brane not visible in ventral view. Distal part of oper-

culum angularly oval-shaped, medially distinctly

reaching beyond meracanthus. Lateral margin short

and straight, angularly bending into long and angu-

larly convex distal margin and angularly bending into

crest of basal part. Medial margin angularly bent at

half-length. Meracanthus reaching beyond opercu-

lum, and just beyond anterior margin of abdominal

segment 2. Distal part of female operculum (fig. 63)

shorter than basal part and angularly rounded, sickle-

shaped.

Abdomen: Male abdomen distinctly inflated. First

tergite very short and medially hidden under metano-

tum. Medial part of 2nd tergite fairly long, about

twice as long as lateral parts. Anterior margin of 2nd

tergite weakly concave medially. Auditory capsules

distinctly swollen. First and 2nd sternites adjacent.

Female caudodorsal beak in dorsal view (fig. 64)

broad, triangular, and bluntly rounded at apex.

Ovipositor sheaths reaching just beyond apex of cau-

dodorsal beak.

Male genitalia: Pygofer in lateral view (fig. 54) with

sharp fold at base of caudodorsal beak. Dorsal margin

of pygofer convex, though weakly concave towards

proximal end, and continuously curved with broad

and stout caudodorsal beak. Distal margin forming

an obtuse angle with margin of beak, and an angular-

ly rounded, inwardly curved lobe just above lateral

protuberance. Ventral margin straight, but angularly

bent near lateral protuberance, though not forming a

distinct or protruding corner (figs. 54-55). Ventral

margins weakly converging to base of pygofer; basal

part of pygofer opening broad, U-shaped.

Caudodorsal beak in dorsal view (fig. 58) very broad

and stout, bicuspidate; medially incised at distal mar-

gin. Lateral lobe of pygofer strongly curving inwards

towards end of distal margin, forming a short, dorso-

ventrally flattened, and distinctly posteriorly project-

Figs. 53-64. Papuapsaltria stoliodes sp. n. — 53, postclypeus in lateral view, holotype; 54, pygofer in lateral view, holotype; 55,

pygofer from aslant, holotype; 56, claspers, holotype; 57, male fore femur, Wau; 58, male caudodorsal beak in dorsal view,

holotype; 59, male operculum, holotype; 60, aedeagus in dorsal view, holotype; 61, aedeagal apex in lateral view, holotype;

62, aedeagus in lateral view, holotype; 63, female operculum, Wau; 64, female caudodorsal beak in dorsal view, Wau.

Figs. 66-75. Papuapsaltria plicata sp. n. — 66, pygofer from aslant, Mt. Missim; 67, pygofer in lateral view, Mt. Missim; 68,

male first and second tergites in dorsal view, Mt. Missim; 69, male caudodorsal beak in dorsal view, Mt. Missim; 70, aedea-

gal apex, Bulldog Road; 71, aedeagus in lateral view, Bulldog Road; 72, claspers, Mt. Missim; 73, male operculum, Mt.

Missim; 74, female operculum, Mt. Missim; 75, female caudodorsal beak in dorsal view, Mt. Missim.

DE BOER: The genus Papuapsaltria

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

ing protuberance. Claspers (fig. 56) very slender, la-

miniform and slightly diverging towards apices.

Dorsal margin straight, curving down at rounded dis-

todorsal corner. Clasper slightly bending inwards dis-

tally of aedeagus. Apical part very long and slender,

with almost membranous and transparent medial

part, arching between apex and base of clasper.

Clasper forming a short, sharply pointed thorn-

shaped lateral protrusion at its base, in paratype dis-

tinctly longer than in holotype. Aedeagus (fig. 62)

short and almost straight, most strongly curved near

apex. Lateral crests broadening, almost angularly am-

plified, towards proximal ends and curving upwards

at distal ends, fusing to a small ridge at apical bending

point of aedeagus. Pair of distinct dorsal ridges (fig.

60) abruptly ending at angularly protruding distal

corners. Aedeagal pore almost round (fig. 61).

Remark: The male from Edi Creek lacks the great-

er part of its abdomen.

Measurements: Body length d: 18.6 & 19.1 mm,

2: 17.6-18.7 mm; tegmen length d': 20.0-25.0 mm,

? : 28.8 mm; head length d: 1.2-1.3 mm, 2: 1.5-1.6

mm; pronotum length d: 1.8-2.0 mm, 9: 2.0-2.4

mm; mesonotum length ó: 3.8-4.1 mm, 9: 4.6-5.0

mm; head width d: 3.1-3.4 mm, 9: 3.7-3.9 mm;

width of pronotal collar d: 5.0-5.2 mm, 9: 5.7-5.8

mm.

Distribution (fig. 3). — P. stoliodes is endemic to the

central western part of the Papuan peninsula.

Etymology. — Stoliodes (Greek) means folded and

refers to the peculiarly folded pygofer.

Papuapsaltria plicata sp. n.

(figs. 3, 65-75)

Type material. — Holotype d : ‘ NEW GUINEA (NE)

/ Mt. Kaindi, 16 km / SW of Wau, 2200 m / 8-

9.vi.1962 [print]; ‘Light Trap / J. Sedlacek / BISHOP?

[print], BPBM. — Paratypes: PAPUA NEW GUINEA: same

data as holotype 16, BPBM; Bulldog Rd., c 14 km S.

Edie Creek, 2450 m, 4-10.vii.1966, G.A. Samuelson,

1d, BPBM; Mt. Kaindi, 2400 m, 27.1.1963, J. Sedla-

Fig. 65. Papuapsal-

tria plicata sp. n. Left

tegmen, male Bull-

dog Road.

cek, 16, BPBM; Mt. Kaindi, 2350 m, 26.v.1967, J.L.

Gressitt, 16, BPBM; Mt. Kaindi, 9.vi.1962, J. & M.

Sedlacek, 1 ®, BPBM; Mt. Missim, 2400-2800 m, 22-

30.v.1968, J.L. Gressitt, R.C.A. Rice & J. Sedlacek,

1d, BPBM; Mt. Missim, 2400 m, 21.iv.1968, J. & M.

Sedlacek, 15, ZMAN; Mt. Missim, Morobe Prov.,

2400 m, 23.iv.1968, J. Sedlacek, 16, BPBM; same

data 1d, ZMAN; Mt. Missim, Wau, Morobe Distr.,

1200-1800 m, 8.xii.1963, H. Clissold, 19, ZMAN;

Mt Strong, 2600-3000 m, 8-10.1.1968, J. & M.

Sedlacek, 1 4, BPBM.

P. plicata closely resembles P. stoliodes, sharing a

very similarly folded pygofer, but is distinctly larger.

Males have a larger operculum and a shorter, bicuspi-

date, clasper.

Description

Body light brown. Females smaller than males, but

with more robust head and thorax, and longer tegmi-

na. Tegmina of males 1.2-1.3X as long as body

length, of females 1.6 X. Male abdomen 1.7-2.2 X as

long as head and thorax, of females 1.1-1.3 X.

Head: Postclypeus fairly narrow, angularly pro-

truding, anterior margin weakly convex, almost con-

tinuous with anterior margins of vertex lobes.

Postclypeus weakly swollen; anterior margin (lateral

view) almost straight. Ocelli large and fairly close to-

gether; distance between lateral ocelli 0.6-0.9 X as

long as distance between lateral ocellus and eye and

1.2-2.0 X the width of frontal ocellus.

Legs: Fore femur with row of three erect spines.

Most proximal spine very short, though longer than

middle spine, much shorter than distance to middle

spine.

Tegmina: Hyaline, venation variable, generally

with 8, but sometimes with 9-10 long and slender ap-

ical areas (fig. 65); additional areas often caused by

splitting of the 8th apical area. Tegmen with distinct

costal area and fairly broad hyaline border along hind

margin.

Tymbals: Six slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin,

and a 7th, most proximal, ridge spanning only about

3/4 of tymbal width. Lateral band of seven intercalary

ridges weakly developed.

Opercula: Male operculum (fig. 73) larger than in

P. stoliodes, almost completely covering tymbal cavity;

folded membrane not visible in ventral view. Distal

part of operculum angular and oblong, medially dis-

tinctly reaching beyond meracanthus. Lateral margin

long and straight, forming a distinct and obtuse angle

with crest of basal part and with distal margin. Distal

margin long and straight, sometimes weakly concave

at about half-length. Medial margin angularly con-

vex. Meracanthus not reaching distal margin of oper-

culum. Distal part of female operculum (fig. 74)

about as long as basal part and oblong-shaped, with

convex distal margin and rounded corners.

Abdomen: Male abdomen distinctly inflated.

Medial part of 2nd tergite fairly long, about twice as

long as lateral parts (fig. 68). Anterior margin of 2nd

tergite weakly concave medially. First and second

sternites adjacent. Auditory capsules weakly inflated.

Female caudodorsal beak in dorsal view (fig. 75) nar-

row triangular and bluntly rounded at apex.

Ovipositor sheaths not reaching beyond apex of cau-

dodorsal beak.

Male genitalia: Pygofer in lateral view (fig. 67) with

sharp fold at base of caudodorsal beak. Dorsal margin

almost straight, convexly bent into stout caudodorsal

beak. Distal margin making an obtuse angle with

margin of beak, and forming an angularly rounded,

inwardly curved lobe just above lateral protuberance.

Ventral margin angularly bent at half-length. Pygofer

from behind with almost parallel ventral margins; ba-

sal part of pygofer opening broad, U-shaped (fig. 66).

Caudodorsal beak in dorsal view (fig. 69) very broad

and stout, bicuspidate; medially incised at distal mar-

gin. Lateral lobe of pygofer strongly curving inwards

towards end of distal margin, forming a long and

slender, dorsoventrally flattened and upwards curv-

ing, but distinctly posteriorly projecting, almost la-

miniform protuberance. Claspers (fig. 72) very slen-

der, almost laminiform, slightly diverging towards

apices. Dorsal margin straight, curving down at

rounded distodorsal corner and slightly bending in-

wards distally of aedeagus. Apical part shorter and

broader than in P. stoliodes, and bicuspidate; ending

in two, often darkly sclerotized, small thorns.

Aedeagus (fig. 71) angularly upcurved at half-length,

but weakly recurving near apex. Lateral crests slightly

broadening towards proximal ends. A small, weakly

developed collar at apical bending point of aedeagus,

not clearly connected to lateral crests. Distinct dorsal

ridge, abruptly ending in angularly protruding distal

corner. Aedeagal pore oval (fig. 70).

Measurements: Body length d: 20.0-22.5 mm

(x 21.2 mm), 2: 16.4-19.1 mm; tegmen length d:

DE BOER: The genus Papuapsaltria

24.3-29.3 mm (x 26.0 mm), 9 : 28.8-30.8 mm; head

length d: 1.3-1.5 mm, 2: 1.5-1.6 mm; pronotum

length d: 1.9-2.2 mm, 9: 2.0-2.5 mm; mesonotum

length d: 3.7-4.7 mm, 9: 4.8-5.2 mm; head width

3: 3.3-3.7 mm, 2: 3.7-4.1 mm; width of pronotal

collar &: 5.0-6.0 mm, ®: 5.8-6.2 mm.

Distribution (fig. 3). — This species appears re-

stricted to the western part of the Papuan peninsula,

all specimens were collected in the viscinity of the

town of Wau.

Etymology. — Plicata (Latin) means folded and re-

fers to the peculiarly folded pygofer.

Papuapsaltria ungula sp. n.

(figs. 3, 76-82)

Type material. — Holotype à : ‘NEW GUINEA: PAPUA

Kokoda-Pitoki 450 m, iii-24-1956 [print]; ‘J. L.

Gressitt Collector’ [print], BPBM. — Paratype: PAPUA

NEW GUINEA: British New Guinea, 1921, R. Neil

(Rothschild Bequest BM 1939 1), 36, BMNH.

Only two males are available of this species. P. un-

gula is presumably closely related to P. stoliodes, as is

suggested by the shared transparent medial part of the

clasper. The species can be recognized by its claw-

shaped clasper, with a very long and spiny apical part

and a sharp lateral protrusion. P. ungula can further

be recognized by its long and curved caudodorsal

beak.

Description

Body greenish brown. Tegmina 1.1 X as long as

body length. Abdomen 1.5-1.6 X as long as head and

thorax.

Head: Vertex and postclypeus with short red-

brown setae. Postclypeus angularly protruding, ante-

rior margin almost continuous with anterior margins

of vertex lobes. Postclypeus not swollen; anterior

margin (lateral view) concave. Ocelli large and fairly

close together. Distance between lateral ocelli 0.7-

0.8 X as long as distance between lateral ocellus and

eye and 1.4-1.7 X the width of frontal ocellus.

Legs: Fore femur with row of three erect spines.

Most proximal spine fairly short, hardly longer than

middle spine and much shorter than distance to mid-

dle spine.

Tegmina: Hyaline, with 8 apical areas, a narrow

but distinct costal area and a fairly narrow hyaline

border along hind margin.

Tymbals: Seven slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin,

and an 8th, most proximal, ridge spanning only

about 3/4 of tymbal width. Lateral band of seven

intercalary ridges weakly developed.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 76-82. Papuapsaltria ungula sp. n., holotype: 76, py-

gofer from aslant; 77, claspers; 78, pygofer in lateral view;

79, caudodorsal beak in dorsal view; 80, aedeagus in lateral

view; 81, aedeagal apex; 82, operculum.

Operculum (fig. 82): Very small, not covering

tymbal cavity; folded membrane hardly visible in ven-

tral view. Distal part of operculum shorter than basal

part, angularly oblong, and erect, not reaching medi-

ally of meracanthus. Lateral margin short and

straight, concavely curving into crest of basal part and

forming an obtuse angle with long and straight distal

margin. Medial margin straight, distomedial corner

almost rectangular. Meracanthus just reaching anteri-

or margin of abdomen.

Abdomen: Not inflated. First tergite short, not

hidden under metanotum. Medial part of 2nd tergite

only slightly longer than lateral parts. Anterior mar-

gin of 2nd tergite convex dorsally, but weakly concave

medially. First and 2nd sternites almost adjacent.

Auditory capsules strongly inflated.

Genitalia: Pygofer in lateral view as in fig. 78. Dorsal

margin straight at base, but broadly convex, continu-

ous with long and slender, convexly bent caudodorsal

beak. Distal margin angularly convex, forming a dis-

tinct and angular distal lobe on pygofer, and forming

an almost right angle with weakly concave margin of

beak. Ventral margin almost straight. Ventral margins

weakly converging towards base of pygofer; basal part

of pygofer opening broad, V-shaped (fig. 76).

Caudodorsal beak in dorsal view (fig. 79) long, trian-

gular, and rounded at apex. Lateral lobe of pygofer

weakly curving inwards towards end of distal margin,

forming a short and conically shaped, dorsoventrally

flattened, and distinctly posteriorly projecting protu-

berance. Claspers (fig. 77) distinctly bifurcate, claw-

shaped, and slightly diverging towards apices. Dorsal

part of clasper forming a globularly rounded protu-

berance adjacent to aedeagus and supporting aedeagus

in upright position. Apical part of clasper very long and

slender, strongly curved down and sharply pointed.

Medial part of clasper membranous as in P. stoliodes

and P. nana, and almost transparent between apex and

base. Clasper forming a short, thorn-shaped, sharply

pointed and downwards curving lateral protuberance

at the base of its globular dorsal protuberance. This lat-

eral protuberance shorter than the apical part of clasp-

er. Aedeagus (fig. 80) shorter than in most foregoing

species, just reaching beyond clasper, and angularly up-

curved at half-length, but weakly recurving near apex.

Lateral crests very slender, slightly broadening towards

their proximal ends and upcurving at their distal ends.

Crests fused to a small collar around apical bending

point of aedeagus (fig. 81), though connection

between collar and crests very vague. Aedeagus with

rounded dorsal ridge and distinct ventral crest.

Aedeagal pore oval-shaped.

Measurements: Body length: 13.2 & 14.8 mm;

tegmen length: 15.0 & 16.8 mm; head length: 1.0 &

1.1 mm; pronotum length: 1.5 mm; mesonotum

length: 2.7 & 2.9 mm; head width: 2.8 & 2.9 mm;

width of pronotal collar: 3.5 & 3.7 mm.

Distribution (fig. 3). — P. ungula is probably en-

demic to the Papuan peninsula.

Etymology. — The name ungula (Latin claw) refers

to the claw-shaped clasper.

Papuapsaltria nana (Jacobi, 1903) n. comb.

(figs. 3, 6, 83-91)

Baeturia beccarii. — Distant 1892: Tab xiv figs. 27, 27a-b

only (non Distant, 1888).

Acrilla nana Jacobi, 1903: 13, fig. 5.

Baeturia famulus Distant, 1906: 157. — Kato 1931: 74; Kato

1932: 184; Blöte 1958: 266; Metcalf 1963: 245-250;

Duffels & Van der Laan 1985: 252; De Boer 1986: 176;

De Boer 1993a: 16. (syn. n.).

Thaumastopsaltria nana. — Distant 1906: 160; Metcalf

1963: 259; De Boer 1992b: 17-18.

The following references to famulus refer to other

species: Baeturia famulus Stäl MS; Distant 1892: 149

in synonymy of Baeturia beccarii is Muda virguncula,

Baeturia famulus, Myers 1928: fig. 18 is Aedeastria

latifrons (Blöte, 1960), and Baeturia famulus, Lalle-

mand 1931: 78, the specimen indicated is a female

belonging to Baeturia macgillavryi De Boer, 1989.

Lectotype designation. — The type series of Acrilla

nana comes from Milne Bay, but the number of type

specimens is not known (Jacobi 1903). In the

Dresden museum I found 34 and 5? from Milne

Bay from the collection of A. Jacobi; 1d and 19 are

labelled ‘typus’ and one 9 ‘cotypus’. One male in the

BMNH from the same locality and from the Distant

collection, is also labelled ‘typus’. All these specimens

belong to the same species. The male with the ‘typus’

label from the Dresden collection is hereby designat-

ed lectotype, the other specimens with typus/cotypus

labels are regarded as paralectotypes, but the remain-

ing 26 and 39 should possibly also be regarded as

paralectotypes.

Nomenclature and synonymy. — Baeturia famulus

is a manuscript name of Stäl which was regarded as a

synonym of Baeturia beccarii Distant, 1888, when it

was mentioned in Distant’s ‘Monograph of Oriental

Cicadidae’ (1892). Some years later Baeturia beccarii

was transferred to the genus Muda Distant, 1897

(Distant 1906), and brought into the synonymy of

Muda virguncula (Walker, 1857) by Moulton (1923).

Distant’s type of Baeturia beccarii in the BMNH cer-

tainly belongs to the genus Muda. The specimen de-

picted by Distant (1892) as Baeturia beccarii (table

xiv figs 27, 27 a-b) is not a Muda however. Distant re-

alized this and in his ‘Synonymic Catalogue of

Homoptera’ (1906) he presents Baeturia famulus sep-

arately from Muda beccarii. Distant does not give a

description of B. famulus, he merely refers to Stâl MS

and Baeturia beccarii (fig only) in his monograph of

1892.

From that moment the name Baeturia famulus

Distant became valid, with as its description the fig-

ure published in 1892 and the depicted specimen as

its holotype, B. famulus Stäl never having been pub-

lished than as a junior synonym (see Int. code art. 11

e). This depicted specimen was identified by Blöte as

one from Bujakori, S New Guinea, and is now in the

BMNH. Though the specimen does not bear any iden-

tification labels of Distant, it can be recognized by the

9 apical areas in its left tegmen. As Blöte realized, this

specimen could not be the one indicated as famulus

Stäl MS, since it was collected in 1890 and Stäl died

in 1878. Apart from some doubt it sheds on the cor-

rectness of Blöte’s identification of the Bujakori spec-

imen as the one depicted in 1892, this is of no further

consequence.

DE BOER: The genus Papuapsaltria

I here follow Blöte and regard the Bujakori speci-

men as the holotype of Baeturia famulus Distant,

1906. Comparison of types revealed that B. famulus

Dist. is a junior synonym of Thaumastopsaltria

nana (Jacobi, 1903), originally described as Acrilla

nana.

Both Metcalf (1963) and Duffels & Van der Laan

(1985) incorrectly regarded Myers as the author of B.

famulus, originating from an incorrect interpretation

of Distant, 1906. Metcalf falsely stated that Distant

(1906) regarded B. famulus to be a synonym of

Baeturia beccarii.

Myers (1928) mentioned ‘an undoubtedly distinct

species B. famulus Dist.’ and depicted its aedeagus.

Depicted however, is not the holotype of B. famulus

Distant, but a specimen of an altogether different spe-

cies. This specimen could be traced in the BMNH and

is labeled ‘Aru; Wallace; 67/66’ and has on a separate

pin, labeled ‘famulus in pencil in Myers hand, a

preparation of part of its abdomen and pygofer glued

on yellow mika. Though these parts are heavily

crushed, the aedeagus is in good condition and

matches Myers’ drawing. The specimen belongs to

Aedeastria latifrons (Blôte, 1960) which will remain

its valid name, the name B. famulus Myers being a

homonym.

Material examined. — Neu Guinea, Staudinger, 19 det.

Acrilla nana, SMFD; PAPUA NEW GUINEA: Bujakori, viii.1890,

L. Loria, 16 det. beccarii, 34, 12, MCSN; same data 1d

holotype Baeturia famulus Distant, 29, BMNH; Finschhafen,

250 ft, 9.ix.1944, H. Hoogstraal, 2 , NCSU; Menapi, Cape

Vogel peninsula, 0-30 m, 25.iv.1953, G.H.H. Tate, 1d,

AMNH; same data but Geoffrey M. Tate, 2d, AMNH; Milne

Bay, 6 typus Acrilla nana paralectotype, BMNH; Milne Bay,

Coll. A. Jacobi 1911-5, 1d Typus Acrilla nana lectotype,

12 typus Acrilla nana paralectotype, 12 cotypus Acrilla na-

na paralectotype, SMTD; Milne Bay, 2d, 32, (paralect-

otypes?) SMTD; Milne Bay, 12, ZILS; Peria Creek, Kwagira

riv., 50 m, 14.viii-6.ix.1953, Geoffrey M. Tate, 5d, 19,

AMNH; same data 1d, ZMAN; Popondetta, 25 m, v.1966,

Shanahan-Lippert, 8d, 12, BPBM; same data but vi.1966,

56,39, BPBM; same data 2d, 12, ZMAN; Popondetta, 60

m, 1-4.ix.1963, J. Sedlacek, 93, 3, BPBM; same data 24,

12, ZMAN; Popondetta, Buka-Bara, 23.ix.1963, P.

Shanahan, 16, BPBM; Pt. Glasgow, Mamai Plnt., 150 m,

6.11.1965, R. Straatman, 1d, BPBM; without locality label:

Sayer, 1d, 19 det. famulus MS Stal, 19, BMNH:

D’ENTRECASTEAUX ISLANDS: Normanby I., Wakaiuna, Sewa

Bay, 15-30.x.1956, W.W. Brandt, 1d, BPBM; same data but

21-30.xi.1956, 19; 11-20.xii.1956, 12, both BPBM.

P. nana lacks the distinctly convex distal margin of

the pygofer, shared by many of the foregoing species.

The clasper of P. nana, however, strongly suggests a

close relationship with P. ungula. The claspers of

these species share a similar apex and a similarly trans-

parent medial part. However, the clasper of P. nana

has a larger dorsal protrusion and a much smaller lat-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

eral protrusion, the latter forming a bluntly rounded

lobe.

Description

Body yellowish brown or olive green. Females

smaller than males, but with more robust head and

thorax and longer tegmina. Tegmina of males 1.0-

1.2X as long as body length, of females 1.2-1.3 X.

Male abdomen 1.4-1.8 X as long as head and thorax,

of females 1.1-1.4X.

Head (fig. 6): Vertex and postclypeus practically

bald. Postclypeus broad and sharply edged, distinctly

protruding beyond vertex lobes, forming an obtuse

angle anteriorly. Anterior margin of postclypeus

sharply curving back at lateral corners, forming al-

most right angles with anterior margins of vertex

lobes. Postclypeus not swollen; anterior margin (later-

al view) concave. Vertex with weak furrows between

ocelli. Distance between lateral ocelli fairly variable,

0.9-1.3 X as long as distance between lateral ocellus

and eye and 1.6-3.2 X the width of frontal ocellus.

Thorax: Pronotum often with distinct medial furrow.

Legs: Fore femur with row of three erect spines.

Most proximal spine generally distinctly longer than

middle spine, but much shorter than distance to mid-

dle spine.

Tegmina: Generally with 8, occasionally with 9-

10, apical areas, a distinct costal area and a broad hya-

line border along hind margin.

Tymbals: Four slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin,

and a 5th, most proximal, ridge almost reaching ven-

tral margin. Lateral band of four intercalary ridges

weakly developed.

Opercula: Male operculum (fig. 87) very small, not

covering tymbal cavity, leaving a wide gap between

operculum and abdomen; folded membrane com-

pletely visible in ventral view. Distal part of opercu-

lum shorter than basal part, angularly oblong, and

erect, not reaching medially of meracanthus. Lateral

margin short and straight, concavely curving into

crest of basal part and forming an obtuse angle with

long and straight distal margin. Medial margin

straight, distomedial corner almost rectangular.

Meracanthus reaching beyond operculum and just

beyond edge of folded membrane. Female operculum

(fig. 90) very similar to that of males, with angularly

oblong distal part.

DE BOER: The genus Papuapsaltria

Abdomen: Male abdomen distinctly inflated. First

tergite short, not hidden under metanotum. Medial

part of 2nd tergite nearly twice as long as lateral parts.

Anterior margin of 2nd tergite concave medially.

Auditory capsules weakly inflated. First and 2nd ster-

nites not adjacent. Female caudodorsal beak in dorsal

view (fig. 91) narrow triangular and rounded, almost

pointed at apex. Ovipositor sheaths reaching just be-

yond apex of caudodorsal beak.

Male genitalia: Pygofer in lateral view as in fig. 84.

Dorsal margin straight, though concave at base of

caudodorsal beak and angularly bent into short and

straight beak. Distal margin weakly convex, not form-

ing a lobe on pygofer, and concavely bent into margin

of beak. Ventral margin almost straight. Pygofer from

behind with almost parallel ventral margins; basal

part of pygofer opening broad U-shaped (fig. 83).

Caudodorsal beak in dorsal view (fig. 88) short trian-

gular, and rounded or truncate at apex. Lateral lobe of

pygofer gradually curving inwards towards end of dis-

tal margin, forming a fairly large, distinctly swollen

and bluntly rounded lateral protuberance. Claspers

(fig. 89) closely resembling those of P. ungula, with

dorsal part forming a globularly rounded, but larger,

protuberance adjacent to aedeagus, and supporting

aedeagus in upright position. Apical parts of claspers

strongly curved down and slightly diverging towards

apices, forming a long, slender, and sharply pointed

ridge along their convex distal margins. This ridge

arches into a small, lobate lateral protuberance at the

base of clasper. Medial part of clasper membranous

and almost transparent between apex and base.

Aedeagus (fig. 85) weakly upcurved, abruptly narrow-

ing and strongly curved down near apex. Aedeagus

with very slender lateral crests and a distinct pair of

dorsal ridges. Aedeagal pore round (fig. 86).

Measurements: Body length d: 15.5-21.0 mm

(x 17.5 mm + 1.1), 2: 14.9-17.0 mm (x 15.9 mm +

0.7); tegmen length d : 16.5-21.0 mm (x 18.8 mm +

1.0), 2: 18.7-21.0 mm (x 19.8 mm + 0.7); head

length d: 1.5-1.9 mm (x 1.6 mm), 9: 1.5-1.8 mm

(x 1.7 mm); pronotum length d : 1.6-2.2 mm (x 1.8

mm), 2: 1.8-2.2 mm (x 1.9 mm); mesonotum

length d: 3.1-4.1 mm (x 3.4 mm), 9: 3.2-3.9 mm

(x 3.4 mm); head width d : 3.0-3.9 mm (x 3.4 mm),

2: 3.5-4.0 mm (x 3.8 mm); width of pronotal collar

d: 4.0-5.4 mm (X 4.4 mm), 2: 4.5-5.0 mm (x 4.8

mm).

Figs. 83-91. Papuapsaltria nana (Jacobi, 1903): 83, pygofer from aslant, Peria Creek; 84, pygofer in lateral view, Peria Creek;

85, aedeagus in lateral view, Popondetta; 86, aedeagal apex, Popondetta; 87, male operculum Popondetta; 88, male caudo-

dorsal beak in dorsal view, Popondetta; 89, claspers, Peria Creek, arrow indicating membranous medial part; 90, female op-

erculum, Kwagira River; 91, female caudodorsal beak, Kwagira River.

Figs. 92-101. Papuapsaltria bidigitula sp. n.: 92, pygofer in lateral view, holotype; 93, pygofer from aslant, Feramin; 94, ae-

deagus in lateral view, holotype; 95, aedeagal apex in lateral view, holotype; 96, aedeagus in lateral view, Wetar; 97, male cau-

dodorsal beak in dorsal view, holotype; 98, female caudodorsal beak in dorsal view, Wetar; 99, male operculum, holotype;

100, claspers, Wetar; 101, female operculum, Wetar.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Distribution (fig. 3). — P. nana is recorded from the

Huon- and Papuan peninsulas of New Guinea, and

from Normanby Island of the D’Entrecasteaux

Islands.

Papuapsaltria bidigitula sp. n.

(figs. 3, 92-101)

Type material. — Holotype d : ‘NEW GUINEA: (NE)

/ Feramin, 150-120 m / June 15-18, 1959’ [print],

‘W. W. Brandt / Collector / BISHOP’ [print], BPBM. —

Paratypes: IRIAN JAYA: Ansus, A.B. Mayer, 1893, 1d,

BMNH; Jobie, R.H.F. Rippon, N.M.W., 1918.93.

16, NMWC; PAPUA NEW GUINEA: same data as holo-

GPe ENsamendatambuar 23 312195, Peg IE

6.vi.1959, 29; 7-14.vi.1959, 14; all BPBM; Damanti,

Finisterre Mts., Morobe Dist., 2-11.x.1964, M.E.

Bacchus, 1d, 49, BMNH; Eliptamin Valley, 1200-

1350 m, 16-30.viii.1959, W.W. Brandt, 1d, ZMAN;

same data but 19-30.viii.1959, 12, BPBM; WETAR IS-

LAND: Wetter near Timor, W. Doherty, 1903.31,

16,19, BMNH.

P. bidigitula shares a bifurcate clasper with P. plic-

ata, P. stoliodes, and P. ungula. The species is easily

recognized by the shape of its clasper base, forming a

broad, ring-shaped collar, often with two distinct par-

amedian protrusions at the base of the anal valves.

The shapes of the clasper and, especially, of the aedea-

gus are variable, suggesting that the material might

belong to several species.

Description

Body reddish brown. Females smaller than males,

but with more robust head and thorax, and longer

tegmina. Tegmina of males 1.2-1.3 X as long as body

length, of females 1.3-1.5X. Male abdomen 1.2-

1.7X as long as head and thorax, of females 1.1-

110308

Head: Vertex and postclypeus with short red-

brown setae. Postclypeus broad, distinctly protrud-

ing beyond vertex lobes and smoothly rounded ante-

riorly. Anterior margin of postclypeus weakly

convex, but sharply curving back at lateral corners,

forming almost right angles with anterior margins of

vertex lobes. Postclypeus not swollen; anterior mar-

gin (lateral view) concave. Distance between lateral

ocelli 1.0-1.4X as long as distance between lateral

ocellus and eye and 1.8-3.0 X the width of frontal

ocellus.

Legs: Fore femur with row of three spines. Most

proximal spine generally very short and strongly bent,

almost adjacent to femur, but longer and more erect

in specimens from Wetar and Ansus. Proximal spine

much shorter than distance to middle spine.

Tegmina: Hyaline, with 8 apical areas, a distinct

costal area and a fairly broad hyaline border along

hind margin.

Tymbals: Six slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin.

Lateral band of seven intercalary ridges weakly devel-

oped.

Opercula: Male operculum (fig. 99) very small, not

covering tymbal cavity in ventral view and generally

leaving folded membrane exposed, but covering most

of folded membrane in specimen from Ansus. Distal

part of operculum shorter than basal part, angularly

oblong and erect, not reaching medially of meracan-

thus. Lateral margin long and straight, concavely

curving into crest of basal part and forming an obtuse

angle with long and straight distal margin. Medial

margin short and straight, distomedial corner almost

rectangular. Meracanthus reaching beyond opercu-

lum, but not reaching to margin of abdominal seg-

ment 2. Female operculum (fig. 101) very similar to

that of male, with angularly oblong distal part.

Abdomen: Male abdomen hardly inflated. First

tergite short, almost completely hidden under meta-

notum middorsally. Medial part of 2nd tergite slight-

ly less than 2X as long as lateral parts. Anterior mar-

gin of 2nd tergite weakly concave medially. First and

2nd sternites generally not adjacent, but adjacent in

specimen from Ansus. Auditory capsules distinctly

inflated. Female caudodorsal beak in dorsal view (fig.

98) triangular and pointed at apex. Ovipositor

sheaths reaching just beyond apex of caudodorsal

beak.

Male genitalia: Pygofer in lateral view as in fig. 92.

Dorsal margin almost straight and angularly concave

at base of caudodorsal beak. Distal margin weakly

convex, forming an obtuse angle with margin of beak.

Ventral part of pygofer lobe forming an angular cor-

ner under lateral protuberance. Ventral margin

straight. Ventral margins converging to base of pygo-

fer; basal part of pygofer opening broad, V-shaped

(fig. 93). Caudodorsal beak in dorsal view (fig. 97)

short triangular, almost pointed at apex. Lateral lobe

of pygofer weakly curving inwards towards end of dis-

tal margin, but recurving near ventral margin, form-

ing a broad, rectangular and weakly swollen lamini-

form protuberance. Claspers (fig. 100) distinctly

bifurcate, very short and broad, angular in lateral

view, and slightly diverging towards apices. Dorsal

margin straight, curving down at rounded distodorsal

corner. Clasper weakly inflated around aedeagus, and

forming a thorn-shaped and sharply pointed apical

protrusion and a generally more rounded lateral pro-

trusion. In some specimens the lateral protrusion is

the longest, in others the apical protrusion. Basal

parts of claspers fuse to a broad and smoothly round-

ed collar around base of anal valves. This collar gener-

ally with two, almost finger-shaped, paramedian pro-

DE BOER: The genus Papuapsaltria

Figs. 102-109. Papuapsaltria spinigera sp. n. — 102, pygofer in lateral view, Korop; 103, aedeagus in lateral view, paratype;

104, claspers, holotype; 105, pygofer from aslant, holotype; 106, male caudodorsal beak in dorsal view, holotype; 107, male

operculum, holotype; 108, female operculum, Okapa; 109, female caudodorsal beak in dorsal view, Okapa.

Figs. 110-118. Papuapsaltria ustulata (Blôte, 1960). — 110, pygofer from aslant, holotype; 111, male caudodorsal beak in dor-

sal view, holotype; 112, pygofer in lateral view, holotype; 113, aedeagus in lateral view, Kutsime; 114, aedeagal apex, Kutsime;

115, claspers, holotype; 116, male operculum, holotype; 117, female caudodorsal beak in dorsal view, Kutsime; 118, female

operculum, Kutsime.

trusions at its dorsal margin, which are only weakly

developed in the specimens from Damanti and

Feramin, and absent in the specimen from Eliptamin.

Aedeagus quite long, slender, and S-curved, with

broad and angular lateral crests and a distinct dorsal

crest. Aedeagus in the western part of the distribution

area (Ansus, Jobi and Wetar) with slender dorsal crest

and gradually curving down to apex (fig. 96), but in

all other specimens, with more angular dorsal crest

and more angularly bent near apex (fig. 94). Aedeagal

pore weakly incised, almost round (fig. 95).

Measurements: Body length d: 13.1-16.2 mm

LA Dr ER Ben

1.3); tegmen length d: 15.7-20.4 mm (x 19.0 mm +

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

1.9), 2: 15.5-22.4 mm (x 20.6 mm + 2.3); head

length d: 1.1-1.4 mm (x 1.2 mm), 9: 1.1-1.4 mm

(x 1.3 mm); pronotum length d: 1.4-1.7 mm (x 1.6

mm), 9: 1.5-2.0 mm (x 1.8 mm); mesonotum

length d: 2.7-3.8 mm (x 3.2 mm), ?: 3.0-3.5 mm

(x 3.2 mm); head width 6: 3.2-3.4 mm (x 3.3 mm),

9: 3.0-3.8 mm (x 3.5 mm); width of pronotal collar

d:3.6-4.4 mm (X 4.0 mm), 9: 3.7-4.9 mm (X 4.5

mm).

Distribution (fig. 3). — P. bidigitula has a very pe-

culiar distribution. The species is recorded from

northern and central Irian Jaya, Yapen Island, and

from Wetar Island. Since the species has not been re-

corded from any of the intermedate Banda islands,

the record from Wetar is considered dubious.

Etymology. — Digitulus is the diminutive form of

digita (Latin) finger; the name refers to the two small

finger-shaped protuberances on the ring-shaped dor-

sal parts of the claspers.

Papuapsaltria spinigera sp. n.

(figs. 3, 102-109)

Type material. — Holotype d : ‘E Highlands Dist.,

Okapa c 5000 ft. 20.xii.1964 R.H. Hornbrook’ [-

print]; ‘M.E. Bacchus BM 1965-120’ [print]; ‘British

Museum’ [print], BMNH. — Paratypes: PAPUA NEW

GUINEA: same data as holotype 1d, 19, BMNH;

Korop Upper Jimi R. 1300 m, 12.vii.1955, J.L.

Gressitt, 1d, BPBM.

P. spinigera is presumably closely related to the five

foregoing species, sharing a similarly bifurcate clasper.

P. spinigera is distinctly smaller than these species and

is easily recognized by the long lateral protrusion of its

clasper, which is longer than the apical part of clasper.

Description

Body greenish brown. Female smaller than males,

but with more robust head and thorax, and longer

tegmina. Tegmina of males 1.2-1.3 X as long as body

length, of female 1.5 X. Male abdomen 1.2-1.4 X as

long as head and thorax, of female 1.0 x.

Head: Vertex and postclypeus with short red-

brown setae. Postclypeus broad,

distinctly protruding beyond vertex lobes and

smoothly rounded anteriorly. Anterior margin of

postclypeus almost straight, but sharply curving back

at lateral corners, forming almost right angles with

anterior margins of vertex lobes. Postclypeus not

swollen; anterior margin (lateral view) concave.

Diverging fissures on vertex weakly developed.

Distance between lateral ocelli in males 1.0 x, in fe-

males 1.2 x, as long as distance between lateral ocellus

and eye and 1.6-1.8 X the width of frontal ocellus.

Legs: Fore femur with row of three erect spines.

Most proximal spine very short, hardly longer than

middle spine and much shorter than distance to mid-

dle spine.

Tegmina: Hyaline, with 8 long and slender apical

areas, a distinct costal area, and a fairly broad hyaline

border along hind margin.

Tymbals: Seven slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin,

and an 8th, most proximal, ridge spanning only

about 3/4 of tymbal width. Lateral band of seven

intercalary ridges weakly developed.

Opercula: Male operculum (fig. 107) very small, not

covering tymbal cavity, but leaving only a narrow gap

between operculum and abdomen; folded membrane

not visible in ventral view. Distal part of operculum

shorter than basal part, angularly oblong, and erect,

not reaching medially of meracanthus. Lateral margin

short and straight, concavely curving into crest of basal

part and forming an obtuse angle with long and

straight distal margin. Medial margin straight, disto-

medial corner almost rectangular. Meracanthus reach-

ing beyond anterior margin of abdominal segment 2.

Female operculum (fig. 108) very similar to that of

males, with angularly oblong distal part.

Abdomen: Male abdomen not inflated. First ter-

gite very short. Medial part of 2nd tergite only slight-

ly longer than lateral parts. Anterior margin of 2nd

tergite straight medially. First and 2nd sternites adja-

cent. Auditory capsules strongly inflated. Abdomen

of the only female specimen available dried in and

twisted. Female caudodorsal beak in dorsal view (fig.

109) narrow, triangular, and bluntly rounded at apex.

Ovipositor sheaths reaching distinctly beyond apex of

caudodorsal beak.

Male genitalia: Pygofer in lateral view as in fig.

102. Dorsal margin almost straight and almost con-

tinuous with straight margin of short and stout cau-

dodorsal beak. Distal margin angularly convex, form-

ing a distinct distal lobe on pygofer, and concavely

bending into margin of beak. Ventral margin straight.

Pygofer from behind with almost parallel ventral

margins; basal part of pygofer opening broad, U-

shaped (fig. 105). Caudodorsal beak in dorsal view

(fig. 106) short triangular, with concave margins and

rounded apex. Lateral lobe of pygofer weakly curving

inwards towards end of distal margin, forming a

small, lobate, almost laminiform protuberance, pro-

jecting beyond pygofer margin. Claspers (fig. 104)

distinctly bifurcate, very short and broad, angular in

lateral view, and slightly diverging towards apices.

Dorsal margin straight, curving down at rounded dis-

todorsal corner and slightly bending inwards distally

of aedeagus. Apical part of clasper short, thorn-

shaped, directed downwards, with a small inwards di-

rected clasper hollow. Clasper forming a long, spiny,

but apically rounded lateral protuberance near its

base. This lateral protuberance is longer than the api-

cal part of clasper. Clasper base not forming a distinct

collar around base of anal valves. Aedeagus (fig. 103)

very long, reaching far beyond apex of caudodorsal

beak, weakly S-curved, narrowing and strongly curv-

ing down near apex. Lateral crests short and restricted

to proximal half of aedeagus. Aedeagal pore round.

Measurements: Body length d: 11.8-12.7 mm, 2:

11.5 mm; tegmen length d : 14.3-16.2 mm, Q: 16.7

mm; head length d': 1.0-1.1 mm, 2: 1.1 mm; prono-

tum length d: 1.4-1.5 mm, 2: 1.5 mm; mesonotum

length d: 2.6-3.0 mm, 2: 3.0 mm; head width d:

2.7-3.0 mm, 2: 3.1 mm; width of pronotal collar d:

3.3-3.8 mm, 2: 4.0 mm.

Distribution (fig. 3). — P. spinigera is probably re-

stricted to northeastern Papua New Guinea.

Etymology. — Spinigera (Latin) means spiny and

refers to the lateral spine on the clasper.

Papuapsaltria ustulata (Blöte, 1960) comb. n.

(figs. 110-118, 129)

Baeturia ustulata Blôte, 1960: 75, fig. 32; Duffels & Van der

Laan 1985: 255; De Boer 1993a: 16.

P. ustulata is a small species with sharply pointed

pick-shaped claspers, males can further be recognized

by a very short distal part of the operculum and an ex-

tremely long meracanthus.

Description

Body bicolorate: head and thorax ochraceous, tin-

ged with green, abdomen more reddish brown.

Females larger than males. Tegmina of males 1.1-

1.2X as long as body length, of females 1.2 x. Male

abdomen 1.4-1.5 X as long as head and thorax, of fe-

males 1.1-1.4X.

Head: Ochraceous. Postclypeus and anterior part

of vertex lobes with some short red-brown setae.

Postclypeus slightly protruding, almost oblong-

shaped and smoothly rounded anteriorly. Anterior

margin of postclypeus weakly convex, almost contin-

uous with anterior margins of vertex lobes.

Postclypeus not swollen; anterior margin (lateral

view) concave, strongly concave in holotype. Lateral

parts of postclypeus very smooth, with some weak

furrows, but almost without rows of parallel ridges.

Ocelli fairly close together. Distance between lateral

ocelli 1.5-1.8 as long as distance between lateral

ocellus and eye and 0.8-1.1 the width of frontal

ocellus.

Thorax: Pronotum with weak medial furrow.

Mesonotum greenish, cruciform elevation ochra-

ceous.

DE Borr: The genus Papuapsaltria

Legs: Femora ochraceous, tibiae and tarsi reddish.

Fore femur with row of three erect spines. Proximal

spine fairly long in holotype, longer than middle

spine and almost as long as distance to middle spine,

but in other specimens only slightly longer than mid-

dle spine and distinctly shorter than distance to mid-

dle spine.

Tegmina: Hyaline, with 8 apical areas and a dis-

tinct hyaline costal area. Hyaline border along hind

margin of tegmen fairly narrow in holotype, distinct-

ly broader in other specimens.

Tymbals: Holotype with seven transverse sclero-

tized ridges from dorsal to ventral margin and an 8th,

most proximal, ridge almost reaching ventral margin.

Other specimens with one complete ridge less.

Opercula: Male operculum (fig. 116) very short,

but partly covering tymbal cavity in ventral view,

leaving a narrow gap between operculum and abdo-

men; folded membrane hardly visible in ventral view.

Distal part of operculum very short and erect, hardly

more than a continuation of the crest along distolate-

ral corner of basal part, only slightly amplifying me-

siad to less than 1/3 as long as basal part. Lateral mar-

gin convexly bent into straight distal margin. Distal

margin forming a sharp angle with short medial mar-

gin. Meracanthus very long and slender, reaching be-

yond anterior margin of abdominal segment 3.

Female operculum (fig. 118) as in male.

Abdomen: Red-brown, distal part green in holo-

type. One specimen with row of dark ventrolateral

spots on segments 3-7. Male abdomen very slender,

not inflated. First tergite very short and middorsally

almost completely hidden under metanotum. Medial

part of 2nd tergite slightly longer than lateral parts.

Anterior margin of 2nd tergite straight medially. First

and 2nd sternites adjacent. Auditory capsules very

large, bulgy, clearly visible in dorsal view. Female ab-

domen even more slender than in male. Female cau-

dodorsal beak in dorsal view (fig. 117) triangular,

rounded at apex. Ovipositor sheaths just reaching be-

yond apex of caudodorsal beak.

Male genitalia: Pygofer in lateral view as in fig.

112. Dorsal margin weakly concave, convexly bent

into straight and slender caudodorsal beak. Distal

margin weakly convex between caudodorsal beak and

lateral protuberance, concavely bent into margin of

beak, not forming a distal lobe on pygofer.

Caudodorsal beak in dorsal view (fig. 111) very slen-

der, with concave margins, rounded or pointed at

apex. Lateral lobe of pygofer sharply bent inwards, es-

pecially in holotype, forming a large angular, slightly

posteriorly protruding and distinctly swollen protu-

berance. Pygofer forming an angular, in holotype

more rounded, corner just below this protuberance.

Ventral margin of pygofer weakly convex. Ventral

half of pygofer opening narrow U-shaped (fig. 110),

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 119-128. Papuapsaltria baasi sp. n.: 119, pygofer in lateral view, Mokai; 120, pygofer from aslant, Mokai; 121, male

caudodorsal beak in dorsal view, Tor River; 122, male caudodorsal beak in dorsal view, Mokai; 123, claspers, Mokai; 124, ae-

deagal apex, Mokai; 125, aedeagus in lateral view, Bodem; 126, male operculum, Mokai; 127, female operculum, Bodem;

128, female caudodorsal beak in dorsal view, Bodem.

with weakly concave basal margin; ventral margins al-

most parallel. Claspers (fig.115 ) very short, hardly

reaching beyond pygofer margin, and weakly diverg-

ing towards apices; pick-shaped, with rectangular

square-shaped dorsal part and slender, slightly recurv-

ing, sharply pointed, and downwards directed apical

part. Dorsal part of clasper curving mesiad, as a shaft

encircling aedeagus, and supporting aedeagus in up-

right position. Apical part of clasper with large and

sharply edged clasper hollow. Basal parts of claspers

curving inwards, but not fusing, forming an almost

continuous collar around base of anal valves.

Aedeagus standing erect between claspers, adjacent to

anal valves. Aedeagus (fig. 113) weakly S-curved, al-

most straight, but angularly bent near apex, with dis-

tinct lateral crests and a weak dorsal ridge. Apical part

of aedeagus abruptly broadening, and forming a

small and angular protrusion at apical bending point.

Aedeagal pore oval, almost round (fig. 114).

Measurements: Body length d: 13.2-13.7 mm, 2:

14.5 & 16.4 mm; tegmen length d: 15.1-16.4 mm,

2: 19.9 mm; head length d: 1.0-1.2 mm, 2: 1.3 &

1.5 mm; pronotum length d: 1.5-1.6 mm, 9: 1.9

mm; mesonotum length d: 2.8-3.0 mm, 2: 3.4 &

3.6 mm; head width d: 2.9-3.3 mm, 9: 3.4 & 3.7

mm; width of pronotal collar d : 3.6-3.9 mm, 9: 4.7

& 4.8 mm.

Material examined. — IRIAN JAYA: Bernhard Camp, 100

m, 10.iv.1939, Neth Ind. — American New Guinea

Expedit., LJ. Toxopeus, & holotype Baeturia ustulata,

RMNH; Karubaka, Swart vall., 1300 m, 1.x1.1958, J.L.

È abaasi 5

À i mdolabrata ||0

{ Ze a vnovariae

= © SS eustulata

LS a SEZ}

1322

136° 140°

Fig. 129. Localities of Papuapsaltria baasi, P. dolabrata, P.

novariae, and P. ustulata.

Gressitt, 24, BPBM; Kutsime, West of Swart vall., 1500 m,

14.x1.1958, J.L. Gressitt, 22, BPBM.

Distribution (fig. 129). — P. ustulata is endemic to

central western New Guinea, north of the central

mountain ranges.

Papuapsaltria baasi sp. n.

(figs. 119-129)

Type material. — Holotype d : ‘NEW GUINEA NETH.

/ Bodem, 100 m, 11 km / SE Oerberfaren / July 7-17,

1959 [print]; “T.C. Maa / Collector / Bishop’ [print],

BPBM. — Paratypes: IRIAN JAYA: same data as holotype

18,49, BPBM; same data 1d, 19, ZMAN; Tor river

(mouth), 4 km E of Hol Maffen, 1.vii.1959, T.C.

Maa, 16, BPBM.

Other material. — PAPUA NEW GUINEA: Mokai vill.,

Torricelli Mts., 700 m, 16-31.xii.1958, W.W.

Brandt, 1d, BPBM; same data but 750 m, 1-

23.1.1959, 16, BPBM.

P. baasi closely resembles P. ustulata and is of about

the same size, but has distinctly larger opercula. Males

can be easily separated from P. ustulata by the clasp-

ers, which are fused to a high and smoothly rounded

collar around the base of anal valves. A similar collar

is found in P. novariae and P. dolabrata, which are de-

scribed below.

Description

Body reddish brown. Females of about the same si-

ze as males, with relatively slightly longer tegmina and

a more robust head and thorax. Tegmina of males

1.1-1.3X as long as body length, of females 1.2-

1.3 X. Male abdomen 1.2-1.5 X as long as head and

thorax, of females 1.1-1.4X.

Head: Reddish brown, with short red-brown setae.

Postclypeus oblong, distinctly protruding beyond

vertex lobes and smoothly rounded anteriorly.

Anterior margin of postclypeus weakly convex, but

sharply curving back at lateral corners, forming al-

most right angles with anterior margins of vertex

lobes. Dorsal surface of postclypeus weakly undulat-

ed; laterally slightly indented. Postclypeus not swol-

len; anterior margin (lateral view) concave. Lateral

parts of postclypeus very smooth, with some weak

furrows, but almost without rows of parallel ridges

along lorum. Distance between lateral ocelli 1.0-

1.2 X as long as distance between lateral ocellus and

eye and 2.1-2.8 X the width of frontal ocellus.

Thorax: Pronotum with weak medial furrow.

Mesonotum greenish, cruciform elevation ochra-

ceous.

Legs: Femora ochraceous, tibiae and tarsi reddish.

Fore femur with row of three erect spines. Proximal

De BOER: The genus Papuapsaltria

spine fairly long, often slightly longer than middle

spine, but much shorter than distance to middle

spine.

Tegmina: Hyaline, with 8 apical areas and a very

narrow costal area; specimen from Tor River with 7

apical areas in right tegmen. Hyaline border along

hind margin of tegmen fairly narrow as in holotype of

P. ustulata, but distinctly broader in one female.

Tymbals: Five transverse sclerotized ridges span-

ning the tymbal from dorsal to ventral margin and a

6th, most proximal, ridge almost reaching ventral

margin. Intercalary ridges weakly developed.

Opercula: Male operculum (fig. 126) small, but

distinctly longer than in P. ustulata, only partly cover-

ing tymbal cavity in ventral view, leaving a narrow

gap between operculum and abdomen; folded mem-

brane hardly visible in ventral view. Distal part of op-

erculum almost as long as basal part. Lateral margin

long and straight, forming an obtuse angle with crest

of basal part and with straight distal margin. Medial

margin weakly convex; distomedial corner rounded.

Meracanthus not reaching to anterior margin of ab-

domen. Female operculum (fig. 127) as in male, but

with shorter distal part.

Abdomen: Red-brown. Male abdomen very slen-

der, not inflated. First tergite very short, almost com-

pletely hidden under metanotum middorsally.

Medial part of 2nd tergite less than 1.5 X as long as

lateral parts. Anterior margin of 2nd tergite weakly

concave medially. First and 2nd sternites adjacent.

Auditory capsules very large, bulgy, clearly visible in

dorsal view. Female abdomen even more slender than

in male. Female caudodorsal beak in dorsal view (fig.

128) triangular and pointed at apex. Ovipositor

sheaths just reaching beyond apex of caudodorsal

beak.

Male genitalia: Pygofer in lateral view as in fig.

119. Dorsal margin weakly concave to base and angu-

larly bent into slender caudodorsal beak. Caudodorsal

beak straight in type specimens, but recurved in

Mokai specimens. Distal margin weakly convex

between caudodorsal beak and lateral protuberance,

but not forming a distal lobe on pygofer; and con-

cavely, in Mokai specimens angularly, bent into mar-

gin of beak. Caudodorsal beak in dorsal view apically

rounded, broadly triangular in type specimens (fig.

121), very slender and with a sharp medial ridge at its

base in Mokai specimens (fig. 122). Lateral lobe of

pygofer slightly curved inwards, but recurved towards

ventral margin, forming a broad, rounded, lamini-

form protuberance. Pygofer of specimen from Tor

River forming an angular corner just below this pro-

tuberance, such a corner is absent in other specimens.

Ventral margin of pygofer almost straight. Ventral

half of pygofer opening narrow V-shaped (fig. 120);

ventral margins converging to a sharp angle at the

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

<® 146

Figs. 130-136. Papuapsaltria novariae sp. n. — 130, aedeagus in lateral view, holotype; 131, aedeagal apex, holotype; 132, py-

gofer from aslant, holotype; 133, claspers, holotype; 134, pygofer in lateral view, holotype; 135, caudodorsal beak in dorsal

view, holotype; 136, operculum, paratype.

Figs. 138-146. Papuapsaltria dolabrata sp. n. — 138, pygofer from aslant, holotype; 139, pygofer in lateral view, holotype; 140,

aedeagus in lateral view, Etna Bay; 141, aedeagal apex, Etna Bay; 142, male caudodorsal beak in dorsal view, holotype; 143,

female caudodorsal beak in dorsal view, Roon; 144, claspers, holotype; 145, male operculum, paratype Waigeu; 146, female

operculum, Roon.

base of pygofer opening. Claspers (fig. 123) very shaped, with rectangular square-shaped dorsal part

short as in P. ustulata, hardly reaching beyond pygof- and slender, downwards directed, slightly recurving

er margin, and slightly diverging towards apices; pick- and sharply pointed apical part. Dorsal part of clasp-

DE BOER: The genus Papuapsaltria

Fig. 137. Papuapsaltria dolabrata sp. n.: left tegmen, male, Waigeu .

er curving mesiad, as a shaft encircling aedeagus, and

supporting aedeagus in upright position. Apical part

of clasper broader than in P. ustulata, with large and

sharply edged clasper hollow. Basal parts of claspers

fused at distinct medial ridge, forming a high and

smoothly rounded continuous collar around base of

anal valves. Aedeagus standing erect between claspers,

adjacent to anal valves. Aedeagus (fig. 125) almost

straight, but strongly curved down, and abruptly nar-

rowing near slightly incised apex (fig. 124), with slen-

der lateral crests and a weak dorsal ridge. Aedeagus in

one specimen from Mokai distinctly broader than in

other specimens.

Measurements: Body length d: 11.7-14.5 mm

(rl Otmm 019); OE 121137 mm & 12:9 mm

+ 0.5); tegmen length d: 13.7-17.5 mm (X 15.5

mm + 1.4), 2: 15.6-16.9 mm (x 16.3 mm + 0.6);

head length d: 1.1-1.3 mm (x 1.2 mm), 9:1.2-1.5

mm (x 1.4 mm); pronotum length d: 1.4-1.6 mm

(x 1.5 mm), 9: 1.5-1.7 mm (x 1.6 mm); mesono-

tum length G : 2.3-3.2 mm (x 2.8 mm), 9: 2.6-3.0

mm (x 2.8 mm); head width &: 2.8-3.3 mm (x 3.0

mm), 2: 3.0-3.3 mm (x 3.2 mm); width of prono-

tal collar d: 3.4-4.1 mm (& 3.7 mm), 9: 3.5-4.1

mm (x 3.8 mm).

Distribution (fig. 129). — P. baasi is endemic to the

mountain ranges of northern New Guinea.

Etymology. — This species is named in honour

of Hendrik Baas in acknowledgement of our long

discussions on the paleogeography of New Gui-

nea.

Papuapsaltria novariae sp. n.

(figs. 129-136)

Type material. — Holotype d : ‘NEW GUINEA NE /

Eliptamin Valley / 1200-1350 m / July 16-31, 1959

[print]; “W.W. Brandt / Collector / Bishop’ [print],

BPBM. — Paratype: IRIAN JAYA: same data as holotype

but 1665-2530 m, 9.vi.1959, 1d, BPBM.

Only two males are known of this species. P. no-

variae closely resembles P. baasi in the shape of the

genitalia: its clasper base forming a similarly high col-

lar around the base of the anal valves. The dorsal cor-

ner of the P. novariae clasper is more rounded, how-

ever. P. novariae is distinctly larger than the two

foregoing species and shows a much wider gap

between operculum and abdomen. The species shares

the latter character with P. dolabrata described next.

P. novariae can further be recognized by its 9 apical

areas in the tegmina.

Description

Body reddish brown. Tegmina 1.2-1.3 X as long as

body length. Abdomen 1.4-1.6 X as long as head and

thorax.

Head: Reddish brown, with short red-brown setae.

Postclypeus oblong, slightly protruding beyond ver-

tex lobes and smoothly rounded anteriorly. Anterior

margin of postclypeus weakly convex, but sharply

curving back at lateral corners, forming almost right

angles with anterior margins of vertex lobes.

Postclypeus not swollen; anterior margin (lateral

view) straight. Lateral parts of postclypeus very

smooth, with some weak furrows, but almost without

rows of parallel ridges along lorum. Distance between

lateral ocelli slightly longer than distance between lat-

eral ocellus and eye and 1.7-1.9 X the width of fron-

tal ocellus.

Legs: Femora ochraceous, tibiae and tarsi reddish.

Fore femur with row of three erect spines. Proximal

spine very short, slightly longer than middle spine,

but much shorter than distance to middle spine.

Tegmina: Hyaline, with 9 apical areas, a distinct

and hyaline costal area, and a fairly narrow hyaline

border along hind margin.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Tymbals: Five transverse sclerotized ridges span-

ning the tymbal from dorsal to ventral margin and a

6th, most proximal, ridge almost reaching ventral

margin. Intercalary ridges hardly visible.

Operculum (fig. 136): Very short. Distal part more

than half as long as basal part, not covering tymbal

cavity, and leaving a very wide gap between opercu-

lum and abdomen; folded membrane clearly visible in

ventral view. Lateral margin long and straight, form-

ing an obtuse angle with crest of basal part and angu-

larly or convexly bending into straight distal margin.

Medial margin straight, forming a sharp angle with

distal margin. Meracanthus reaching to about half-

way the gap between operculum and abdomen.

Abdomen: Red-brown, more inflated than in the

two foregoing species. First tergite very short, almost

completely hidden under metanotum middorsally.

Medial part of 2nd tergite almost 2 X as long as later-

al parts. Anterior margin of 2nd tergite distinctly con-

cave medially. First and 2nd sternites not adjacent.

Auditory capsules weakly swollen and not visible in

dorsal view.

Genitalia: Pygofer in lateral view as in fig. 134.

Dorsal margin concave, but convexly, almost angu-

larly, bent into straight and slender caudodorsal beak.

Distal margin weakly convex between caudodorsal

beak and lateral protuberance, but not forming a dis-

tal lobe on pygofer, and concavely bent into margin

of beak. Caudodorsal beak in dorsal view (fig. 135)

broadly triangular and bluntly rounded at apex.

Lateral lobe of pygofer slightly curved inwards, but

recurved towards ventral margin, forming a broad,

rounded, and laminiform protuberance. Pygofer

forming a small rectangular corner just below this

protuberance. Ventral margin of pygofer almost

straight. Ventral half of pygofer opening narrow V-

shaped (fig. 132); ventral margins converge to a sharp

angle at base of pygofer opening. Claspers (fig. 133)

closely resembling those of P. baasi, but almost par-

allel, only slightly diverging near apices. Claspers

hardly reaching beyond pygofer margin, their basal

parts fused at distinct medial ridge, forming a high

and smoothly rounded continuous collar around base

of anal valves. Dorsal part of clasper more rounded

than in P. baasi, curving mesiad, as a shaft encircling

aedeagus. Apical part of clasper recurving, directed

downwards and sharply pointed, with a large and

sharply edged clasper hollow. Aedeagus standing erect

between claspers, adjacent to anal valves, in holotype

reaching far above claspers, in paratype just reaching

above dorsal margin of clasper. Aedeagus (fig. 130) al-

most straight, but strongly, almost angularly, curved

down towards its distinctly incised apex (fig. 131).

Aedeagus with slender lateral crests and a weak dorsal

ridge.

Measurements: Body length: 16.0 & 16.4 mm;

tegmen length: 19.3 & 21.1 mm; head length: 1.2

mm; pronotum length: 1.7 & 1.8 mm; mesonotum

length: 3.4 & 3.8 mm; head width: 3.3 & 3.5 mm;

width of pronotal collar: 4.3 & 4.4 mm.

Distribution (fig. 129). — P. novariae is only known

from Eliptamin valley, in the central mountain rang-

es of western Papua New Guinea.

Etymology. — The name novariae is derived from

the latin words novem (nine) and ariae (areas) and re-

fers to the nine apical areas of tegmen.

Papuapsaltria dolabrata sp. n.

(figs. 129, 137-146)

Type material. — Holotype d : ‘INDONESIA / Irian

Jaya / A.J. de Boer / A.L.M. Rutten & / R. de Vos’

[print]; © Roon Island / YENDE / 60 m. at light /

7.x1.1993 [print], ZMAN. — Paratypes: IRIAN JAYA:

same data as holotype 26, ZMAN; Etna baai,

25.xi.1939, Nieuw Guinea Exp., KNAG 1939, 2d,

RMNH; Roon, Fruhstorfer, 16, 19, BMNH; Waigeu,

Camp Nok, 2500 ft., iv.1938, L.E. Cheesman, 36,

59, BMNH; same data 1d, 19, ZMAN; same data but

v.1938, 39.

P. dolabrata closely resembles P. baasi in size and

shape of body, but shares a wide gap between opercu-

lum and abdomen with P. novariae. P. dolabrata is

easily separated from these two species by its sharply

downwards directed dorsal margin of the claspers; the

distal margin of the clasper is almost continuous with

the ring-shaped clasper base (lateral view). P. dolabra-

ta has an extremely narrow hind margin of tegmen.

Description

Body olive green to light brown. Females smaller

than males, but with more robust head and thorax,

and longer tegmina. Tegmina of males 1.1-1.2X as

long as body length, of females 1.3-1.4X. Male ab-

Figs. 147-155. Papuapsaltria toxopei sp. n.: 147, pygofer in lateral view; 148, pygofer from aslant; 149, aedeagus in lateral

view; 150, aedeagal apex; 151, male caudodorsal beak in dorsal view; 152, claspers; 153, male operculum; 154, female oper-

culum; 155, female caudodorsal beak in dorsal view.

Figs. 156-164. Papuapsaltria dioedes sp. n.: 156, pygofer from aslant, holotype; 157, pygofer in lateral view, holotype; 158,

aedeagus in lateral view, holotype; 159, male fore femur, Pengagl; 160, male caudodorsal beak in dorsal view, holotype; 161,

female caudodorsal beak in dorsal view, Pengagl; 162, claspers, holotype; 163, male operculum, holotype; 164, female oper-

culum, Pengagl.

DE BOER: The genus Papuapsaltria

“TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

domen 1.4-1.7 X as long as head and thorax, of fe-

males 1.1-1.5 X.

Head: Olive green, with short red-brown setae.

Postclypeus oblong, distinctly protruding beyond

vertex lobes and smoothly rounded anteriorly.

Anterior margin of postclypeus weakly convex, but

sharply curving back at lateral corners, forming al-

most right angles with anterior margins of vertex

lobes. Dorsal surface of postclypeus weakly undulat-

ed; laterally slightly indented. Postclypeus not swol-

len; anterior margin (lateral view) concave. Lateral

parts of postclypeus very smooth, with some weak

furrows, but almost without rows of parallel ridges

along lorum. Distance between lateral ocelli 1.0-

1.2 X as long as distance between lateral ocellus and

eye and 1.7-2.6 X the width of frontal ocellus.

Legs: Femora ochraceous, tibiae and tarsi reddish.

Fore femur with row of three erect spines. Proximal

spine fairly long, slightly longer than middle spine,

but much shorter than distance to middle spine.

Tegmina: Hyaline, with 8 apical areas and distinct

and hyaline costal area. Venation often dark red-

brown. Hyaline border along hind margin of tegmen

extremely narrow, hardly visible.

Tymbals: Five transverse sclerotized ridges span-

ning the tymbal from dorsal to ventral margin and a

6th, most proximal, ridge almost reaching ventral

margin. Intercalary ridges weakly developed.

Operculum: Male operculum (fig. 145) not cover-

ing tymbal cavity, leaving a very wide gap between

operculum and abdomen; folded membane clearly

visible in ventral view. Distal part of operculum short,

less than half as long as basal part. Lateral margin

straight, forming an obtuse angle with crest of basal

part and angularly or convexly bent into straight dis-

tal margin. Medial margin straight, forming a nar-

rowly rounded corner with distal margin.

Meracanthus not reaching beyond folded membrane.

Female operculum (fig. 146) as in male, but much

shorter. Distal part as in P. ustulata, erect and hardly

more than a continuation of the crest along distolate-

ral corner of basal part, and only slightly amplifying

mesiad to less than 1/5 X as long as basal part.

Abdomen: Male abdomen hardly inflated. First

tergite very short, almost completely hidden under

metanotum middorsally. Medial part of 2nd tergite

about 2 X as long as lateral parts. Anterior margin of

2nd tergite weakly concave medially. First and 2nd

sternites not adjacent. Auditory capsules weakly swol-

len, and hardly or not visible in dorsal view. Female

caudodorsal beak in dorsal view (fig. 143) broadly tri-

angular and rounded, almost pointed, at apex.

Ovipositor sheaths almost reaching to apex of caudo-

dorsal beak.

Male genitalia: Pygofer in lateral view as in fig.

139. Dorsal margin straight, continuous with, or

concavely bent into, slender caudodorsal beak. Distal

margin weakly convex between caudodorsal beak and

lateral protuberance, and concavely bent into margin

of beak. Caudodorsal beak in dorsal view (fig. 142)

broad and triangular, almost rectangular at apex.

Lateral lobe of pygofer slightly curved inwards, but

recurving towards ventral margin, forming a very

large, weakly indented, and lobate laminiform protu-

berance, almost triangular in lateral view. Pygofer

forming a small but distinct rectangular corner just

below this protuberance. Ventral margin of pygofer

straight. Ventral half of pygofer opening narrow V-

shaped (fig. 138), ventral margins converging to a

sharp angle at base of pygofer opening. Claspers quite

different from the three foregoing species. Whereas in

these three species the dorsal margin of the clasper

part, which bends around aedeagus, ends in almost

right angle on the fused ring-shaped clasper base (see

fig. 144), in P. dolabrata this margin merges with the

basal ring under a very slight angle (fig. 152).

Moreover, the apical part of clasper is less strongly

bent down than in the three foregoing species.

Claspers pick axe-shaped almost parallel, only slightly

diverging near apices. Apical part of clasper recurved

and sharply pointed, with a large and sharply edged

clasper hollow. Claspers closely encircling aedeagus at

their bending point. Aedeagus, unlike the three fore-

going species, not erect between claspers, but slightly

directed posteriad. Aedeagus (fig. 149) weakly S-

curved, with slender lateral crests and a very distinct,

rounded dorsal ridge. Aedeagus weakly incised at

apex; aedeagal pore almost round (fig. 150).

Measurements: Body length d: 14.1-14.9 mm (x

14.4 mm = 0:3); SETA rn Ge 1228 imma

0.6); tegmen length d: 16.1-17.2 mm (x 16.8 mm +

0.5), 2: 16.0-18.6 mm (x 17.1 mm + 0.8); head

length d: 1.2-1.5 mm (x 1.3 mm), 9: 1.2-1.6 mm

(x 1.4 mm); pronotum length d': 1.5-1.7 mm (x 1.6

mm), 9: 1.4-1.8 mm (x 1.7 mm); mesonotum length

di: 2.7-3.1 mm & 2.9 mm); 9:2732 mm (& 2:9

mm); head width 6: 3.1-3.3 mm (x 3.2 mm), 2:

3.1-3.5 mm (x 3.3 mm); width of pronotal collar d:

3.7-4.1 mm (x 3.9 mm), 2: 3.8-4.4 mm (x 4.1 mm).

Distribution (fig. 129). — P. dolabrata is distribut-

ed in western New Guinea, recorded from Etna Bay,

Roon Island, and Waigeu Island, but not from the

intermediate Birds Head peninsula.

Etymology. — Dolabrata means shaped as a dolabra

(Latin) pickaxe. The name refers to the shape of the

claspers.

Papuapsaltria toxopei sp. n.

(figs. 4, 147-155)

Type material. — Holotype: ‘Neth. Ind.-American /

New Guinea Exped. / Mist Camp 1800 m /

23.1.1939 LJ. Toxopeus’ [print]; ‘Paratype’ (print,

orange label]; ‘Baeturia | toxopeusî [written]; ‘Mus.

Leiden’ [print], d, RMNH. Paratype: IRIAN JAYA: same

data as holotype but 19.1.1939 and without identifi-

cation label, 19 paratype B. toxopeusi, RMNH.

P. toxopei presumably belongs to this genus, as is

indicated by the small male opercula and short femo-

ral spines, but its phylogenetic position within the ge-

nus is not clear. P. toxopei is very similar, and presum-

ably closely related, to the four foregoing species. The

hook-shaped and sharply pointed claspers of P. toxo-

pei somewhat resemble those of P. woodlarkensis, but

similar claspers occur in several related genera.

Description

Body reddish brown, mesonotum greenish. Female

smaller than male, but with more robust head and

thorax, and longer tegmina. Tegmina of male 1.3 X

as long as body length, of female 1.5 X. Male abdo-

men 1.5X as long as head and thorax, of female

Head: Vertex and postclypeus with short red se-

tae. Postclypeus angularly protruding. Anterior mar-

gin of postclypeus almost continuous with anterior

margins of vertex lobes. Postclypeus not swollen; an-

terior margin (lateral view) concave. Vertex with

fairly distinct diverging fissures. Distance between

lateral ocelli 0.9-1.0 X as long as distance between

lateral ocellus and eye and 1.5-1.9 X the width of

frontal ocellus.

Legs: Fore femur with row of three erect spines.

Most proximal spine very short, slightly longer than

middle spine, but much shorter than distance to mid-

dle spine.

Tegmina: Hyaline, with 8 long and slender apical

areas, a distinct hyaline costal area, and a broad hya-

line border along hind margin.

Tymbals: Six slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin,

and a 7th, most proximal, ridge almost reaching ven-

tral margin. Lateral band of seven intercalary ridges

weakly developed.

Opercula: Male operculum (fig. 153) very small,

not covering tymbal cavity in ventral view, leaving a

wide gap between operculum and abdomen; folded

membrane exposed in ventral view. Distal part of op-

erculum narrow, and angularly oblong, shorter than

basal part. Operculum not reaching medially of mer-

acanthus. Lateral margin very long and straight, grad-

ually curving into crest of basal part and forming an

obtuse angle with short and straight distal margin.

Medial margin straight, distomedial corner angularly

rounded. Meracanthus very long, reaching beyond

anterior margin of abdomen. Female operculum (fig.

De BOER: The genus Papuapsaltria

154) very similar to that of male, but with shorter and

broader distal part.

Abdomen: Male abdomen long and slender, weak-

ly inflated. First tergite not hidden under metano-

tum. Medial part of 2nd tergite less than 2 X as long

as lateral parts. Anterior margin of 2nd tergite straight

medially. First and 2nd sternites not adjacent.

Auditory capsules strongly inflated and protruding,

visible in dorsal view. Female caudodorsal beak in

dorsal view (fig. 155) long and slender, rounded at

apex. Ovipositor sheaths not reaching to apex of cau-

dodorsal beak.

Male genitalia: Pygofer in lateral view as in fig.

147. Dorsal margin convex, concave near base, and

continuous with convexly bent caudodorsal beak.

Distal margin weakly convex, forming an obtuse an-

gle with concave margin of beak. Ventral margin

straight. Ventral margins weakly converging to base

of pygofer; basal part of pygofer opening V-shaped

(fig. 148). Caudodorsal beak in dorsal view (fig. 151)

slender, triangular, narrowly rounded at apex. Lateral

lobe of pygofer strongly curving inwards towards end

of distal margin, forming a bluntly rounded and dis-

tinctly swollen protuberance. Claspers (fig. 152)

hook-shaped and parallel, only slightly diverging near

sharply pointed apices. Dorsal part of clasper forming

an angularly protruding corner, bending around ae-

deagus. Apical part of clasper smoothly rounded, lo-

bate, with large and sharply edged clasper hollow.

Aedeagus (fig. 149) weakly S-curved, strongly curved

down near apex, with long and slender lateral crests

and distinct dorsal ridge. Lateral crests curving up-

wards at their distal ends and fusing to a very small

collar at apical bending point of aedeagus. Aedeagal

pore round.

Measurements: Body length d: 19.4 mm, 2: 17.2

mm; tegmen length d: 23.5 mm, 2: 25.0 mm; pro-

notum length d: 1.7 mm, 2: 2.1 mm; mesonotum

length d: 3.8 mm, 9: 3.0 mm; head length d: 1.2

mm, ®: 1.5 mm; head width 6: 3.4 mm, 9: 3.9

mm; width of pronotal collar d: 4.8 mm, 2: 5.7

mm.

Distribution (fig. 4). — P. toxopei is known from

only one locality in northern Irian Jaya.

Etymology. — This species is named in honour of

Mr L.J. Toxopeus who collected the holotype, and

the types of several other species on which the current

revision of the ‘ Baeturia and related genera complex’

is based.

Papuapsaltria dioedes sp. n.

(figs. 4, 156-164)

Type material. — Holotype d : ‘New Guinea: NE /

Mur Mur P. 2800 m / 28.xii.67-6.1.68 [print]; “Tawi

/ Collector / BISHOP’ [print], BPBM; Paratypes: PAPUA

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 165-174. Papuapsaltria woodlarkensis sp. n.: 165, female genital segment in lateral view, paratype; 166, pygofer in later-

al view, holotype; 167, pygofer from aslant, holotype; 168, aedeagal apex, paratype; 169, male caudodorsal beak in dorsal

view, holotype; 170, aedeagus in lateral view, paratype; 171, claspers, holotype; 172, male operculum, holotype; 173, female

operculum, paratype; 174, female caudodorsal beak in dorsal view, paratype.

NEW GUINEA: 5°15’ S/ 141°05’ E, 2280 m, 4.v.1970,

O. McCaw, 1d, Moul; Bosavi Mt. 2300 m,

7.x.1973, J.L. Gressitt & Goya, 1d, BPBM; Giluve,

2700 m, J. & M. Sedlacek, 16, ZMAN; Giluve Mt.,

2500-2650 m, 28.v.1963, J. Sedlacek, 16, BPBM;

same data but 2500-2750 m, 30.v.1963, 19, BPBM;

Kepilam, 2420-2540 m, 21.vi.1963, J. Sedlacek, 19,

BPBM; same data but 2450-2600 m, 22.vi.1963, 19,

ZMAN; Kiunga, 5°15’ S / 141°05° E, 7500 ft,

4.v.1970, O. McCaw, 16, Moul; Kubor Range, W

Highlands, 2950 m, 23.v.1966, J.L. Gressitt, 1d,

BPBM; Wilhelm Mt., east slopes, Pengagl Camp, 2770

m, 17.vii.1959, sixth Archbold Exped. to Papua New

Guinea, L.J. Brass, 16, AMNH; same data but

26.vii.1959, 22, AMNH.

The generic allocation of this species is somewhat

uncertain. P. dioedes shares the supposed synapomor-

phous very short proximal spine on the fore femur

with most species of Papuapsaltria, while the tegmina,

with very long and slender apical areas and a very

broad hyaline border, also indicate a relation with this

genus. For these reasons the species is preliminary in-

cluded in Papuapsaltria. However, its claspers are very

similar to those of Guineapsaltria pennyi. The male

opercula somewhat resemble those of Guineapsaltria

chinai and those of P. goniodes and P. angulata.

Description

Body light yellowish brown to reddish brown.

Females smaller than males, but with more robust

head and thorax, and longer tegmina. Tegmina of

males 1.1-1.4X as long as body length, of females

1.5-1.6X. Male abdomen 1.4-2.0X as long as head

and thorax, of females 1.1-1.4X.

Head: Vertex and postclypeus with short red setae.

Postclypeus angularly protruding beyond vertex

lobes. Anterior margin of postclypeus sharply curving

back at lateral corners, forming almost right angles

with anterior margins of vertex lobes. Postclypeus not

swollen; anterior margin (lateral view) concave.

Vertex with fairly distinct diverging fissures. Distance

between lateral ocelli 0.9-1.3X as long as distance

between lateral ocellus and eye and 1.4-2.5X the

width of frontal ocellus.

Legs: Fore femur very slender (fig. 159) with row

of three erect spines. Most proximal spine extremely

short, rudimentary, often shorter than middle spine

and much shorter than distance to middle spine.

Tegmina: Hyaline, with 8 long and slender apical

areas, a distinct hyaline costal area and a very broad

hyaline border along hind margin.

Tymbals: Six slightly sclerotized parallel ridges

spanning the tymbal from dorsal to ventral margin.

Lateral band of seven intercalary ridges weakly devel-

oped.

Opercula: Male operculum (fig. 163) quite large,

but only partly covering tymbal cavity and leaving a

wide gap between operculum and abdomen; folded

membrane only partly visible in ventral view. Distal

part of operculum fairly long, more than twice as long

as basal part, and medially elongate, with greatest

length at medial margin. Operculum not reaching

medially of meracanthus. Lateral margin long and

straight, forming an obtuse angle with crest of basal

part, close to distolateral corner of basal part, and

convexly bent into much shorter, straight distal mar-

gin. Medial margin long and straight, distomedial

corner almost rectangular. Meracanthus reaching be-

yond operculum. Female operculum (fig. 164) very

short, its distal part shorter than basal part and angu-

larly oblong. Distolateral corner rounded, distomedi-

al corner rectangular, medial margin very short.

Abdomen: Male abdomen strongly inflated. First

tergite very short, partly hidden under metanotum.

Medial part of 2nd tergite about twice as long as lat-

eral parts. Anterior margin of 2nd tergite strongly

concave medially. First and 2nd sternites generally

not adjacent. Auditory capsules weakly inflated.

Female caudodorsal beak in dorsal view (fig. 161)

narrow triangular and bluntly rounded, almost point-

ed at apex. Ovipositor sheaths almost reaching to

apex of caudodorsal beak.

Male genitalia: Pygofer in lateral view as in fig.

157. Dorsal margin weakly convex, concavely bent

into straight, erect and very slender caudodorsal beak.

Distal margin weakly convex, concavely bent into

margin of beak. Ventral margin convex, forming a

small angular corner under lateral protuberance.

Ventral margins converging towards base of pygofer;

basal part of pygofer opening V-shaped (fig. 156).

Caudodorsal beak in dorsal view (fig. 160) very slen-

der, oblong, and rounded at apex. Lateral lobe of py-

gofer strongly curving inwards towards end of distal

margin, forming a weakly swollen, bluntly rounded

lateral protuberance. Claspers (fig. 162) very different

from other species of this genus, almost straight and

parallel towards rounded apices. Claspers adjacent,

DE BOER: The genus Papuapsaltria

but diverging at half-length, curving around aedea-

gus. Aedeagus not adjacent to base of anal valves.

Claspers almost fused, dorsally separated by narrow

furrow between aedeagus and base of anal valves.

Apical part of clasper weakly curved down, with large

and sharply edged ventral hollow. Aedeagus (fig. 158)

weakly S-curved, with long and slender lateral crests.

Aedeagal pore round.

Measurements: Body length d: 20.0-26.5 mm (x

22.3 mm + 2.3), ®: 18.2-21.1 mm (x 20.0 mm

+ 0.9); tegmen length d: 23.6-31.4 mm (x 27.1 mm

+ 2.4), 2: 29.1-33.3 mm (x 31.0 mm + 1.5); head

length d: 1.3-1.7 mm (x 1.4 mm), 2: 1.5-1.8 mm

(x 1.7 mm); pronotum length d: 1.8-2.5 mm (x 2.1

mm), 9: 2.3-2.8 mm (x 2.5 mm); mesonotum length

d: 4.2-5.2 mm (X 4.7 mm), 9: 4.7-5.8 mm (x 5.1

mm); head width d: 3.4-4.1 mm (X 3.7 mm), ©:

4.1-4.3 mm (x 4.2 mm); width of pronotal collar d:

5.3-7.0 mm (x 5.8 mm), ®: 6.5-7.3 mm (x 6.8 mm).

Distribution (fig. 4). — P. dioedes is found in the

central mountain ranges of western Papua New

Guinea, at high altitudes, and on Mt. Bosavi, just

south of these mountain ranges.

Etymology. — Dioedes (Greek) means swollen and

refers to the swollen abdomen of this species.

Papuapsaltria woodlarkensis sp. n.

(figs. 4, 10, 165-174)

Type material. — Holotype d : ‘NEW GUINEA: PAPUA

/ Woodlark (Murua) / Kulumandau Hill / Feb. 3.

1957’ [print]; “W.W. Brandt / Collector’ [print],

BPBM. — Paratypes: WOODLARK ISLAND: same data as

holotype 18, 19, ZMAN; 29, BPBM; same data but

28301.19574.12:2101119572.12:216:1419579235

DMSA ENIS ul 05745 EN 57E

16-22.iv.1957, 18, 19, all BPBM; Kulumandau, 0-

100 m, 3.xi.1956, 5th Archbold Exped. to New

Guinea, L.J. Brass, 19, AMNH; same data but

10.x1.1956, 16, 19, AMNH.

P. woodlarkensis is easily recognized by the conical,

thorn-shaped, and inwardly curved protuberances at

the ventral margin of the pygofer.

Description

Body of males almost uniformly ochraceous-

brown, but tinged with green. Females more reddish

brown or bicolorate, with greenish head and thorax

and reddish brown abdomen. Females distinctly

smaller than males, but with more robust head and

thorax, and longer tegmina. Tegmina of males 1.0-

1.1 X as long as body length, of females 1.2-1.3 x.

Male abdomen 1.5-1.6 X as long as head and thorax,

of females 1.2-1.3 X.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 175-181. 175. Papuapsaltria spinigera sp. n.: first ulnar area of right tegmen, holotype. 176-181. Papuapsaltria phyllo-

phora (Blöte, 1960): 176, first ulnar area of right tegmen, Bokondini; 177, male first and second tergite in dorsal view,

Bokondini; 178, male head in dorsal view, Bokondini; 179, female genital segment in lateral view, Bokondini; 180, female

caudodorsal beak in dorsal view, Bokondini; 181, female operculum, Bokondini.

Head: Postclypeus slightly angularly protruding,

smoothly rounded anteriorly, its anterior margin

weakly convex, almost continuous with anterior mar-

gins of vertex lobes. Postclypeus not swollen; anterior

margin (lateral view) straight. Rows of short parallel

ridges form a broad, though weakly developed, band

along lorum. Distance between lateral ocelli 0.9-

1.2 X as long as distance between lateral ocellus and

eye and in males 2.0-2.2 X, in females 2.2-3.1 X,

the width of frontal ocellus.

Tegmina: Hyaline, with 8 long and slender apical

areas, a fairly distinct costal area, and a very broad

hyaline border along hind margin.

Legs: Fore femur (fig. 10) with row of three or four

erect spines, diminishing in length towards tibia.

Proximal spine fairly long, distinctly longer than mid-

dle spine and only slightly shorter than distance to

middle spine.

Tymbals: Five parallel transverse sclerotized ridges

spanning the tymbal from dorsal to ventral margin

and a Gth ridge almost reaching ventral margin. Six

short intercalary ridges clearly visible.

Opercula: Male operculum (fig. 172) not covering

tymbal cavity in ventral view, leaving a wide gap

between operculum and abdomen; folded membrane

clearly visible in ventral view. Distal part of opercu-

lum angularly oblong and distinctly longer than basal

part. Lateral margin long and straight, convex to base,

ending in almost right angle on crest of basal part and

reaching that crest close to lateral corner. Distal part

of basal crest very short, and lower than its lateral

part. Distolateral corner rounded, distal and medial

margins straight, distomedial corner rectangular.

Meracanthus longer than operculum, but not reach-

ing to margin of abdominal segment 2. Female oper-

culum (fig. 173) resembling that of male, but much

shorter, with its distal part shorter than basal part,

and oblong. Lateral margin as in male. Distal margin

weakly convex, medial margin short and straight.

Abdomen: Male abdomen strongly inflated. First

tergite rather long; medially about half as long as 2nd

tergite. Anterior margin of 2nd tergite weakly convex

medially. First and 2nd sternites not adjacent.

Auditory capsules weakly developed, hardly protrud-

ing and not visible in dorsal view. Female abdomen

stout. Female caudodorsal beak (fig. 174) oblong and

rounded at apex. Ovipositor sheaths reaching dis-

tinctly beyond apex of caudodorsal beak (fig. 165).

Male genitalia: Pygofer in lateral view as in fig.

166. Dorsal margin concave, but convexly, almost

angularly, bent into straight, short, and stout caudo-

dorsal beak. Distal margin weakly convex between

caudodorsal beak and lateral protuberance, forming

an obtuse angle with straight margin of caudodorsal

beak. Caudodorsal beak in dorsal view (fig. 169)

short and broad, weakly convex, almost truncate at

apex. Lateral lobe of pygofer with large swollen and

bluntly rounded protuberance. Ventral part of pygo-

fer lobe strongly curving inwards towards ventral

margin, forming a slender conically rounded, almost

thorn-shaped and inwardly directed, protrusion just

below lateral protuberance (fig. 167). This protrusion

is not visible in lateral view. Ventral margin of pygo-

fer in lateral view weakly concave below lateral protu-

berance, but convex to base. Ventral half of pygofer

opening narrowly oblong, forming a distinctly swol-

len ridge along its margins (fig. 167). Claspers (fig.

171) very short, hardly reaching beyond pygofer, di-

verging towards sharply pointed and downwardly di-

rected apices. Clasper forming a weak dorsal crest.

Rectangular distodorsal corner of that crest bending

mesiad, distally of aedeagus and supporting aedeagus

in upright position. Distal margin of clasper almost

straight. Apical part of clasper with large, sharply

edged hollow, ending at base in weak lateral swelling.

Clasper base forming a ring-shaped collar around

base of anal valves, which is medially sharply indent-

ed around aedeagus. Aedeagus standing erect between

claspers, close to anal valves, its apical part slightly

curving away from anal valves. Aedeagus (fig. 170)

strongly S-curved, with long and distinct lateral crests

and a weak dorsal ridge. Apex of aedeagus weakly

pointed in lateral view, with oval, almost round, pore

(fig. 168).

Measurements: Body length d: 19.7-21.0 mm (x

20.3 mm + 0.5), 2: 18.0-19.8 mm (x 18.9 mm +

0.6); tegmen length d : 20.5-21.8 mm (x 21.2 mm +

0.5), 2: 22.6-24.5 mm (x 23.3 mm + 0.6); head

length d: 1.4-1.6 mm (x 1.5 mm), 9: 1.6-1.8 mm

(x 1.7 mm); pronotum length d: 2.2-2.3 mm, 9:

2.2-2.5 mm (X 2.3 mm); mesonotum length d: 4.0-

4.4 mm (x 4.2 mm), 2: 4.2-4.7 (x 4.4 mm); head

width G : 4.0-4.4 mm (x 4.2 mm), 9 : 4.3-5.0 mm (x

4.8 mm); width of pronotal collar d: 5.2-5.7 mm (x

5.5 mm), 2: 5.7-6.4 mm (x 6.0 mm).

DE BOER: The genus Papuapsaltria

Distribution (fig. 4). — P. woodlarkensis is presum-

ably endemic to Woodlark Island of the

D’Entrecasteaux islands.

Etymology. — The name of this species is derived

from the type locality: Woodlark island.

Papuapsaltria phyllophora (Blöte, 1960) comb. n.

(figs. 4, 176-190)

Baeturia phyllophora Blöte, 1960: 79, figs. 45, 46; Duffels &

Van der Laan 1985: 254; De Boer 1993a: 16.

The generic allocation of this species is dubious. P.

phyllophora shares none of the apomorphies on which

any of the genera of the ‘Baeturia and related genera

complex’ are based. Furthermore, the apomorphy of

that complex as a whole, an S-curved aedeagus with

wing-shaped lateral crests, can hardly be recognized

in the strongly deviating aedeagus of P. phyllophora;

this deviating shape is here regarded as autapomor-

phous. The small male opercula and smoothly vault-

ed vertex suggest that the species either belongs to

Guineapsaltria or to Papuapsaltria. The species is pre-

liminary included in Papuapsaltria since its ocelli are

more closely together than is generally the case in

Guineapsaltria. Current phylogenetic analysis con-

cerning all species of the ‘Baeturia and related genera

complex’ indicate that P. phyllophora either as the sis-

ter group of Guineapsaltria, or as part of Papua-

psaltria both belong to the most parsimonious solu-

tions. Males and females of P. phyllophora can easily

be recognized by a swelling of the costa at the distal

end of the first radial area, and by the quintangular-

shaped second ulnar area. Males are unmistakable by

their peculiar aedeagus.

Description

Body of males unicoloured red-brown or olive

green, of females often darker brown or greenish.

Females on average slightly longer than males, but

with distinctly more robust head and thorax and lon-

ger tegmina. Tegmina of males 1.2-1.4X as long as

body length, of females 1.3-1.5X. Male abdomen

1.2-1.5 X as long as head and thorax, of females 1.0-

122

Head (fig. 178): Reddish brown or greenish, often

reddish along anterior margins of postclypeus and

vertex lobes. Vertex and postclypeus with short red-

brown setae. Postclypeus very broad in dorsal view,

3.0-4.3 X as broad as long and oblong-shaped, hard-

ly protruding beyond vertex lobes. Anterior margin of

postclypeus almost continuous with anterior margins

of vertex lobes. Postclypeus not swollen, its anterior

margin (lateral view) concave. Lateral surface of post-

clypeus quite smooth, with only weakly developed

rows of parallel ridges. Distance between lateral ocelli

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 182-190. Papuapsaltria phyllophora (Blöte, 1960): 182, pygofer from aslant, Bokondini; 183, pygofer in lateral view,

Bokondini; 184, male operculum, Bokondini; 185, claspers and aedeagus, Bokondini; 186, aedeagus in lateral view, Wau;

187, aedeagus from aslant, Wau; 188, male caudodorsal beak in dorsal view, Bokondini; 189, male fore femur, Bokondini;

190, detail male fore femur, Bokondini.

0.8-1.2 X, though in most specimens practically 1.0

X, distance between lateral ocellus and eye and 2.2-

3.5 X the width of frontal ocellus (1.3X in one fe-

male).

Thorax: Red-brown or olive green. Pronotum with

very weak medial furrow. Lateral corners of pronotal

collar often slightly swollen proximally.

Legs: Ochraceous, fore tibiae and tarsi reddish.

Fore femur with row of three erect spines, dimin-

ishing in length towards tibia. Proximal spine slightly

longer than middle spine, but shorter than distance to

middle spine (figs. 189-190).

Tegmina: Hyaline, with 8 fairly short apical areas,

a distinct costal area and a broad hyaline border along

hind margin. Costa distinctly inflated towards distal

end of first radial area. Second ulnar area definitely

quintangular shaped, practically quadrangular in all

related species (compare figs. 175 and 176).

Tymbals: Eight darkly sclerotized transverse ridges

spanning the tymbal from dorsal to ventral margin,

sometimes with a partly developed 9th ridge, closely

along proximal tymbal margin. Eight short and much

lighter coloured intercalary ridges seem to form a

band across the tymbal.

Opercula: Male operculum (fig. 184) very small

and angular, not covering tymbal cavity, and leaving

a narrow gap between operculum and abdomen; fold-

ed membrane hardly visible in ventral view. Distal

part of male operculum angular, oblong-shaped, and

erect. Lateral margin of distal part straight, forming

an obtuse angle with straight distal margin, and with

crest around distolateral corner of basal part. Medial

margin straight, distomedial corner angularly round-

ed. Meracanthus reaching well beyond operculum, to

sternite 2. Female operculum (fig. 182) very similar

to that of male, but slightly shorter. Distal part ob-

long-shaped and erect, with straight margins and al-

most rectangular distomedial corner.

Abdomen: Male abdomen ochraceous brown or

greenish, not inflated. Segmental hind margins some-

times reddish. First tergite short and broad, partly

hidden under metanotum. Medial part of 2nd tergite

about 1.5X as long as lateral parts (fig. 177).

Anterior margin of second tergite almost straight me-

dially. Auditory capsules distinctly swollen and visible

in dorsal view (fig. 177). First and 2nd sternites adja-

cent. Female abdomen more slender than that of

males. Female pygofer (fig. 179) robust, in lateral

view broader than long. Female caudodorsal beak in

dorsal view (fig. 180) stout, broadly rounded at apex.

Ovipositor sheaths reaching just beyond apex of cau-

dodorsal beak.

Male genitalia: Pygofer in lateral view as in fig.

183. Dorsal margin slightly convex, concavely bent

into stout, straight and erect caudodorsal beak. Distal

margin convex between base of beak and lateral pro-

tuberance. Ventral margin almost straight. Ventral

margins converge to a sharp angle at base of pygofer

opening; ventral part of pygofer opening V-shaped

(fig. 182). Pygofer lobe curved inwards towards distal

margin and forming a small, globularly rounded, lat-

eral protuberance. Caudodorsal beak in dorsal view

(fig. 188) short, stout and broadly rounded at apex.

Claspers very slender and parallel to rounded apices

(fig. 185). Proximal part of clasper very long and

straight, adjacent to membranous part of aedeagus.

Apical part of clasper long and slender, with very shal-

low ventral hollow, strongly bent downwards. Dorsal

part of clasper base forming a broad rounded ridge at

base of anal valves. Dorsal ridges of both claspers dis-

tally elongated. Elongated parts curving mesiad, al-

most fusing, and forming a small hood over aedeagus.

Clasper forming a rounded medial protrusion, sup-

porting aedeagus. Aedeagus (figs. 186-187) very pe-

culiar in shape, resembling a pick-axe; consisting of a

stem ending in long and slender, recurving, dorsal

and ventral protrusions. A square-shaped, semi-trans-

parent membrane connecting dorsal protrusion with

stem. This membrane strengthened by a crest along

its proximal margin. Ejaculatory duct following ven-

tral protrusion, ending in posteriorly directed pore.

Aedeagus with pair of short, semi-circular, wing-

shaped crests ventrally along the stem. Aedeagal crests

bending slightly laterad and resting on medial protru-

sions of claspers.

Measurements: Body length d: 12.0-15.0 mm (x

13.3 mm + 0.8), 2: 11.6-15.8 mm (x 14.0 mm +

1.0); tegmen length d: 14.6-18.2 mm (x 16.8 mm +

0.7), 2: 15.6-23.0 mm (x 19.1 mm + 1.7); prono-

tum length d: 1.4-1.7 mm (x 1.5 mm), 9: 1.5-2.0

mm (x 1.7 mm); mesonotum length d: 2.5-3.2 mm

(x 2.8 mm), 9: 2.8-4.1 mm (x 3.4 mm); head length

4: 1.0-1.3 mm (x 1.1 mm), 2 1.1-1.6 mm (x 1.3

mm); head width d: 3.0-3.4 mm (x 3.2 mm), 2:

3.1-4.1 mm (x 3.5 mm); width of pronotal collar d:

3.6-4.2 mm ( 3.9 mm), 2: 4.2-5.4 mm (X 4.6 mm).

Material examined. — IRIAN JAYA: Bewani R. territ., 1200

m, 1939, W. Stüber leg., 19, RMNH; Bokondini, 10 km E,

40 km N of Baliem Val., 1300 m, 24.xi.1961, L. & S.

Quate, 166, 119, BPBM; same data 2d, 29, ZMAN;

Waigeu, Camp Nok, 2500 ft, iv.1938, L.E. Cheesman, 34,

DE BOER: The genus Papuapsaltria

19, BMNH; same data but iv-v.1938, 26, 59, BMNH; 1d,

1, ZMAN; PAPUA NEW GUINEA: Arau, 40 km E of Kainantu,

1400 m, 15.x.1959, T.C. Maa, 19, BPBM; Arau, Kratke

Mts., Valley of Upper Wanton R., 1400 m, 7.x.1959, L.J.

Brass, 19, AMNH; Baiyer River, 14.1.1978, R.B. Lachlan,

19, Moul.; Big Wau Creek, Wau, 1200 m, xii.1965, J.

Sedlacek, 14, BPBM; Coviak Rdg., Wau, 763 m, 7.xii.1963,

H. C., 1d, 59, BPBM; Damanti, Finisterre Mts., Madang

Dist., 3550 ft, 2-11.x.1964, M.E. Bacchus, 18, 19, BMNH;

Kumur, Upper Jimi V., 1000 m, 12.vii.1955, J.L. Gressitt,

16, BPBM; same data but 13.vii.1955, 12, BPBM; Garaina S,

900-1800 m, 8-14.x.1968, 19, BPBM; Loloipa, Goilala,

Owen Stanley Range, 11-20.xii.1957, W.W. Brandt, 19,

BPBM; Mafulu, 4000 ft, i.1934, L.E. Cheesman, d holotype

Baeturia phyllophora Blöte, 19 paratype, 19 allotype,

BMNH; Mt. Missim, Wau, 950-1300 m, xii.1965, J.

Sedlacek, 19, BPBM; Mt. Missim, Wau, Morobe Dist.,

1200-1800 m, 8.xii.1963, H. Clissold, 16, 19, ZMAN;

same data but 1600 m, 17.11.1963, Gressitt, 19; 1800 m,

22.11.1966, 19, both BPBM; Mt. Missim, 7°16° S 146°48’

E, 1500-1800 m, 7.1.1970, M. Sedlacek, 16, BPBM;

Moroka, 1300 m, vii-xii.1895, Loria, 19 paratype Baeturia

phyllophora Blôte, BMNH; Nakata Ridge, Wau, 1800 m,

18.xii.1963, H. Clissold, 16, 19, BPBM; Sattelberg, Huon

Golf, 1898, Biré, 38, 29, TMB; Upper Baiane, Nr.

Bulowat, 1100 m, 3.vi.1979, J.L. Gressitt, 19, BPBM;

Wanuma, Adelbert Mts., 800-1000 m, 26.x.1958, J.L.

Gressitt, 2d, 59, BPBM; Wau, 1200 m, 21.xi.1963, J.L.

Gressitt, 19, BPBM; same data but 25-30.ix.1964, J. & M.

Sedlacek, 19; 1-9.ix.1965, J. Sedlacek, 19; 11.xii.1965, J.

& M. Sedlacek, 16; 14.iii.1966, Gressitt & Wilkes, 14,

19, all BPBM; Wau, 1150-1250 m, 17.11.1966, J. Sedlacek,

19, BPBM; same data but 1100-1300 m, 12.1.1966, 16,

BPBM; Wau Area, Morobe P., iii-iv.1984, T.R. New, 16,

Moul; Wau, 10 km S, st. 054, 22.v.1988, J. van Stalle, 19,

ISNB; Wau, Ecology Inst., 1220 m, 18-20.vi.1981, J. van

Goethem, 19, ISNB; Wau, Hospital Ck., 1200 m,

17.11.1965, J. Sedlacek, 19, BPBM; same data but 1250-

1300 m, 7.iii.1965, 19, BPBM; Wau, Morobe Distr., 14.-

viii.1972, G.G.E. Scudder, 16, BPBM; same data but 1000

m, 1.111.1963, H.W. Clissold, 18; 1090 m, 25.1.1963, J.

Sedlacek, 19; 1100 m, 26.x.1961, J. Sedlacek, 19, all

BPBM; Wau, Morobe Distr., 1200 m, 1-20.xi.1961, J.

Sedlacek, 12; same data but 2-10.xi.1961, J. & M.

Sedlacek, 1 9 ; 19.xi.1961, J.H. Sedlacek, 1 d ; 22.xi.1961, J.

& J.H. Sedlacek, 16; 2.xii.1961, J. Sedlacek, 19; 20-

26.v.1962, J. Sedlacek, 19; 10.11.1963, J. Sedlacek, 19;

15.1.1963, J. Sedlacek, 16, all BPBM.

Remark: The female paratype from Kokoda, mentioned

by Blôte (1960) does not belong to this species but to

Guineapsaltria stylata (Blöte, 1960) (see De Boer 1993a).

Distribution (fig. 4). — P. phyllophora is distributed

throughout northern New Guinea, including the is-

land of Waigeu, and the Huon and Papuan peninsu-

las, but is not recorded from Cendrawasih.

ACKNOWLEDGEMENTS

For the loan of material I am indebted to: Dr R. T. Schuh

(AMNH); Dr W. J. Knight and Mr M. D. Webb (BMNH); Mr

G. M. Nishida and Mr K. Arakaki (BPBM); Mr J. van Stalle

(ISNB); Dr A. Neboiss (MVMA); Dr L. L. Deitz (NCSU); Dr A.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

H. Kirk Spriggs (NMWC); Mr J. van Tol (RMNH); Dr H.

Schröder (SMFD); Mr F. Heller (SMNS); Dr R. Emmrich

(SMTD); Dr T. Väsärhelyi (TMB); Dr R. Danielson (ZILS);

and to Mr M. S. Moulds for the loan of specimens from his

private collection.

I would like to thank Mr G. Verlaan for technical assis-

tance and and Mr D.A. Langerak for preparing the maps

(figs. 3-4, 32 and 129). I am indebted to Professor Dr F. R.

Schram and Dr J. P. Duffels (Instituut voor Systematiek en

Populatie Biologie, University of Amsterdam) for their crit-

ical reading and comments on the manuscript. My special

thanks go to the ‘Uyttenboogaart-Eliasen stichting’ for the

grant I recieved to visit New Guinea during the months of

October and November 1994. This trip enabled me to col-

lect the holotype of P. dolabrata.

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VII. Part 2. Tibicinidae. — North Carolina State College,

Raleigh, N.C.: i-iv + 1-492.

Moulds, M.S., 1990. Australian cicadas. — New South

Wales University Press, Kensington: 1-217.

Moulton, J. C., 1923. Cicadas of Malaysia. — Journal

Federated Malay States Museum 11: 69-182 + pls. 1-5.

Myers, J. G., 1928. Cicadidae. Insects of Samoa and other

Samoan terrestrial Arthropoda 2 (2): 55-65. — Trustees of

the British Museum, London.

Pigram, C. J. & H. L. Davies, 1987. Terranes and the accre-

tion history of the New Guinea orogen. — BMR Journal of

Australian Geology and Geophysics 10: 193-212.

Swofford, D.L., 1993. PAUP: Phylogenetic Analysis Using

Parsimony, version 3.1.1. — Computer program distributed

by the Illinois Natural History Survey, Champaign, Illinois.

Wheeler, T., 1988. Papua New Guinea a travel survival kit.

— Lonely Planet Publications, Victoria, Australia: 1-340.

Received: 9 January 1995

Accepted: 4 April 1995

P. E. BRAGG

Ilkeston, United Kingdom

A REVIEW OF THE SUBFAMILY KORINNINAE

(PHASMIDA: PSEUDOPHASMATIDAE), WITH THE

DESCRIPTION OF A NEW SPECIES

Bragg, P. E., 1995. A review of the subfamily Korinninae (Phasmida: Pseudophasmatidae),

with the description of a new species. — Tijdschrift voor Entomologie 138: 45-50, figs. 1-9.

[ISSN 0040-7496]. Published 15 June 1995.

The subfamily Korinninae is reviewed. A key to genera and species is provided. Kalocorinnis

pulchella (de Haan), was previously placed in a different suborder, was found to be the senior

synonym of Kalocorinnis calopteryx Günther. Variation in the males of K. pulchella is discussed

and illustrated. A new species, Kalocorinnis wegneri, from Borneo, is described and illustrated.

The female of Korinnis errans Günther is illustrated. The egg of K. wegneri is the first egg to be

described and illustrated from this subfamily.

P.E. Bragg, 51 Longfield Lane, Ilkeston, Derbyshire, DE7 4DX, United Kingdom.

Key words. — Phasmida, Korinninae, Korinnis, Kalocorinnis, new species, Borneo.

This paper is produced as the result of a visit to

Leiden (RMNH) to examine the type specimens of de

Haan. During this visit it was discovered that one of

de Haan’s species appeared to be the same as the par-

atype of a species described by Günther in 1944

which is also in the RMNH collection. According to

the literature however the two species were consid-

ered to be in different suborders. Careful cleaning

and examination of de Haan’s type specimen showed

that it had been placed in the wrong suborder by

Redtenbacher in 1908.

The RMNH collection has a large number of un-

identified specimens, many from Borneo. These were

examined and several more specimens of this subfam-

ily were located, including one species which is de-

scribed here for the first time.

KORINNINAE

Korinninae Günther 1953: 550. — Type genus: Korinnis

Günther 1932.

The subfamily belongs to the suborder Areolatae

and family Pseudophasmatidae. This is a small sub-

family which is restricted to south east Asia. The sub-

family contains only four species which fall into two

genera. The two suborders of Phasmida are distin-

guished by the presence of a triangular depression on

the underside of the apex of the middle and hind tib-

iae in the Areolatae, and the absence of this feature in

the Anareolatae. The use of Bradley & Galil’s key to

families (1977: 178) places these species in the

Pseudophasmatidae because the tarsi have five seg-

ments, the first abdominal segment is longer than

metanotum, the metanotum is longer than it is wide,

and the antennae are long and filiform. The key to

subfamilies of Pseudophasmatidae (Bradley & Galil

1977: 200) refers these species to the Korinninae be-

cause they possess normally formed elytra, four un-

armed carinae on the femora, and they lack ocelli.

The following key serves to distinguish the genera

and species of the subfamily.

Key to the subfamily

1. Base of fore femora curved, mesonotum with lar-

ge tubercules, wings not coloured, or only tinted

(Korinnis spp.)

— Base of femora straight, mesonotum smooth,

wings strongly coloured (Kalocorinnis spp.) ..... 3

2. Mesonotum evenly tapering and not armed with

bluntispines mesa Korinnis errans Günther

— Mesonotum tapers in two distinct stages and is

armed with eight blunt spines … nen.

bgn CI DIM I) È, Korinnis potameis Günther

3. Mesonotum evenly tapering and not armed with

blunt spines ..... Kalocorinnis pulchella (de Haan)

— Mesonotum tapers in two distinct stages and is

armedawithtsixablumbspines ren

PO Pe, Se EE Kalocorinnis wegneri sp.n.

Ragge (1955), in his study of wing venation in

Phasmida, did not examine any representative of the

Korinninae. The wing venation agrees with the gen-

eral pattern which Ragge found in all groups that he

examined (1955: 390). In the Korinninae the costa,

subcosta, radius, cubitus, and first anal are all un-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

branched, the media branches near the base to form

the anterior and posterior media. Ragge considered

an unbranched radius to be unusual, this state occur-

ring only ‘in isolated genera (or even species) in two

subfamilies (Aschiphasminae and Necrosciinae)’. The

only difference within the Korinninae is the length of

the subcosta; this is between three quarters and four

fifths of the wing length in Korinnis but only about

two thirds in Kalocorinnis. The females of all four

members of the subfamily have an operculum with a

deep notch in the apex. This condition is rare in

Aschiphasmatinae (present in one out of 13 species

examined) although not unusual in the Pseudophas-

matinae.

The distribution of the subfamily is probably limit-

ed to the islands of the East Indies and the Malay

Peninsula. All four species occur in Borneo; reliable

records from outside Borneo refer to only one species,

Kalocorinnis pulchella (de Haan).

SYSTEMATIC PART

Kalocorinnis Günther

Kalocorinnis Günther, 1944: 77. — Type species Kalocorinnis

calopteryx Günther 1944 by original designation [= K.

pulchella (de Haan, 1842)].

Kalocorinnis pulchella (de Haan) comb.n.

(figs. 1-4)

Phasma (Necroscia) pulchellum de Haan, 1842: 120, pl. XV

fig. 5. — Holotype d, Sumatra, Batang Singalang

(RMNH) [examined].

Necroscia pulchella (de Haan). — Westwood 1859: 152;

Kirby 1904: 377.

Tagesoidea pulchella (de Haan). — Redtenbacher 1908: 565.

Kalocorinnis calopteryx Günther, 1944: 78, fig. 5. —

Holotype: ©, Nordborneo (RMNH) [examined],

Paratypes: 9, Central-Borneo (SMTD); d, Peninsular

Malaysia, Selangor, Bukit Kutu, 1100m, 13-iii-1931,

H.M. Pendlebury (RMNH) [examined]. Syn. n.

Material examined. — Holotypes: Phasma (Necroscia) pul-

chellum, &; Kalocorinnis calopteryx, 2. Paratype:

Kalocorinnis calopteryx, 3. — Others: Midden O-Borneo,

14.viii.1925, H. C. Siebers, 19 (RMNH); E. Borneo, 125m

Tabang, Bengen River, 28.x.1956, A.M.R. Wegner, 3d

(RMNH); Sarawak, J.E.A. Lewis, 1d (BMNH, 1910-116);

Sarawak, Wallace, 168 (OXUM); Sabah, Danum Valley,

220m, light trap sample, roadside, secondary forest,

10.ix.1987, A.H. Kirks-Spriggs, 19 (NMWC,

NMW.Z.1987.094).

Examination of the holotype of pulchella showed

that it belongs in the subfamily Korinninae; the orig-

inal description makes no mention of the characteris-

tic which is used to distinguish the two suborders.

Direct comparison of de Haan’s male holotype, and

Günther’s male paratype leave no doubt that they are

the same species.

Rather unusually for phasmids, the wings of this

species are brightly coloured and have a distinctive

pattern. In view of the fact that the species had al-

ready been described and well illustrated (de Haan

1842), and had been recorded from Borneo

(Westwood 1859: 152), Günther’s description of

Kalocorinnis calopteryx as a new species from Borneo

might appear strange. However some research into

the background suggests a rational explanation of

this.

The most recent comprehensive work on Phasmida

is the three part monograph by Brunner von

Wattenwyl (1907) and Redtenbacher (1906, 1908),

this is still used as the basis for identification of phas-

mids by most workers. Redtenbacher (1908) placed

pulchella in his “Tribus Necrosciini (now the subfam-

ily Necrosciinae). This was presumably based only on

de Haan’s description and illustration; the species is

not represented in Redtenbacher’s collection. De

Haan’s description is quite brief and makes no men-

tion of the feature which distinguishes the two subor-

ders of Phasmida: the sunken triangular area on the

middle and hind tibiae. It appears that Günther had

not seen de Haan’s type specimen; Leiden Museum

records show no evidence of a visit by Günther. The

illustrations in de Haan’s publication (1842) are very

accurately drawn but are hand coloured; examination

of two copies (RMNH and Oxford University library)

has shown that the pattern on the wings varies (al-

though less than can occur in the insects themselves).

Westwood (1859: 152) correctly identified Wallace’s

specimen although it differs considerably from de

Haan’s specimen (figs. 1, 4). Günther probably did

not consider the possibility that Redtenbacher had

placed the species in the wrong suborder, the species

is so distinctive that he would almost certainly have

recognised it despite variations in de Haan’s illustra-

tions.

The type specimens have suffered damage since

they were originally described. The holotype of put

chella has only two legs remaining, the right fore leg

and left mid leg; the tarsi of both these legs are miss-

ing. Günther illustrated the calopteryx holotype

(1944: 75, fig. 5) at which time it had one antenna

missing, it has since lost both front and both middle

legs; the male paratype lacks both front legs, the left

mid leg, both antennae, and the end of the abdomen.

This species shows variation in both colour and

pattern of the wing patches. There is considerable

variation in the shape of the patches of the males (figs.

1-4), this is particularly noticeable in Wallace’s speci-

men which has greatly reduced patches. The wing

patches of all three females which have been exam-

ined, and the two male types, are cream coloured.

One of the three Tabang males has pale blue-green

patches, the other two have white patches with a blue-

green tinge to the veins. Wallace’s male also has blue-

green patches. The BMNH specimen has yellow

patches. Table 1 gives measurements of the female

collected by Siebers and one of the males collected by

Wegner, part of the abdomen of the male is missing.

This species appears to be quite widespread, having

been recorded from Borneo, peninsular Malaysia and

Sumatra. Within Borneo it has been recorded from

Sabah (Günther’s male paratype and NMWC speci-

men), Sarawak (Westwood, 1859: 152, and Lewis)

and Kalimantan (Wegner, Siebers, and Giinther’s fe-

male paratype).

Kalocorinnis wegneri sp.n.

(fig. 5)

Type material. — Holotype © : East Borneo, 125m

Tabang, Bengen River, 3.ix.1956, A.M.R. Wegner

(RMNH). — Paratype 9: Sabah, Sepilok, 23-11-1983,

Shinji Nagai (C.L. Chan).

The holotype is in poor condition; all the internal

parts of the head and thorax have been eaten, as have

all thoracic sternites; the one remaining antenna is

broken; the left hind and mid legs are missing and on-

ly one tarsus remains; the abdomen is shrunken and

distorted. Despite the poor condition, the wings are

very distinctive and this species is unlikely to be con-

fused with any other.

The paratype is in better condition than the holo-

type, but lacks any front legs, antennae, left mid leg

and right mid tarsus. It differs from the holotype by

being slightly longer (57 mm), and by being more

brightly coloured. It is likely that the colour of the

holotype was originally similar to the paratype. The

abdomen is much fatter, while some of this is due to

dorso-ventral compression during preservation, it

clearly shows that the abdomen of the holotype is

badly shrunken.

Measurements of the holotype and paratype are

given in table 1. Figure 5 is based on the holotype but

has been modified to take some account of the dam-

age and distortion of the specimen; the right wing has

been omitted.

Head globular, longer than wide, smooth. Eyes

prominent. Antennae filiform; light brown at the

base, becoming darker (broken off).

Pronotum mid brown, lateral surfaces green in the

paratype; smooth, with a deep transverse indentation

about one third of the way from the front margin.

Mesonotum granulose; narrow, with parallel sides at

the anterior; widening and swelling suddenly one

third of the way back; with two pairs of laterally

pointing blunt spines near the front edge, and one

BRAGG: Review of Korinninae

blunt spine on each side, slightly in front of the mid

point of the swelling. Anterior third of holotype light

brown, posterior portion mid to dark brown; para-

type evenly mid brown dorsally and green on lateral

surfaces. Metanotum and abdominal segments light

brown; paratype with green lateral surfaces. It is pos-

sible that when live the whole of the paratype’s abdo-

men may have been green.

Abdominal segments of the holotype are shrunken

and distorted, those of the paratype are flattened, but

the abdomen appears to narrow evenly along its

length. Segments 1-7 becoming evenly shorter, 8th

segment noticeably shorter than 7th; 9th and 10th

much shorter and of equal length. The slender lami-

na supraanalis is as long as the 10th segment and ta-

pers to a point. Operculum covered in bristles, with a

deep notch at the apex.

Elytra almost circular; with an obvious hump near

the outer margin; evenly covered in prominent fine

veins.

Wings well rounded. Holotype with costal region

chocolate brown, anal region of the wing tessellated

dark brown and white, with an arc of large white

patches about one two thirds of the way from the base

of the wing, and small white triangular spots on the

margin between the veins. Paratype with costal region

of wings green in front of the main vein and mid

brown behind; anal region dark brown with very few

white tessellations; an arc of pale blue patches corre-

sponds to the white patches of the holotype; the white

triangles on the margin are present but some have a

pale blue tint.

Femora all with four carinae; smooth, apart from

fine bristles mainly on the carinae. The fore femora

are straight and narrow only slightly at the base. Hind

femora slightly enlarged and laterally compressed.

Tibiae with four indistinct carinae; only slightly

widened at the apices. Fore and mid tarsi with five

segments; first tarsomere one third of the total; fourth

tarsomere very short. (Middle tarsi missing from both

specimens).

Male: Unknown.

Egg. — Several eggs (figs. 7-9) were removed from

the body of the paratype by rehydration and partial

dissection of the abdomen. While some eggs appear

to be fully developed, there may be some slight differ-

ences in surface detail between these and eggs which

are laid. Some of the eggs have been slightly damaged

during the original preservation or by their subse-

quent rehydration; this has caused some distortion of

the ventral surface, as a consequence the true curva-

ture of the ventral surface may differ slightly from

that shown in the illustration (fig. 8).

Whole of capsule mid brown. Capsule twice as

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

DEN ETSER

FA LL TE I

DEEE

DER De

Een

Katar > Bere en

LEAD

ANS

; : SSS == 2

N x =

n N <a = I TT

d INI PI Si e =

II N —

i ROS = Son

I SS È

| Le \ NN : NT n > È

N x

IE NOR NC

4 \ \ \ \

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eenen

ORC mare ES

RES

ET —=

Figs. 1-9. Korinninae. — 1-4. Wing patterns of male Kalocorinnis pulchella (de Haan): 1, De Haan’s holotype; 2, Giinther’s

paratype; 3, One of Wegner’s specimens; 4, Wallace’s specimen. — 5. Kalocorinnis wegneri sp. n., holotype 2. — 6. Korinnis

errans Günther, 2. — 7-9. Egg of Kalocorinnis wegneri n.sp. (removed from paratype): 7, dorsal view; 8, lateral view; 9, inter-

nal view of dorsal surface. Scale line A (1 cm) for figs. 5 and 6, scale line B (1 mm) for figs. 7-9.

long as wide, tapering slightly at polar end, dorsal

surface sloping steeply towards the ventral surface at

the anterior end; ventral surface longitudinally al-

most flat, laterally sharply convex; dorsal surface

strongly convex. A sharp ridge circles the egg, separ-

ating the dorsal and ventral surfaces; a similar ridge

runs from the micropyle to the polar end.

Operculum circular; at the anterior end of the dor-

sal surface. Micropylar plate indistinct externally;

internally the plate is almost circular, open, with a

triangular notch at the polar end (fig. 9). Typical

measurements: length 3.8 mm, height 1.5 mm,

width 2.0 mm.

Etymology. — Wegneri, after the collector A.M.R.

Wegner.

Korinnis Günther

Korinnis Günther, 1932: 66. — Type species K. potameis

Günther, 1932 by original designation.

Korinnis potameis Günther

Korinnis potameis Günther, 1932: 67, fig. 1. — Holotype 9,

Kalimantan, Noesa Djangkai on the lower reaches of the

River Serawai, 19-xi-1924.

Korinnis potameis. — Günther 1943: 151.

Material examined. — Kalimantan, Mahakam, ex-

pedition of Dr Nieuwenhuis, 1894 [det. Günther,

1943: 151], 12 (RMNH); same data [det. Günther,

but not mentioned in Günther 1943], 2 nymph

(RMNH); Kalimantan Tengah, Sungai Ratu Miri,

Ratu Miri logging camp, to lights, 22-viii-1993, P.E.

Bragg, 19 (P.E. Bragg, PEB-1999).

Table 1. Measurements of specimens examined in detail

BRAGG: Review of Korinninae

Table 1 gives measurements of the two adult spec-

imens which have been examined. The specimen col-

lected by the author was kept alive for four days but

did not lay any eggs. Colour transparencies (Koda-

chrome 64 ASA film) were taken and used in con-

junction with the preserved specimen for the follow-

ing description of the coloration of the insect.

Head, pronotum, mesonotum and leading edge of

elytra a glossy mid brown. Spines on the mesonotum

with black tips. Head with a large almost round pea-

green spot between the eyes. Sides and underneath of

body, and whole of abdomen, blueish-green; final

three abdominal segments with a chocolate brown

longitudinal stripe. Costal and subcostal areas of wing

blueish-green, radial and medial areas pea-green; radi-

al vein distinctly orangy-brown with a narrow white

line on the hind margin of the subcostal area. The

anal region of the wing pale rose-pink. Elytra blueish-

green on the leading edge, glossy mid-brown on the

humped portion, pea-green on the outer trailing edge

and bright yellow on the inner trailing edge. All fe-

mora, fore tibiae and fore tarsi glossy mid-brown, fore

femora yellowish green at the base. Mid and hind tib-

iae very pale green, brown at apices. Mid and hind

tarsi very pale green at the base becoming mid brown

at the apices. Antennae very dark brown, almost

black, with some pale bands towards the apices.

Remarks. — Maps of Kalimantan are of limited val-

ue for finding precise localities, they are often marked

‘data incomplete’ and not all rivers and villages are

named. Attempts to locate Noesa Djangkai on avail-

able maps have been unsuccessful, however a village

called Nanga Serawai has been located on the conflu-

ence of the rivers Melawei and Serawai; this suggests

Total >28

Length (mm) 3 pulchella

Antennae >26

Head 2.5

Pronotum 1.5

Mesonotum 4

Metanotum & median segment 6.5

Elytra IS

Wing 21

Fore femora 6.5

Fore tibiae 5

Fore tarsi 5

Mid femora 5

Mid tibiae 4

Mid tarsi 4

Hind femora 8

Hind tibiae 6.5

Hind tarsi 5

Kalocorinnis Korinnis

9 pulchella 3 wegneri 9 potameis © errans

38 53-57 58-59 48

>16 >6 25 >13

3 4.5-5 4 3.5

? 2 2.5 1.5

5 5.5-6 6.5-7 4.5

5 10-11 9-10 8

2.5 4-4.5 4.5-5 4

27 40 40-41 35

Y 9 10.5-11.5 9.5

4.5 7 7-9 7,

4.5 6 6-7 6

5 VS 7.5-8.5 7

3.5 5 5-6.5 5

3.5 - 4.5-5.5 4.5

8 10.5-11 11.5-13 11

6 9 8-10.5 8

5 7 5.5-7 6

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

that Noesa Djangkai is probably in the region around

0° 15°S, 112° 45°E. The specimens from Dr. Nieu-

wenhuis’ expedition give ‘Mahakam’ as the locality;

this is rather vague as the Mahakam, in eastern Kali-

mantan, is one of the longest rivers in Borneo.

Neither the Ratu Miri logging camp, or the River

Ratu Miri are marked on any available maps; the

camp is estimated to be approximately 0° 30’S, 113°

35E.

Korinnis errans Günther

(fig. 6)

Korinnis errans Günther, 1938: 125. — Holotype 9, Borneo?

[Günther stated that the specimen was incorrectly label-

led ‘Sibsagar, Nordost Assam, S.E. Peal’] (NZSI).

Material examined. — [from Siebers’ collection],

1® (RMNH); Sabah, Mt Kinabalu, Silau Silau trail, c.

1580m, 10.xi.1991, C. L. Chan (C. L. Chan) 9;

Sabah, Sepilok, 13.vi.1982, C. L. Chan & S. Nagai,

12 (C. L. Chan).

The female from RMNH which has been examined

has no original data label other than the number 181

written in pencil. However there are two labels added

by Klante in 1965; the first is a determination label,

the second is a note that the specimen is ‘without

doubt from box 39 of the Siebers collection’. The

RMNH collection contains a large number of phasmids

collected in Borneo by Siebers in 1925. This supports

Günther’s suspicions about the locality of the holo-

type. This is confirmed by the two specimens in the

collection of Mr C.L. Chan of Kota Kinabalu, Sabah.

The RMNH specimen agrees with Günthers de-

scription. Although the total length is only 3 mm

shorter, there are three measurements notably differ-

ent from those given by Günther: the length of the

mesonotum is shorter (4.5 mm compared to 6.5 mm

in Günther’s description), the fore femora are shorter

(9.5 mm compared to 11 mm), and the elytra are

larger (4 mm compared to 3 mm); other size differen-

ces are minor. The length of the fore femora agree if

it is assumed that Günther included the trochanter in

his measurement. A complete set of measurements of

Siebers’ specimen is given in table 1, and the speci-

men is illustrated in figure 6.

The two specimens from the collection of C.L.

Chan appear to differ considerably from each other.

The Mt Kinabalu specimen, length 40 mm, is much

smaller than the other specimens although the gener-

al proportions agree closely with the RMNH speci-

men. The Sepilok specimen, length 48 mm, has a

more slender appearance due to the longer mesotho-

rax (5.5 mm), and longer legs; its proportions agree

closely with those given by Günther.

In most respects these specimens agree closely with

Günther’s description; the differences may be due to

intraspecific variation. There is a possibility that there

are two species represented here but at present insuf-

ficient material is available to determine this with

confidence.

It is worth noting that although the other members

ofthe Korinninae have a few hairs on the radius of the

wing, these specimens have numerous hairs all over

the costal region; although those of the Sepilok speci-

men are less densely packed than the RMNH and

Sepilok specimens.

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AND ADJACENT COUNTRIES

Andersen, N.M. & P.P. Chen, 1995. A taxonomic revision of the ptilomerine genus

Rhyacobates Esaki (Hemiptera: Gerridae), with five new species from China and adjacent coun-

tries. — Tijdschrift voor Entomologie 138: 51-67, figs. 1-37, table 1. [ISSN 0040-7496].

Published 15 June 1995.

The taxonomy of the lotic-adapted water strider genus Rhyacobates Esaki, 1923, is revised, all

previously known species redescribed, and five new species described from China and adjacent

countries: R. abdominalis sp. n. from Guangdong, À. edentatus sp. n. from Guangdong and

Guangxi, À. recurvus sp. n. from Jiangxi, R. scorpio sp. n. from Sichuan, all in China, and R.

malaisei sp. n. from Burma, Thailand, and China (Yunnan). R. esakii Miyamoto & Lee, 1963,

syn. n. is synonymized with À. chinensis Hungerford & Matsuda, 1959. A key to all species of

the genus is provided. The status of ‘Rhyacobates imadatei Miyamoto, 1967, is discussed.

Finally, the results of a cladistic analysis of relationships between species of Rhyacobates and re-

lated genera of Ptilomerinae are given.

Correspondence: Dr. Nils M. Andersen, Zoological Museum, Universitetsparken 15, DK-

2100 Copenhagen, Denmark

Key words. — Gerridae; Ptilomerinae; Rhyacobates; taxonomy; key; new species; China, Korea,

Burma, Thailand.

The subfamily Ptilomerinae Bianchi, 1896 com-

prises medium-sized or large water striders which all

are strongly adapted to life on the water surface of lot-

ic habitats including torrents and fast-flowing streams

and rivers. Except for one genus from Madagascar,

the ptilomerines are confined to the eastern Palearctic

Region and the Oriental Region including New

Guinea (Andersen 1982). Matsuda (1960) presented

a very useful account of the taxonomic morphology

and generic taxonomy of the Gerridae including most

genera of the Ptilomerinae. Apart from the taxonom-

ic revision by Hungerford & Matsuda (1965) of the

large genus Ptilomera Amyot & Serville and the revi-

sion of the genus Potamometropsis Lundblad by Zettel

(1994), the species taxonomy of the ptilomerine gen-

era is still incompletely known.

The following account of the genus Rhyacobates

Esaki (1923) is the first part of a series of studies of

the classification and phylogeny of ptilomerine gene-

ra undertaken by the senior author. It is also a contri-

bution towards improving the knowledge about the

fauna of semiaquatic bugs (Heteroptera, Gerromor-

pha) of China and adjacent countries.

All measurements are in millimeters. ‘Length’ has

been measured from the anterior margin of the head

along the central body axis to the posterior margin of

the last visible abdominal tergum in apterous (wing-

less) specimens, or to the apex of the forewings in

macropterous (winged) specimens. ‘Dealated’ speci-

mens are macropterous specimens which have shed

their wings by self-inflicted damage or autotomy (see

Andersen 1982). ‘Width’ is the maximum width of

the body and is measured across the middle pair of

acetabula (coxal cavities). Measurement of the femur

does not include the trochanter. Measurements of the

distal leg segments (tibia and/or tarsus) are usually in-

accurate since these segments are extremely thin and

usually coil up when the specimen is dried.

The genital segments of female Rhyacobates (seg-

ments 8-10) are usually completely withdrawn into

the tubular, pregenital abdomen (e.g., figs. 6-7) and

difficult to remove for examination and dissection. In

order to examine the genital segments (segments 8-

10) of the male (fig. 8), the segments are detached

from the abdomen (dry specimens may be softened

by placing them in alcohol for a few hours), macerat-

ed in hot 10% potassium hydroxide (KOH) for about

ten minutes, and cleared in lactic acid (50% aqueous

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

solution) for a couple of hours. The phallic organ is

pulled out of the pygophore (segment 9) and the dis-

tal part of the phallus (vesica) pushed out of the basal

part (phallotheca) with fine needles.

The specimens dealt with in this paper were bor-

rowed from or deposited in the following collections

(with abbreviations used in the lists of material exam-

ined): Entomological Laboratory, Kyushu University,

Fukuoka, Japan (ELKU); the Natural History Museum

(formerly British Museum, Natural History),

London, U.K. (BMNH); the John T. Polhemus collec-

tion, Englewood, Colorado, U.S.A. (JTPC); the Nico

Nieser collection, Tiel, the Netherlands (NC); Natural

History Museum, Stockholm, Sweden (NHMS);

Natural History Museum, Vienna, Austria (NHMV);

Nankai University, Tianjin, China (NUC); Snow Mu-

seum of Entomology, Kansas University, Lawrence,

Kansas, U.S.A. (SEMK); National Museum of Natural

History, Smithsonian Institution, Washington,

D.C., U.S.A. (USNM); Zoological Institute, Acade-

mia Sinica, Beijing, China (ZIB); Zoological

Museum, University of Copenhagen, Denmark

(ZMUC).

SYSTEMATIC PART

Genus Rhyacobates Esaki

Rhyacobates Esaki, 1923: 387 (descr.). Type species by ori-

ginal designation: Rhyacobates takahashii Esaki, 1923;

Hungerford & Matsuda, 1960: 8 (key); Matsuda, 1960:

273-276; Andersen, 1982: 424 (key; many further refe-

rences throughout book).

Esakobates Lundblad, 1934: 22. Type species by monotypy:

Esakobates svenhedini Lundblad, 1934. Synonymized by

Hungerford & Matsuda (1959: 69).

Description. — Chiefly black dorsally and pleurally,

covered by silvery pubescence (fig. 1). Head yellowish

with median black spot. Pronotum with large median

pale spot; pronotal lobe (macropterous form) dark

with brownish lateral margins. Mesonotum (apterous

form) with median pale stripe not reaching anterior

margin. Mesosternum chiefly black, rest of venter yel-

lowish.

Female larger than male, not modified ventrally.

Head with anterior margin not bent ventrad; eyes

globular. Antenniferous tubercles prominent; anten-

nae almost as long as body in male, slightly shorter in

female: first antennal segment longer than three fol-

lowing segments together (fig. 2); second segment

not longer than third; fourth segment shortest,

curved and with elongate, impressed whitish hair-pad

ventrally on distal two fifths (fig. 3). Pronotum of

apterous form short, subquadrangular, with lateral

margins slightly converging posteriorly. Mesonotum

about three times as long as pronotum in male, slight-

ly longer in female. Pronotum of macropterous form

large, covering mesonotum and wing bases; posterior

margin broadly rounded. Fore leg longer than body

(fig. 1); femur robust, slightly tapering apically; tibia

with inner apical process more distinct in female than

in male; first tarsal segment 1.6-2x as long as second

segment in male, longer in female. Middle femur

about twice or a little less than twice as long as tibia;

femur in male without dense fringe of long hairs on

inner margin, with scattered dark pegs or spines ven-

trally, not forming a distinct row of spines; middle

tibia about half as long as femur, with distinct hair-

fringe ventrally; middle tarsus less than half as long as

tibia, first tarsal segment much longer than second

segment. Hind coxae longer than wide, without spine

on apical margin; hind femur slightly longer than

middle femur; hind tibia about one third as long as

femur, without hair-fringe; hind tarsus less than one

tenth as long as tibia, first tarsal segment shorter than

second segment. Claws absent from both middle and

hind legs. Forewing venation very similar to that of

Ptilomera (Andersen, 1982: fig. 430), with M and Cu

separated from near base, connected by a cross-vein

before middle of wing.

Male abdomen shortened, seventh segment about

1.5x sixth segment ventrally. Eighth segment reduced

in length ventrally (fig. 8), hind margin slightly pro-

duced. Pygophore prolonged, rounded on apical mar-

gin. Proctiger widened (fig. 10), sometimes expanded

laterally in basal parts. Parameres large (fig. 9), sym-

metrical and falciform, not conspicuously setose.

Phallotheca cylindrical but weakly sclerotized; con-

junctivum with two pairs of lobes; endosoma with

dorsal vesical sclerite only partly sclerotized (fig. 11),

bifurcated apically and fused to slender ventral scler-

ite basally; there are no well-defined lateral sclerites.

Female abdomen moderate in length in most spe-

cies, posterior segments curved dorsad (fig. 6; apter-

ous form), in some species to oblique or even vertical

position. Seventh segment about twice or over twice

as long as sixth segment ventrally (fig. 4), tubular,

prolonged and enclosing genital segments. Connexi-

vum with postero-lateral corners more or less distinct-

ly prolonged. Hind margin of seventh sternum pro-

duced medially but without distinct median lobe.

Ovipositor illustrated by Matsuda (1960: fig. 680, À.

lundbladi).

Type species by original designation and monoty-

py: Rhyacobates takahashii Esaki, 1923.

Biology. — Species belonging to the genus

Rhyacobates live on the surface of swift streams or

small rivers. Esaki (1923: 389-390) aptly characteriz-

es the habitat and behaviour of Rhyacobates takahashii

as follows: ‘This curious water strider was found at

first on a very rapid stream in a rocky ravine at Sozan

MOLLER ANDERSEN & CHEN: Revision of Rhyacobates

EEL = e D nn |

— o

SSI

Fig. 1. Rhyacobates scorpio; apterous male, dorsal habitus (Reproduced with permission from Andersen, 1982: fig. 448).

“TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

[in Taiwan]. The insects glide swiftly on the surface

of the water in all directions and are hardly recog-

nizable owing to the disturbance of the water. They

look, however, somewhat like whirligig beetles. Some

of them were found climbing on the rocks near by.

Numerous of specimens in copula were also captured.

Numerous examples were found on the Shinten River

which is a very much larger stream than the Sozan.

Here also they live on a rapid current, but at Shinten

only the males and the nymphs of the last instar were

found. The nymphs were not seen at Sozan.’

The last observation seems to indicate that the spe-

cies is protandrous, i.e., that males complete their last

moult before the females. The modifications of the

female abdomen with genital segments withdrawn

into a tubular seventh abdominal segment may be an

adaptation to discourage copulation except when the

female is newly emerged and teneral.

Comparative notes. — Rhyacobates is most closely

related to Heterobates Bianchi (1896) and Pleciobates

Esaki (1930). Matsuda (1960: 264-265) pictured

these genera as a (presumably monophyletic) group

with unresolved mutual relationships in his diagram

of ‘evolutionary relationships of genera of the

Ptilomerinae. The genus Potamometroides Hunger-

ford (1951) from Madagascar was also included but is

not considered in the following discussion.

Rhyacobates shares the following characters with

Heterobates and Pleciobates: (a) meso- and metanotum

chiefly dark, covered with a dense layer of silvery pu-

bescence; (b) first antennal segment much longer

than three following segments together; (c) fourth an-

tennal segment short, curved and with elongate ovate,

impressed whitish area in distal two fifths to half, (d)

male fore femora relatively slender, without ventral

modifications before apex; (e) claws absent from mid-

dle and hind tarsi; (f) male parameres moderate in

size, falciform, not conspicuously setose; (g) apical

segments of female pregenital abdomen strongly

modified. According to Matsuda (1960: 204), the ba-

sally strongly reflexed connexiva (female abdomen)

also unites the three genera. This character, however,

is not present in all Rhyacobates species.

Rbyacobates differs from both Heterobates and

Pleciobates in the following characters: (h) middle fe-

mur of male without ventral row of black spines; (i)

eighth abdominal segment of male reduced ventrally,

shorter than seventh sternum; (j) posterior segments

of female abdomen usually curved dorsad (apterous

form); (k) seventh segment of female abdomen tube-

like prolonged, completely enclosing the genital seg-

ments.

The genus Heterobates is above all characterized by

the distinctly demarcated, flattened area on the ven-

tral surface of the female body (Matsuda 1960: fig.

684). The hind margin of female abdominal sternum

7 has a square, median lobe (absent in Rhyacobates

and Pleciobates).

The genus Pleciobates was not available to Matsuda

(1960) for direct study but its affinity was discussed

based on the original description by Esaki (1930). In

females of the type species, P. tuberculatus Esaki,

1930, the seventh abdominal segment is prolonged,

with a finger-like process arising from the hind corner

of the sixth connexival segment (see also Cheng &

Fernando 1969). More recently, Thirumalai (1986)

has described two species of Pleciobates from southern

India in which the females lack this connexival pro-

cess.

Key to the species of Rhyacobates

(male of R. recurvus sp. n. not available)

ldRemalesf Leti eo e o 2

i MAIS ET te Tee MELIA 10

2. Seventh abdominal segment abruptly turned dor-

sad at an angle of about 90 degrees (fig. 28).

Posterolateral corners of segment with very short

process (figs. 29-30). China (Sichuan) ...............

Dee R. scorpio sp. n.

— Seventh abdominal segment not modified as

above asen e RER URN LE 3

3. Abdomen distinctly curved dorsad towards the

end (figs. 6, 12, 16, 33). Posterolateral corners of

seventh abdominal segment either blunt or trian-

gularly produced, but never bilobate … … … … 4

— Abdomen almost straight or only faintly curved

dorsad towards the end (fig. 18). Posterolateral

corners of seventh abdominal segment more or

lessidlistinetyabilobate ns 8

4. Dorsal margin of seventh abdominal segment dis-

tinctly shorter than ventral margin when viewed

from the side (fig. 16). Posterior margin of sev-

enth abdominal sternum with median, recurved

process (fig. 17). China (Guangdong) ................

a ae ei R. recurvus sp. n.

— Dorsal margin of seventh abdominal segment al-

most as long as ventral margin when viewed from

the side (figs. 6, 12, and 33). Seventh abdominal

sternum at most with a median, pointed but not

recuivedi process (el) re 5

5. Posterior margin of seventh segment (figs. 34-35)

without any projections. China (Guangdong,

Guanda R. edentatus sp. n.

— Posterior margin of seventh segment with one or

two pairs of lateral projections and one median,

ventral'processü...:.. ttes er tn 6

6. Posterior margin of seventh segment (fig. 12)

with a blunt, angular projection above laterad of

which is a faint process (fig. 13). China (Guang-

dong) ito R. abdominalis sp. n.

— Posterior margin of seventh segment with one or

10.

12.

15;

14.

two pairs of prominent, lateral projections. Dor-

sal margin of segment usually furnished with a

rowiofierectidarkdhrairspes sets 7

. Posterior margin of seventh segment (fig. 4) with

one triangular projection above which is directed

obliquely downward. First abdominal tergite

swollen. Length 9.1-9.5 mm. Taiwan

pale tals ne à TR he LEN RER R. takahashii Esaki

Posterior margin of seventh abdominal segment

(figs. 6-7) with two projections: one short, trian-

gular projection above laterad of which is a long-

er, postero-ventrally directed projection. First ab-

dominal tergite with two, distinctly separated

swellings. Length 8.0-9.3 mm. China (Neimeng-

gu, Hebei, Hubei, Hunan, Tianjin), Korea … …

EAN R. chinensis Hungerford & Matsuda

Small species, length 7.0-7.6 mm. Posterior mar-

gin of seventh abdominal segment with two pro-

jections (figs. 18-19): one long, triangular projec-

tion above laterad of which is a short, pointed

process. Burma, China (Yunnan), Thailand … …

Pee bode enen R. malaisei sp. n.

Larger species, length 11-12.2 mm. Posterior end

of abdomen not modified as before … … … … …. 9

Posterolateral corners of seventh abdominal seg-

ment (figs. 26-27) with two lobes; inner lobe nar-

row and pointed, outer lobe broad and rounded

at tip. Length 12.2 mm. China (Zhejiang) .........

Ee nnen Rhyacobates lundbladi Hungerford

Posterolateral corners of seventh abdominal seg-

ment as above, but inner lobe very small (figs. 24-

25) and outer lobe broad and obliquely cut off at

tip. Length 11-12 mm. China (Fujian, Sichuan)

ENE Rhyacobates svenhedini Lundblad

Small species, length 6.0-7.2 mm .................. 11

Large species, length 7.5 mm or more ........... 13

Middle and hind femora subequal in length.

Proctiger with small, hook-shaped process on

each side (fig. 21). Length 6.0-6.5 mm ..............

NEE rite Ah Portes ILE R. malaisei sp. n.

Middle femur shorter than hind femur. Proctiger

laterally widened (fig. 10), but without hook-

shapedsprocessesiin Ben PA. nue 12

Parameres rather broad in middle one-third, with

apices more or less suddenly narrowed (fig. 9) …

TEN R. chinensis Hungerford & Matsuda

Parameres slender and falciform, with distinctly

hook-shaped apices (as viewed from behind) ......

RR R. takahashii Esaki & R. abdominalis sp. n.

Eighth segment distinctly impressed ventrally,

with transverse ridge in basal part. Length 8.4

DOD A AL ia R. svenhedini (Lundblad)

Eighth segment not modified ventrally as before

RE ARIA REA Aura «Brin SUES de duna Soria: 14

Eensch7.5:mm...... R. lundbladi (Hungerford)

Kensth18.0-8°lemmer mn er R. scorpio sp. n.

MOLLER ANDERSEN & CHEN: Revision of Rhyacobates

Rhyacobates takahashii Esaki

(figs. 2-5)

Rhyacobates takahashii Esaki, 1923: 388, pl. 1. Holotype fe-

male [not examined] from Taiwan, Sosan near Taihoku

(ELKU); Esaki, 1925: 60; Matsuda, 1960: 273-276, figs.

656-658, 660, 663, 674-676, 678.

Material. — TAIWAN: 2d 2% (apterous), Formosa,

Keishinryo, 15.iv.1965, leg. T. Saigusa (det. S. Miyamoto)

(rec); 1d (apterous), Ural, Taipei, Taiwan, July 1957, leg.

L.C. Chen (TPC); 16 19 (apterous), Taiwan, Shinten near

Taihoku, 7.vi.1940, leg. S. Miyamoto (R. takahashii, det.

Esaki) (USNM).

Descriptive notes. — The original description

(Esaki, 1923) of the apterous male and female of À.

takahashii is excellent and well illustrated, and

leaves no doubt about the identity of this species.

The length is given as 6.5 mm (male) and 9.5 mm

(female). Matsuda (1960) offers useful illustrations

of both sexes of this species including details of

head, appendages, male genitalia, and female abdo-

men.

The apterous specimens examined by us have the

following dimensions: length 6.8, 2 9.1; width of

head d 1.2 mm, @ 1.4; width of thorax d 1.8, ©

2.6. Colour pattern as in generic description.

Antennal segments 1-4 of male measure: 3.52 : 0.98 :

1.05 : 0.82. Relative lengths of leg segments (femur:

tibia: first tarsal segment: second tarsal segment): fore

leg: 3.60 : 3.00 : 1.00 : 0.62; middle leg: 10.05 : c. 4.9

: c. 2.0 (segment 1+2); hind leg: 10.92 : c. 3.2 : 0.10

:0.20.

Male genital segments have the following structure

(Matsuda 1960: figs. 658, 674): eighth segment

slightly impressed ventro-laterally; pygophore large,

ovate; proctiger with small, angular projection on

each side; parameres (fig. 5) slender and falciform,

with distinctly hook-shaped apices (as viewed from

behind). Miyamoto & Lee (1963: fig. 5) illustrated

the vesical sclerites and Miyamoto (1967: fig. 58) the

structure of the male endosomal conjunctivum.

Abdomen of apterous female curved upward to-

wards the end. First abdominal tergite with a me-

dian, yellowish brown swelling. Connexiva reflect-

ed, margins of third to fifth segment almost meeting

each other above the tergites. Seventh abdominal

segment slightly longer than two preceding seg-

ments together ventrally; dorsal margin describing a

regular curve when viewed from the side (fig. 4),

furnished with dense, erect pubescence; posterior

margin with a triangular projection above which is

directed obliquely downward; median, ventral pro-

cess small but sharp, continued as a low carina on

seventh sternum.

Esaki (1925: 60, figs. 11-12) described and illus-

trated the macropterous form.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

RARES

Figs. 2-11. — 2-5. Rhyacobates takahashii, 2 left antenna, 3 fourth antennal segment, ventral view, 4 abdominal end of female,

lateral view, 5 left paramere of male, two different views. Scales 2, 4 0.5 mm, 3 0.2 mm, 5 0.1 mm. — 6-11. Rhyacobates chi-

nensis, 6 female body, lateral view (mn mesonotum, pn pronotum, t1 first abdominal tergum), 7 abdominal end of female,

lateral view (s7 seventh abdominal sternum), 8 abdominal end of male, ventral view (pa paramere, py pygophore, s7 seventh

abdominal sternum, s8 eighth abdominal sternum), 9 left paramere of male, two different views, 10 proctiger of male, dorsal

view, 11 vesical sclerites of male, lateral view (ds dorsal sclerite, vs ventral sclerite). Scales 6 1 mm, 7, 8 0.5 mm, 9, 11 0.1 mm,

10 0.2 mm.

Rhyacobates chinensis Hungerford & Matsuda

(figs. 6-11)

Rhyacobates chinensis Hungerford & Matsuda, 1959: 69.

Holotype [not examined] from China, Neimenggu

[Inner Mongolia], Djalantun, Great Khingan Mts.

(SEMK).

Rhyacobates esakii Miyamoto & Lee, 1963: 43. Holotype male

[examined]: Korea (S), Nam Te Cheon (ELKU). Syn. n.

Rhyacobates takahashii Esaki; Esaki, 1940: 128 (Miyamoto

& Lee, 1963: 43)

Material. — CHINA: Paratypes 1d 19 (apterous)(R. chi-

nensis), Djalantun, Gr. Khingan, VI.38, Bot. from M.

Weymarn (JTPC). — Hebex 76 319 (apterous), 12 (mac-

ropterous), Wu-Ling Shan (40.6N, 117.4E), VIII.29.1973,

leg. Liu Shengli (NUC, ZMUC); 6% (apterous), same locality,

VIII.1974 (nuc); Huber: 19 (apterous), Shen-Nong-Jia

Nat. Res., Song-Bai (31.7N, 110.6E), VII.17.1977, leg.

Zou Huan-guang (NUC); Hunan: 68 69 (apterous),

Zhang-jia-jie Nat. Res. (29.1N, 110.4E), Jin-bian Stream,

X.14.1985, leg. Zou Huan-guang (NUC); 1d (apterous),

NW-Hunan, Bez. Dayong, Wulingvuan, Zhiangjiajie,

Forest NP, Shuiraosinum, X.30.1993, 600 m, leg. H.

Schönmann (NHMV); Neimenggu [Inner Mongolia]: 11ó

10 (apterous), Xilin Gol Meng (43.9N, 116.0E), livestock

breeding farm, VIII.1974, leg. Jiang Zhong-ming (NUC);

Tianjin: 28 32 (apterous), Ji County (40.0N, 117.3E),

Xia-Ying Xiang, Chang-zhou Village, VII.29, 1985, leg. Liu

(NUC). KOREA: Holotype d (apterous) (R. esakii): Nam Te

Cheon, Sam Bangcheon, Ham Nam, 31.vii.1937, leg. K.

Doi (ELKU); 3d 39 (apterous), S. Korea, Seolak Mt.

8.9.1976, collector unknown (ex Lee coll.) (R. chinensis, det.

P. Chen & J.T. Polhemus) (JTPC).

Description. — Apterous form: Dimensions.

Length d 6.4-6.8, 2 8.0-9.3; width of head d 1.2-

1.3 mm, ® 1.4-1.5; width of thorax d 1.9-2.1, ©

2.8-3.2.

Colour as in generic description. Median black

spot of head posteriorly bifurcated. Antennae dark

brown. Pronotum with a large, triangular brownish

yellow spot in middle. Median, brownish yellow

stripe of mesonotum extends onto metanotum and

first abdominal tergite in female. Connexival margin

brownish in female. Prosternum, legs and distal parts

of acetabula light brown. Mesosternum chiefly dark

with a median subtriangular, yellowish spot.

Male elongate with relatively short abdomen.

Relative lengths of antennal segments (1-4): 3.50 :

0.93 : 0.98 : 0.83. Relative lengths of leg segments

(femur: tibia: first tarsal segment: second tarsal seg-

ment); fore leg: 3.50 : 3.00 : 0.90 : 0.56; middle leg:

10.30 : 5.80 : c. 3.5 (1st + 2nd tarsal segment); hind

leg: 11.20 : c. 3.6 : 0.11 : 0.21. Eighth segment ven-

tro-laterally impressed (fig. 8); pygophore large,

ovate; proctiger (fig. 10) with a small angular projec-

tion on each side. Parameres (fig. 9) rather broad in

middle one-third, with apices more or less suddenly

narrowed. Phallic conjunctivum with two pairs of

lobes, of which the anterior one is sclerotized on dor-

MOLLER ANDERSEN & CHEN: Revision of Rhyacobates

sal wall; dorsal sclerite of vesica partly sclerotised (fig.

11), with a hook-shaped distal part; ventral sclerite

short and elongate leaf-like (see also Miyamoto &

Lee, 1963: 44 and fig. 2).

Female robust. Relative lengths of antennal seg-

ments (1-4): 4.50: 1.15 : 1.20 : 0.93. Relative lengths

of leg segments (femur: tibia: first tarsal segment: sec-

ond tarsal segment); fore leg: 4.40 : 3.80 : 1.75 : 0.93;

middle leg: 12.30 : 7.20 : c. 3.9 (1st + 2nd tarsal seg-

ment); hind leg: 13.00 : c. 5.0 : 0.16 : 0.24. Abdomen

relatively short, with distal half distinctly curved up-

ward to oblique (fig. 6) or nearly vertical position.

First abdominal tergite swollen and with a pair of

prominent, widely spaced tubercles. Connexiva erect

on first two segments, converging along dorsal mid-

line of third through sixth segments. Tube-like sev-

enth abdominal segment (fig. 7) as long as three pre-

ceding segments together ventrally; dorsal margin

describing a regular curve; posterior margin with a

short, triangular projection above terminating each

connexivum, laterad of which is a longer, postero-

ventrally directed projection; median, ventral process

slender and sharp, sometimes continued as a low, lon-

gitudinal carina on ventral surface of seventh ster-

num.

Comparative notes. — Miyamoto & Lee (1963) de-

scribed Rhyacobates esakii from Korea, seemingly

without knowing À. chinensis Hungerford & Matsu-

da (1959) described from Neimenggu, China. A

comparison between a pair of paratypes of the former

species and the holotype male and a topotypical fe-

male of the latter species, lead us to the conclusion

that they are conspecific. R. chinensis can be separated

from À. takahashii by the slightly larger size, paired

swellings of the first abdominal tergite of female,

more distinctly curved female abdomen, and especial-

ly by having five instead of three apical projections on

the seventh abdominal segment of the female. The

parameres of R. chinensis males are more robust and

less curved, and the structure of the vesical sclerites is

different (see Miyamoto & Lee, 1963: figs. 2 and 5).

While males of R chinensis are rather uniform in

structure throughout the geographical range of the

species, the shape and relative size of the apical pro-

jections of the female abdomen seem to be quite var-

iable although characteristic of the species.

So far, R. chinensis has been recorded from Nei-

menggu Province [Inner Mongolia] (Hungerford &

Matsuda, 1959), Hebei, Hubei, and Hunan

Provinces, China, and from North, central and South

Korea (Miyamoto & Lee, 1963). The records by

Esaki (1940) of R. takahashii from Kullin, Kiangsi

[Jiangxi] and Jehol, Manchuria [Heilongjiang], both

localities in China, and from Korea, probably refer to

R. chinensis.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Rhyacobates abdominalis sp. n.

(figs. 12-15)

Type material. — CHINA: Holotype © (apterous):

Guangdong Province, Ruyang Nat. Res., Lao-Peng Stream,

1100 m, VIII.14. 1990, leg. P.P. Chen, C9012 (NUC). —

Paratypes 96 89 (apterous), same label data as above (NC,

NHMV, NUC, ZMUC).

Description. — Apterous form. Dimensions. Length

d 6.2-6.8, 2 8.6-9.3; width of head d_1.2-1.3, 2

1.47-1.52; width of thorax 1.9-2.2, 9 3.1-3.3.

Colour as in generic description. Median black

spot of head posteriorly bifurcated. Antennae dark

brown to blackish. Pronotum with a large, triangular

brownish yellow spot in middle. Median, brownish

yellow stripe of mesonotum extends onto metanotum

and first abdominal tergite in female. Connexival

margin brownish yellow in female. Prosternum, legs

and distal parts of acetabula light brown. Mesoster-

num chiefly dark with a median subtriangular, yel-

lowish spot.

Male elongate with relatively short abdomen.

Relative length of antennal segments (1-4): 3.15 :

0.89 : 1.05 : 0.84. Relative lengths of leg segments

(femur: tibia: first tarsal segment: second tarsal seg-

ment); fore leg: 3.37: 2.64 : 0.95 : 0.62; middle leg:

9.20 : 5.20 : c. 3.0 (lst + 2nd tarsal segment); hind

leg: 8.98 : c. 3.84 : 0.12 : 0.18. Eighth segment ven-

tro-laterally impressed; pygophore large, ovate; proc-

tiger with a small angular projection on each side.

Parameres (fig. 15) relatively slender, broad on basal

2/5, tapering towards hook-shaped apices.

Figs. 12-17. 12-15. Rhyacobates abdominalis, 12 female body, lateral view, 13 abdominal end of female, lateral view, 14 ab-

dominal end of female, ventral view, 15 left paramere of male, two different views. Scales 12 1 mm, 13-14 0.5 mm, 15 0.1

mm. — 16-17. Rhyacobates recurvus, 16 female body, lateral view, 17 abdominal end of female, lateral view. Scales 16 1 mm,

17 0.5 mm.

Female robust. Relative length of antennal seg-

ments (1-4): 4.06 : 1.10 : 1.31 : 0.96. Relative lengths

of leg segments (femur: tibia: first tarsal segment: sec-

ond tarsal segment): fore leg: 4.24 : 3.51 : 1.95 : 0.91;

middle leg: 12.55 : 7.09 : c. 4.23 (1st + 2nd tarsal seg-

ment); hind leg: 12.38 : c. 6.1 : 0.14 : 0.18. Abdomen

relatively short, with distal half distinctly curved up-

ward to oblique position (fig. 12). First abdominal

tergite slightly swollen. Connexiva erect, only con-

verging distally along mid-line of seventh segment, or

never converging. Tube-like seventh abdominal seg-

ment (fig. 13) ventrally as long as two preceding seg-

ments together; dorsal margin describing a regular

curve; posterior margin with a short, angular projec-

tion above terminating each connexivum, laterad of

which is a blunt projection; median ventral process

(fig. 14) slender and sharp, sometimes continued as a

low, longitudinal carina on ventral surface of seventh

sternum.

Etymology. — abdominalis refers to the structure of

the female abdomen.

Comparative notes. — Females of R. abdominalis sp.

n. share the distinctly curved abdomen with the two

preceding species, but the projection terminating

each connexivum is much shorter. The median pro-

cess of seventh sternum is slender and sharp, but not

recurved as in the following species.

Rhyacobates recurvus sp. n.

(figs. 16-17)

Type material. — CHINA: Holotype ® (apterous): Lushan

[Lu Shan in Jiangxi Province], 9.VIII.30, leg. Chenfu Wu

(BMNH).

Description. — Apterous form. Dimensions.

Length ® 7.7; width of head ® 1.4; width of thorax

Colour as in generic description. Head with one

large, median spot and two smaller, posterior black

spots. Antennae dark brown to blackish. Pronotum

with a large, triangular brownish yellow spot in mid-

dle. Median, brownish yellow stripe of mesonotum

extends onto metanotum and first abdominal tergite

in female. Connexival margin brownish yellow in fe-

male. Prosternum, legs and distal parts of acetabula

light brown. Mesosternum chiefly dark with a me-

dian subtriangular, yellowish spot.

Female stout. Relative lengths of antennal seg-

ments (1-4): 3.70 : 1.10 : 1.22 : 0.88. Relative lengths

of leg segments (femur: tibia: first tarsal segment: sec-

ond tarsal segment); fore leg: 4.18 : 3.35 : 1.48 : 0.88;

middle leg: 11.50 : ? : 2 : 2; hind leg: 12.12 : c. 4.0: ?

: ?. Abdomen very short, with distal half distinctly

M@LLER ANDERSEN & CHEN: Revision of Rhyacobates

curved upward to oblique or nearly vertical position

(fig. 16). First abdominal tergite slightly swollen.

Connexiva erect on first two segments, converging

along dorsal mid-line of third through sixth seg-

ments, reflexed above abdominal tergites. Tube-like

seventh abdominal segment (fig. 17) longer than

three preceding segments together ventrally; dorsal

margin distinctly shortened; posterior margin angular

above, almost straight below, ventrally produced into

a median, hook-shaped process.

Male unknown.

Etymology. — recurvus refers to the recurved me-

dian process of the female seventh abdominal ster-

num.

Comparative notes. — The short, distinctly curved

abdomen and recurved median process of seventh

sternum separates the female of À. recurvus sp. n.

from those of other Rhyacobates species known to us.

Rhyacobates malaisei sp. n.

(figs. 18-23)

Type material. — BURMA: Holotype d (apterous): N.

Burma, Bumgahtuang-Hpungan, river, 17.11.1934, leg. R.

Malaise (NHMS). — Paratypes 24 49 (apterous), same label

data as holotype (NHMS, ZMUC). THAILAND: 4d 39 (apter-

ous), Chiang Mai Prov., Nam Chai R. above hydro station

intake at Fang Hort. Stat, CL 2197, 15.xi.1985, leg. J.T. &

D.A. Polhemus (JTPC). CHINA: 26 dé (apterous), Yunnan,

100 km W Kunming, Diaolin Nat. Res. V.22-VI.6.1993,

leg. E. Jendek & O. Sousa (NHMV).

Description. — Apterous form. Dimensions.

Length d 6.0-6.5, ® 7.0-7.6; width of head d 1.1-

1.2 9711 9o-Fswidchroßthorax CLEO ROME

DD,

Colour as in generic description. Median black

spot of head posteriorly bifurcated. Antennae dark

brown. Pronotum with a broad, triangular brownish

yellow spot in middle. Median, brownish yellow

stripe of mesonotum does not extend onto metano-

tum and first abdominal tergite. Connexival margin

chiefly blackish, connexival processes of female light

brownish. Prosternum, legs and distal parts of aceta-

bula light brown. Mesosternum chiefly dark with a

median subtriangular, yellowish spot.

Male elongate with relatively short abdomen.

Relative lengths of antennal segments (1-4): 2.70 :

0.78 : 1.05 : 0.75. Relative lengths of leg segments

(femur: tibia: first tarsal segment: second tarsal seg-

ment); fore leg: 2.90 : 2.28 : 0.75 : 0.48; middle leg:

8.60 : 4.45 : 1.85 : 0.35; hind leg: 8.82 : 2.10: 0.10:

0.18. Eighth segment ventro-laterally impressed; py-

gophore large, ovate; proctiger widened (fig. 21),

with a small hook-shaped projection on each side.

DI,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Parameres (fig. 22) rather broad, almost spatulate, in ments (1-4): 2.85 : 0.70 : 0.92 : 0.75. Relative lengths

distal part. Armature of phallic vesica as illustrated of leg segments (femur: tibia: first tarsal segment: sec-

(fig. 23). ond tarsal segment); fore leg: 2.80 : 2.30 : 1.10 : 0.72;

Female elongate. Relative lengths of antennal seg- middle leg: 8.20 : 4.65 : 1.92 : 0.28; hind leg: 8.10 :

271]

Figs. 18-27. — 18-23. Rhyacobates malaisei, 18 female body, lateral view, 19 abdominal end of female, dorsal view, 20 abdom-

inal end of female, lateral view, 21 proctiger of male, dorsal view, 22 left paramere of male, two different views, 23 vesical

sclerites of male, lateral view. Scales 18 1 mm, 19, 20 0.5 mm, 21, 23 0.1 mm, 22 0.2 mm. — 24-25. Rhyacobates svenhedini,

24 abdominal end of female, dorsal view, 25 abdominal end of female, lateral view. Scale 24, 25 0.5 mm. mm. — 26-27.

Rhyacobates lundbladi, 26 abdominal end of female, dorsal view, 27 abdominal end of female, lateral view. Scale 26, 27 0.5

mm.

2.18 : 0.12 : 0.15. Abdomen relatively long, almost

straight in lateral view (fig. 18). First abdominal ter-

gite not modified. Connexiva erect on first segment,

reflexed upon abdominal tergites, running parallel to

each other for most of the abdominal length, meeting

each other towards the end. Tube-like seventh ab-

dominal segment shorter than two preceding seg-

ments together ventrally; dorsal margin describing a

regular curve (fig. 20); posterior margin (fig. 19) with

a long, triangular projection above terminating each

connexivum, laterad of which is a small, pointed pro-

cess; hind margin of seventh sternum straight, with-

out median process.

Etymology. — Named for the Swedish entomolo-

gist René Malaise, the collector of the type series from

northern Burma.

Comparative notes. — This is the first Rhyacobates

species known from outside China and Korea (the

status of ‘Rhyacobates’ imadatei Miyamoto from

Borneo and Thailand is discussed below). Apterous

females of R. malaisei sp. n. can be separated from

other species by their nearly straight abdomen and

very characteristic structure of the abdominal end.

The shape of the male proctiger and parameres is also

unique.

Rhyacobates svenhedini (Lundblad)

(figs. 24-25)

Esakobates svenhedini Lundblad, 1934: 23-25, fig. 10, plate

2. Holotype male [examined]: China, N.E. Szechuan

[Sichuan] (NHMS).

Material. — CHINA: Holotype d (apterous): Kina

[China], N.O. Szechuan [Sichuan], Sven Hedin Exp. Cır.

Asien, leg. Dr. Hummel (NHMS). — Paratypes 1d 29 (ap-

terous), same label data as holotype (NHMS, JTPC, ZMUC). —

1? (apterous), Fukien [Fujian], S. China, Kienow: Fengio,

leg. T. Maa (JTPC); 3? (apterous), Ku-ling, Sichuan, collec-

tor unknown (ZIB).

Descriptive notes. — Lundblad (1934) gave excel-

lent descriptions and illustrations of both the apter-

ous male and female of R. svenhedini. He gives the

length as about 9 mm (male) and 12 mm (female),

but the apterous specimens examined by us have the

following dimensions: length d 7.5, ® 11.0; width

of head d 1.3, 2 1.6; width of thorax 2.2, 2.7.

Colour pattern (see Lundblad 1934: plate 2, fig. 4) as

in generic description except that the dark mark on

the dorsal head surface is reduced, distinctly bifid or

dissolved in smaller spots posteriorly (Lundblad

1934: fig. 10A). Antennal segments 1-4 of male

measure: 4.46 : 1.18 : 1.22 : 0.97. Relative lengths of

leg segments (femur: tibia: first tarsal segment: second

MOLLER ANDERSEN & CHEN: Revision of Rhyacobates

tarsal segment): fore leg: 4.60 : 3.78 : 1.25 : 0.67;

middle leg: 12.85 : 6.42 : 2.71 : 0.46; hind leg: 12.64

: 3.38 : 0.12 : 0.19.

Male genital segments have the following structure:

eighth segment slightly impressed ventro-laterally;

pygophore large, ovate; proctiger slightly expanded

on each side; parameres slender and falciform

(Lundblad 1934: figs. 10H-I), with hook-shaped api-

ces (as viewed from behind).

Abdomen of apterous female relatively long, mod-

erately curved upward towards the end. First abdom-

inal tergite not swollen. Connexiva reflected with

margins converging posteriorly as in the female illus-

trated by Lundblad (1934: plate 2, fig. 4) or the con-

nexiva are meeting each other above the tergites.

Seventh abdominal segment shorter than two preced-

ing segments together ventrally; dorsal margin only

moderately curved when viewed from the side (fig.

25); posterolateral corners of each connexivum (fig.

24) usually produced into two lobes: one long, poste-

riorly directed outer lobe and a much smaller, point-

ed inner lobe (lacking in one paratype); posterior

margin of seventh sternum angularly produced in

middle, but without a median tooth.

Comparative notes. — When describing Esakobates

svenhedini, Lundblad (1934) compared it with

Rhyacobates takahashii Esaki (1923). The structure of

the female abdomen, however, are quite different in

the two species. R. svenhedini seems to be closest to

the A. lundbladi Hungerford but separated by the

shape and size of the lobes arising from the posterola-

teral corners of each connexivum (compare figs. 25

and 27).

Rhyacobates lundbladi (Hungerford)

(figs. 26-27)

Esakobates lundbladi Hungerford, 1957: 33-36, pls 1-2.

Holotype male [not examined]: China, Zhekiang, Tien

um Shan (SEMK).

Rhyacobates lundbladi (Hungerford); Matsuda, 1960: 274-

276, figs. 659, 661, 671, 672, 677, 679, 680.

Type material. — CHINA: Paratype 1d (dealated macro-

pterous), Tien um Shan, China, 9.21.37, leg. E. Suenson

(JTPC).

Descriptive notes. — The original description of À.

lundbladi Hungerford (1957) is excellent and well il-

lustrated and leaves no doubt about the identity of

this species. The type series is only composed of

macropterous, although dealated, males and females,

which makes it difficult to compare with other

Rhyacobates species which usually are known only in

the apterous adult form. Matsuda (1960) offers useful

illustrations of structural details of head, female abdo-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

men and ovipositor, and male terminalia. Hunger-

ford (1957) gives the following dimensions of the

apterous form: length d 8.4, 2 12.2; width across

pronotum d 2.6, 9 2.64. Colour as in generic de-

scription except as follows for the pronotum of the

macropterous form (Hungerford, 1957: plate 1); pro-

notum black, margined on the sides and rear with yel-

lowish brown, a large triangular or nearly round yel-

lowish brown spot on anterior part of pronotum with

a median line of same colour on pronotal lobe which

may surpass humeri. Antennal segments 1-4 of male

measure: 4.30 : 1.10 : 1.32 : 1.00. Relative lengths of

leg segments (femur: tibia: first tarsal segment: second

tarsal segment): fore leg: 4.40 : 3.52 : 1.32 : 0.84;

middle leg: 12.25 : 6.30 : 2.42: 0.46; hind leg: 12.36

: c. 4.3 : 0.13 : 0.22.

The male genital segments have the following

structure (Hungerford, 1957: plate 2; Matsuda,

1960: figs. 672, 679): eighth segment distinctly im-

pressed ventrally, with transverse ridge basally; pygo-

phore large, ovate; proctiger slightly expanded on

each side; parameres slender and falciform, with

faintly hook-shaped apices.

Abdomen of apterous female relatively long, only

slightly curved upward towards the end. First abdom-

inal tergite not swollen. Connexiva more or less verti-

cal but on last segment reflected and meeting each

other above the seventh tergite. Seventh abdominal

segment shorter than two preceding segments togeth-

er ventrally; dorsal margin almost straight when

viewed from the side (fig. 27); posterolateral corners

of each connexivum (fig. 26) produced into two, pos-

teriorly directed lobes: outer lobe broad and rounded

at tip; inner lobe narrow and pointed; posterior mar-

gin of seventh sternum only slightly produced in mid-

dle.

Comparative notes. — Hungerford (1957) placed

this species in Lundblad’s genus Esakobates together

with E. svenhedini (see above), but later (Hungerford

& Matsuda, 1959) synonymized this genus with

Rhyacobates. The relatively long and moderately

curved female abdomen unites R. lundbladi and sven-

hedini, but the relative size and shape of the connexi-

val lobes is quite different in the two species (compare

figs. 25 and 27). The ventral modifications of the

eighth abdominal segment may be helpful in distin-

guishing the males.

Rbyacobates scorpio sp. n.

(figs. 1, 28-32)

Type material. — CHINA: Holotype 2 (apterous): Sze-

chuan [Sichuan], Song Ch’i, Kuanhsien, 3500-5000’,

August 1938, leg. D.C. Graham (USNM). — Paratypes 1d

6® (apterous), same label data as holotype (JTPC, USNM,

ZMUC); 36 69 (apterous), Song Ch’i Hsien, July 1938, leg.

D.C. Graham (JTPC, USNM); 38 49 (apterous), 1 nymph,

Wen Chuan, 4000-6000’, August 1938, leg. D.C. Graham

(USNM); 48 59 (apterous), Ku-ling, collector unknown

(z1B); 58 3@ (apterous), 1 nymph., Mt. Emei (29.5N,

103.3E), Jie-Yin Temple, VII.14, 1957, leg. Zheng Le-yi

(NUC, ZMUC).

Description. — Apterous form. Dimensions.

Length d 8.0-8.1, 2 9.7-9.9; width of head d 1.5,

Q 1.6-1.7; width of thorax d 2.6-2.7, 9 3.4-3.5.

Colour as in generic description. Median black

spot of head posteriorly bifurcated. Antennae dark

brown. Pronotum with an elongate triangular brown-

ish yellow spot in middle. Median, brownish yellow

stripe of mesonotum extends onto metanotum and

first abdominal tergite of female. Connexival margins

brownish yellow in female. Prosternum, legs and dis-

tal parts of acetabula light brown. Mesosternum

chiefly dark with a median subtriangular, yellowish

spot.

Male rather stout with relatively short abdomen.

Relative lengths of antennal segments (1-4): 4.60 :

1.45 : 1.55 : 1.15. Relative lengths of leg segments

(femur: tibia: first tarsal segment: second tarsal seg-

ment); fore leg: 4.90 : 4.20 : 1.40 : 0.90; middle leg:

12.95726379.2€.3.0:.052: hindlee il 3 502e55E

0.18 : 0.25. Eighth segment ventro-laterally im-

pressed; pygophore large, ovate; proctiger with a nar-

row, angular projection on each side. Parameres slen-

der and falciform (fig. 31), with weakly hook-shaped

apices (as viewed from behind). Armature of phallic

vesica as illustrated (fig. 32).

Female stout with relatively short abdomen.

Relative lengths of antennal segments (1-4): 4.98 :

1.45 : 1.68 : ?. Relative lengths of leg segments (fe-

mur: tibia: first tarsal segment: second tarsal seg-

ment); fore leg: 5.12 : 4.30 : 2.28 : 1.15; middle leg:

13:90: 8:52 > 472 +058; hindileg: 14°42 5:30::

0.20 : 0.28. Abdomen relatively short, curiously

bent upwards towards end (fig. 28). First abdominal

tergite distinctly swollen. Connexiva narrow, erect

on first to sixth segment, reflexed upon seventh ter-

gite. Sixth sternum depressed medially toward pos-

terior margin. Seventh abdominal segment (fig. 29)

dorso-ventrally flattened, about as long as two pre-

ceding segments together, bent upwards at a right

angle to the preceding abdominal segments (fig. 28);

posterior margin (fig. 30) with a short, pointed

process terminating each connexivum; hind mar-

gin of seventh sternum distinctly produced in mid-

dle.

Etymology. — scorpio refers to the peculiar, up-

turned abdominal end of the female, resembling the

way a scorpion carries its tail.

Comparative notes. — Females of À. scorpio sp. n.

are immediately recognized by the very characteristic

shape of the abdomen. Males can be separated from

those of other Rhyacobates species by their larger size

and slender parameres.

\ we Mall tite,

MOLLER ANDERSEN & CHEN: Revision of Rhyacobates

Rhyacobates edentatus sp. n.

(figs. 33-36)

Type material. — CHINA: Holotype ® (apterous):

Guangdong Province, Lian County (24.7N, 112.3E), Yao-

An Xiang, X.28.1962, leg. Zheng Le-yi & Cheng Han-hua

\ n ag 007 il,

DRE (LI STADIO I N EAN A

;

TN,

Figs. 28-36. — 28-32. Rhyacobates scorpio, 28 female body, lateral view, 29 abdominal end of female, lateral view, 30 abdom-

inal end of female, ventral view, 31 left paramere of male, two different views, 32 vesical sclerites of male, lateral view. Scales

28 1 mm, 29, 30 0.5 mm, 31, 32 0.1 mm. — 33-36. Rhyacobates edentatus, 33 female body, lateral view, 34 abdominal end of

female, lateral view, 35 abdominal end of female, ventral view, 36 left paramere of male, two different views. Scales 33 1 mm,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

(NUC). — Paratypes 54 39 (apterous), 16 (macropterous),

1 nymph, same label data as holotype (NUC, ZMUC); 19 (ap-

terous), Guangxi, Bez. Lipu, 120 km S Guilin, 80 km E

Liuzhou, Siuren, XI.12.1993, 360 m, leg. H. Schillhammer

(NHMV).

Description. — Apterous form. Dimensions.

Length d 6.0-6.6, 2 7.6-8.0; width of head d 1.2-

1.3, 2 1.5; width of thorax d 1.7-1.8, 2 3.0-3.1.

Colour as in generic description. Median black

spot of head posteriorly bifurcated. Antennae dark

brown. Pronotum with a large, triangular brownish

yellow spot in middle. Median, brownish yellow

stripe of mesonotum extends onto metanotum and

first abdominal tergite in female. Connexival margin

brownish in female. Prosternum, legs and distal parts

of acetabula light brown. Mesosternum chiefly dark

with a median subtriangular, yellowish spot.

Male elongate with relatively short abdomen.

Relative lengths of antennal segments (1-4): 3.00 :

0.77 : 0.93 : 0.65. Relative lengths of leg segments

(femur: tibia: first tarsal segment: second tarsal seg-

ment); fore leg: 3.18 : 2.63 : 0.95 : 0.53; middle leg:

9.05 : 4.58 : c. 2.3 (1st + 2nd tarsal segment); hind

leg: 9.08 : c. 2.93 : 0.10 : 0.15. Eighth segment ven-

tro-laterally impressed; pygophore large, ovate; proc-

tiger with a small angular projection on each side.

Parameres (fig. 36) slender with broader base; apices

hook-shaped.

Female robust. Relative lengths of antennal seg-

ments (1-4): 3.70 : 1.00 : 1.20 : 0.90. Relative lengths

of leg segments (femur: tibia: first tarsal segment: sec-

ond tarsal segment); fore leg: 3.86 : 3.16 : 1.90 : 0.90;

middle leg: 13.05 : 6.10 : c. 2.50 (1st + 2nd tarsal seg-

ment); hind leg: 13.3 : c. 4.40 : 0.14 : 0.18. Abdomen

relatively short, with distal half distinctly curved up-

ward to oblique or nearly vertical position (fig. 33).

First abdominal tergite not swollen. Connexiva erect

on first two segments, converging along dorsal mid-

line of third through sixth segments. Seventh abdom-

inal sternum (fig. 35) half as long as three preceding

sterna together; dorsal margin straight, posterior mar-

gin slightly arched, without any projections or pro-

cesses (fig. 34).

Etymology. — edentatus refers to the absence of any

projections or processes on the female abdominal

end.

Comparative notes. — The female abdomen of A.

edentatus is relatively short and wide and the seventh

abdominal segment is lacking any of the projections

or processes found in most of the other Rhyacobates

species

Classification of Rhyacobates imadatei Miyamoto

Miyamoto (1967: 238-241) described Rhyacobates

imadatei from Brunei (N. Borneo) and northern

Thailand. Through the courtesy of Drs S. Miyamoto

and O. Tadauchi (ELKU), we have been able to exami-

ne the holotype d and allotype © (both apterous) la-

belled ‘Amo, Brunei, 24.11.1962, leg. G. Imadaté’.

The following brief description will serve to clarify

the taxonomic position of this species.

Dimensions. Length d 6.7-7.2, 2 9.5-10.0; width

of head d 1.8-1.2, 2 1.3-1.4; width of thorax d

21052 10218310)

Colour. Head yellowish brown with median black

markings. Pronotum black with large yellow marking

in middle. Mesonotum all black.

Female bigger than male, not modified ventrally.

Antenna with first segment much longer than the

three following segments together; second segment

distinctly shorter than third segment; fourth segment

curved, with elongate, impressed whitish area in dis-

tal two thirds. Middle femur of male with distinct

row of dark spines along ventral margin. Middle and

hind legs with small but distinct claws. Eighth ab-

dominal segment of male shorter than seventh ster-

num; pygophore prolonged, narrowed posteriorly;

proctiger widened with angular lateral projections.

Parameres long and slender but not conspicuously se-

tose. Female abdomen relatively long. Seventh seg-

ment prolonged but not enclosing genital segments;

connexival corners with a short spinose process.

Seventh sternum flattened but otherwise simple,

without median lobe or process.

The completely dark mesonotum, the distinct row

of dark spines ventrally on middle femur of male, and

the distinct claws on middle and hind tarsus, excludes

R. imadatei from the genus Rhyacobates as defined here.

However, the classification of ‘Rhyacobates imadatei

Miyamoto cannot be settled until the generic taxono-

my of the Ptilomerinae has been further clarified.

PHYLOGENY

A cladistic analysis of phylogenetic relationships

between the species of Rhyacobates was performed

using the parsimony program Hennig86 (Farris

1988). The species ‘ Rhyacobates imadatei (see above)

and the genera Heterobates and Pleciobates were also

included as well as a hypothetical ‘ancestor’ compo-

sed of character states common in other ptilomerine

gerrids (Matsuda 1960; Andersen 1982).

The following characters and character states were

considered:

1. Colour of meso- and metanotum: uniformly

dark (0), dark with median pale stripe (1).

2. Relative length of first antennal segment: sube-

qual to or shorter than three distal segments together

M@LLER ANDERSEN & CHEN: Revision of Rhyacobates

ù Rhyacobates Fig. 37. Cladogram

Ss no, © ss N | È é SD 26 © relationships between

oe x 20 € S we RS RN we x se the species of Rhyacoba-

DE CARS cite OMO, NAMES Oe Ae SS SS se tes, the genera Heteroba-

Ÿ aA N N se > 9 5 © tes and Pleciobates, and

“Rhyacobates’ imadatei.

15-2 10-0 15-0 :

Numbers on internodes

13-1 8-2 refer to apomorphic

11-0 character states. See text

> 12-1 12-2 72 for further explanations.

7-1

8-1

15-1

10-1

4-1

6-1 6-1

9-1

14-1

11-1

Sail

RIA

(0), much longer than the three distal segments to-

gether (1).

3. Ventral surface of fourth antennal segment: with

an elongate ovate, impressed whitish area in distal

two fifths to one half (1), impressed area shorter than

before (0).

4. Ventral row of dark spines on middle femur of

male: present (0), absent (1).

5. Middle and hind tarsal claws: present (0), absent

(1).

6. Ventral length of eighth abdominal segment of

male: longer than seventh sternum (0), shorter than

seventh sternum (1).

7. First abdominal tergite of apterous female: not

modified (0), slightly swollen (1), distinctly swollen

(2).

8. Shape of dorsal abdomen of apterous female (la-

teral view): almost straight (0), distinctly curved dors-

ad towards apex (1), abruptly turned dorsad to nearly

vertical position towards apex (2).

9. Abdominal laterotergites (connexiva) of apte-

rous female: almost parallel throughout (0), reflexed

and converging towards each other along dorsal mid-

line (1).

10. Seventh abdominal segment of female: prolon-

ged, longer than fifth and sixth sterna together, al-

most completely enclosing the genital segments (1),

shorter, at least distal parts of genital segments visible

(0).

11. Posterior margin of seventh abdominal seg-

ment of female: not modified or at most with a small

process (0), modified, with angular projections (1).

12. Posterior margin of seventh abdominal seg-

ment of female (if modified): with one large, angular

projection (0), with one small and one large projec-

tion (1), with one small, angular projection (0).

13. Posterolateral margin of seventh abdominal

segment of female: simple or slightly produced (0),

with triangular process (1).

14. Posterior margin of seventh abdominal ster-

num of female: simple, straight (0), angularly produ-

ced in middle (1), with square median lobe (2).

15. Posterior margin of seventh abdominal ster-

num of female (if angularly produced): simple (0);

pointed (1), pointed and recurved (2).

The character state matrix is shown in Table 1. The

states of all characters were coded as additively (or-

dered) except for the characters 8, 12, and 14 where

the states were coded as non-additively (unordered).

Using the hypothetical ‘ancestor’ as outgroup, the

Hennig86-analysis found only one most parsimoni-

ous tree, 25 steps long (Consistency Index = 0.80 and

Retention Index = 0.86). This tree (or cladogram) is

shown in fig. 37 with synapomorphies (given as char-

acter number and state) inserted on the internodes.

The cladogram supports the monophyly of the genus

Rhyacobates, excluding ‘Rhyacobates’ imadatei. Four

monophyletic species-groups are delimited in

Rhyacobates: (1) svenhedini + lundbladi, (2) malaisei,

(3) abdominalis + recurvus, and (4) takahashii + chi-

nensis + scorpio + edentatus. Finally, the genus

Heterobates is placed as the sister-group of

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Table 1. Character state matrix for the species of Rhyacobates, ‘Rhyacobates’ imadatei, the genera Heterobates and Pleciobates,

and a hypothetical ‘ancestor’. ? = missing observation, - = character not applicable. Further explanation in text.

Character

Taxon

Rhyacobates

takahashii

chinensis

abdominalis

recurvus

malaisei

svenhedini

lundbladi

scorpio

edentatus

R. imadatei

Pleciobates

Heterobates

Ancestor

OO Fe mm

Oe sO TC pd rr

One ei ei id di pi

oooom Ke ee He tv eS

On On i nm on Y

OO On Va mi

Rhyacobates. It must be emphasized, however, that the

relationships between Rhyacobates and other ptilome-

rine genera depicted in fig. 37 are preliminary, await-

ing a more comprehensive cladistic analysis of the

genera of Ptilomerinae.

As here defined, Rhaycobates has a predominantly

East Asian distribution with one species (R. malaisei

sp. n.) found in the border region between SE

China, Burma, and Thailand. Until a more compre-

hensive cladistic analysis of the genera of Ptilo-

merinae is available, speculations about the histori-

cal biogeography of Rhyacobates and its allies are

premature.

ACKNOWLEDGEMENTS

We are grateful to Liang Aiping, Beijing (ZIB); Le-

yi Zheng, Tianjin (NUC); John T. Polhemus,

Englewood, Colorado (JTPC); the late Peter D.

Ashlock, Lawrence, Kansas (SEMK); M.D. Webb,

London (BMNH); R.C. Froeschner, Washington,

D.C. (USNM); Per Lindskog, Stockholm (NHMS);

Herbert Zettel, Vienna (NHMV); and S. Miyamoto

and O. Tadauchi, Fukuoka, Japan (ELKU) for making

types and other material available for study. We also

thanks Dan A. Polhemus, Honolulu, for offering use-

ful comments on an earlier version of the manuscript.

REFERENCES

Andersen, N.M., 1982. The Semiaquatic Bugs (Hemiptera,

Gerromorpha). Phylogeny, adaptations, biogeography,

and classification. — Entomonograph 3: 1-455.

Cheng, L. & C.H. Fernando, 1969. A taxonomic study of

the Malayan Gerridae (Hemiptera: Heteroptera) with

notes on their biology and distribution. — Oriental Insects

3: 97-160.

Esaki, T., 1923. An interesting new water strider from

Formosa. — Philippine Journal of Science 22(4): 387-391.

SCOOONN'VYOOrRFKNN

„m

| mr © NN à © ©

| or OON im

SISTSZSEIDIDEIISESE Rea

OOO? Bey ee ne

SeeSee EEE

OO OFTEN gr er AE

SISISISTISISHSESESHSESE ES

oONOO mn nm

| ©

Esaki, T., 1925. New or little-known water striders from the

Oriental region. Philippine Journal of Science 26(1): 57-

65.

Esaki, T., 1927. Revision of the Ptilomera-group of the

Gerridae, with descriptions of three new species

(Heteroptera). — Eos 3: 251-268.

Esaki, T., 1930. New or little-known Gerridae from the

Malay Peninsula. — Journal of the Federated Malay States

Museum 16: 13-24.

Esaki, T., 1940. Some aquatic and semiaquatic Heteroptera

from China. — Notes d’Entomologie Chinoise 7: 123-

130.

Farris, J.S., 1988. Hennig86, version 1.5. Program and doc-

umentation. — J.S. Farris, Port Jefferson Station, New

York.

Hungerford, H.B., 1951. An interesting new gerrid from

Madagascar (Hemiptera). — Journal of Kansas

Entomological Society 24: 131-133.

Hungerford, H.B., 1957. A new Gerrid from China

(Gerridae, Hemiptera). — Journal of Kansas

Entomological Society 30: 33-36.

Hungerford, H.B. & R. Matsuda, 1958. Another new ge-

neric entity of the Gerridae. — Entomological News 69:

258-260.

Hungerford, H.B. & R. Matsuda, 1959. Synonymy of the

genera Rhyacobates Esaki 1923 and Esakobates Lundblad

1934, and a description of a new species of Rhyacobates

from China (Hemiptera: Gerridae). — Journal of Kansas

Entomological Society 32: 69-72.

Hungerford, H.B. & R. Matsuda, 1960. Keys to subfami-

lies, tribes, genera and subgenera of the Gerridae of the

World. — Kansas University Science Bulletin 41: 3-23.

Hungerford, H.B. & R. Matsuda, 1965. The genus

Ptilomera Amyot and Serville (Gerridae: Hemiptera). —

Kansas University Science Bulletin 45: 397-515.

Lundblad, O.M., 1934. Schwedisch-Chinesische Wissen-

schaftliche Expedition nach den Nordwestlichen

Provinzen Chinas. — Arkiv för Zoologie 27A (14): 1-31,

pls. 1-2.

Matsuda, R., 1960. Morphology, evolution and a classifica-

tion of the Gerridae (Hemiptera-Heteroptera). — Kansas

University Science Bulletin 41: 25-632.

Miyamoto, S., 1967. Gerridae of Thailand and North

Borneo taken by the Joint Thai-Japanese Biological

Expedition 1961-62. — Nature and Life in Southeast Asia

5: 217-257.

Miyamoto, S. & C. Lee, 1963. Water striders of Korea

(Hemiptera, Heteroptera). — Kontyü 31: 33-47.

Thirumalai, G., 1986. On Gerridae and Notonectidae (Hete-

roptera: Hemiptera: Insecta) from the Silent Valley, Kerala.

— Records of the Zoological Survey of India 84: 9-33.

MOLLER ANDERSEN & CHEN: Revision of Rhyacobates

Zettel, H., 1994. Revision der Gattung Potamometropsis

Lundblad (Insecta: HeteropteraL Gerridae). — Annalen

des Naturhistorischen Museums Wien (B) 86: 75-98.

Received: 20 June 1994

Revised and accepted: 27 September 1994

ende vide rm ee

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À Dre ra of ie ade

entr “ado Giu he gi

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tuer Hubs 1 « dr

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Nico NIESER

Tiel, The Netherlands

NINE NEW SPECIES OF PSEUDOVELIA AND A NEW

XIPHOVELIA (HETEROPTERA: VELIIDAE) FROM

SULAWESI (INDONESIA) AND MINDANAO (PHILIPPINES)

Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), V.

Nieser, N., 1995. Nine new species of Pseudovelia and a new Xiphovelia (Heteroptera: Veliidae)

from Sulawesi (Indonesia) and Mindanao (Philippines). Notes on Malesian aquatic and semi-

aquatic bugs (Heteroptera) V. — Tijdschrift voor Entomologie 138: 69-87, figs. 1-58, table 1

[ISSN 0040-7496]. Published 15 June 1995.

Nine new species of Pseudovelia from Sulawesi (including the northern islands of Sangihe and

Karakelong) and Mindanao are described: P. aflia sp. n., P. afrofila sp. n., P. epimekta sp. n.,

P. koutali sp. n., P. mystax sp. n. and P. pyrokrene from Sulawesi; P. sangihe sp. n. from Sangihe;

P. argyropardala sp. n. from Karakelong and Mindanao and P. kalami from Mindanao.

Additional data on two species and a revised key to Pseudovelia of the Oriental Region are also

given. Finally a new species of Xiphovelia: X. skoteina sp. n. from Sulawesi is described.

Dr. N. Nieser, Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands.

Key words. — Sulawesi, Karakelong, Sangihe, Mindanao; Veliidae; Pseudovelia, key, new spe-

cies; Xiphovelia, new species.

The present paper belongs to a series on the

Nepomorpha and Gerromorpha of Sulawesi started

by P. P. Chen and N. Nieser (papers listed in Nieser

& Chen 1993). The series is a contribution to the

Fauna Malesiana programme and is based on speci-

mens mainly collected by the staff members of the

Zoological Museum Amsterdam and the National

Museum of Natural History Leiden, Dr. G.

Zimmermann (Marburg, FRG) and the author.

Veliidae constitutes the second largest family of

Gerromorpha but as its species are on average dis-

tinctly smaller and often more cryptic than Gerridae

they are less well known. There are six subfamilies of

which five are represented in the Oriental region.

Most often encountered are the Rhagoveliinae with

its principal genus Rhagovelia. Its representatives live

on streams, often on exposed sites. The Sulawesi fau-

na has been revised recently by Nieser & Chen

(1993). Veliinae, which are with Velia the predomi-

nant subfamily in Europe are represented with 1 spe-

cies only in the Oriental Region (Andersen 1982).

The endemic subfamily Perittopinae consists of a few

cryptic species rare in collections. Haloveliinae with

an Oriental-Papuan distribution live mostly on shel-

tered sites in mangrove or on coral banks.

Pseudovelia belongs to the large cosmopolitan

subfamily Microveliinae. Representatives of this

subfamily are characterized by having one segment-

ed fore tarsi and two segmented middle and hind

tarsi. They are small to very small insects many of

which have more or less cryptic habits. Due to this

they have been relatively little studied. On the other

hand it is a species rich taxon with considerable

morphological differentiation. Expectations are that

detailed Studies in the future will reveal that there

are several more genera to be distinguished.

Especially the large genus Microvelia as conceived at

present appears to be polyphyletic; from time to

time a distinct group in Microvelia is split off as a ge-

nus. This is the way in which Pseudovelia came to

existence. The key to genera below may serve sort

out oriental Microveliinae.

METHODS

Measurements are in mm, and are presented as the

range or mean of five specimens, taken from the

sample containing the holotype (if available). Length

and width refer to the maximum value of the speci-

fied body part, if unspecified they refer to body length

and width. Length is measured in dorsal view from

anterior margin of head to apex of abdomen (inclusi-

ve genital segments) in apterous and to apex of he-

mielytra in macropterous specimens. If the tibial

comb projects, length of tibia is measured to apex of

comb, leg measurements are presented in Table 1.

Width of head is measured across eyes. Antennal me-

asurements have been lumped for sexes due to consi-

derable overlap, on average the means in males tend

to be 0.01 mm below and those in females 0.01 mm

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

above the lumped mean for each segment. Angles are

given in radians (TV).

The apical tarsal segments have, apart from the

claws, a dorsal and a ventral arolium placed in the

ventral incision. Of these the dorsal one is always bris-

tle shaped and, except for fig. 10, not drawn in the

figures. The ventral arolium is usually narrow but

may be broad, blade-like, as in figs. 45 and 52.

Pilosity is omitted in the drawings except when it is

striking or diagnostic.

Depositories registered according to Arnett,

Samuelson & Nishida (1993): BMKB (Kota Batu,

Brunei), JTPC (Englewood, Co., U.S.A.), MBBJ (Bo-

gor, Indonesia), MUDH (The Hague, The Nether-

lands), NHMW (Wien, Österreich), NMSC (Singapore),

RMNH (Leiden, The Netherlands), SEMC (Lawrence,

Kan. U.S.A), uscP (Cebu City, Philippines), ZMAN

(Amsterdam, The Netherlands), ZMUC (Kobenhavn,

Danmark). Not registered: BPUH (Bagian Pertanian

Universitas Haluoleo, Kendari, Sulawesi), NC (Nieser

Collection, Tiel, The Netherlands), PCHC (P. P.

Chen Collection, Beijing, P. R. China), ZC (Zimmer-

mann Collection (Goßfelden (Marburg) B.R.D.).

Some additional data on the localities in Sulawesi

Tenggara sampled by N. Nieser can be found in

Nieser & Chen (1991). Specimens collected by

Nieser are in NC unless otherwise specified.

SYSTEMATIC PART

Key to Oriental genera of Microveliinae

Adapted from Andersen 1982.

1. Claws all subterminal, inserted very close to the

apexioßtarsusillrian)gee en Velohebria

— Claws all preterminal, inserted distinctly before

APEK Of TAFSUS ET D

2. Larger species, length at least 3.5 mm. First an-

tennal segment slender and very long, at least 3/4

tnewidthiofshead\berweenieyest nn 3

— Smaller species, length less than 3.5 mm. First

antennal segment shorter and stouter .............. 5)

3. Femora modified (with hair tufts, tubercles etc.)

on posterior surface (at least in males).............. 4

— Femora not modified on posterior surface

(Oriental. Be rm eee Neoalardus

4. Eyes distinctly removed from lateroanterior an-

gles of pronotum (W Malaysia) ...... Lathriovelia

— Eyes touching or very close to anterolateral angles

Ofpronotuiml Oriental) eee Baptista

5. First antennal segment incrassate, extending

more than 2/3 of its length beyond apex of head.

Apical cells of forewings reduced (or macropter-

QUSHO LIU IO VI) E: es eee eee 7

— First antennal segment more slender and usually

much shorter; if not, then with normal forewing

venation or shortened pronotal lobe (apterous

FOR e ne 6

6. Middle tarsi with three leaflike structures subapi-

cally (claws and ventral arolium, fig. 52) (Orien-

tal) ER Eee DI Nea Xiphovelia

— Middle tarsi not modified as above (Cosmopoli-

FAT) cee ere cece eer AE Microvelia

7. Parameres very small, segment 8 of male strongly

modified ventrally (Palaeotropical and Far East-

EEN) ee Pseudovelia

— Parameres large, segment 8 in male not strongly

modified (Hlımalayan) en Geovelia

Genus Pseudovelia Hoberlandt, 1950

The genus Pseudovelia was erected somewhat la-

conically by Hoberlandt (1950 p. 38: ‘Stout antennae

and chiefly long first antennal joint, extending by

more than 2/3 of its length beyond the apex of head

are the characters on the basis of which it is possible

to establish for these species a separate subgenus,

Pseudovelia subg. n.°). Recently the genus has been

thoroughly redescribed in a revision of the Oriental

species by Andersen (1983). For practical purposes

characteristics mentioned in the key to genera above

will serve to recognize the genus. Apart from body

size and relative length of (parts of) appendages good

diagnostic characteristics on the species level are

mainly found in secondary sexual characteristics of

males. The parameres which are very small and look

more or less the same in different species (fig. 2) in

this genus are of little use. However, the ventral struc-

ture of first genital segment and the length of the well

developed tibial combs on fore leg, which curve

around the apical spur-like process on tibia provide

excellent characteristics.

The following key is based on that in Andersen

(1983) with insertion of the new species. P. tjurupen-

sis (Lundblad), which is only known in female sex, is

not included.

Key to males of Pseudovelia in the Oriental

Region

1. Eyes densely hairy. Ventral arolium leaflike flatte-

ned !(fig 525) AEN EEE NIE 2

— Eyes naked except for two ocular setae. Ventral

aroliumibristleslile Aen 3

2. Length (apterous form) d 2-2.2, ? 2.5-2.8 mm.

male hind tarsus distinctly shorter than tibia; tar-

sal segment 1 with a row of long bristles along its

entire length [Indian Subcontinent to Thailand

and! Sumatera eee ae RENEE

nee P. sexualis (Paiva) [= P. crassipes (Lundblad)]

— Length (apterous form) d 2.8, 2 3.0-3.2 mm.

Male hind tarsus as long as tibia; tarsal segment 1

10.

with a basal tuft of long bristles [S. China (Fu

Tar). ea 0 P. longitarsa Andersen

Ventral lobes of head strongly produced back-

wards (fig. 46). Hind tarsus of male distinctly

longer than tibia; tarsal segment 1 modified,

twice as long as segment 2 [Thailand]

moti EINE P. buccula Andersen

Ventral lobes of head not produced backwards.

Elinditarsustnotasfabovenen en, 4

Fore tibia swollen proximally and distally, result-

ing in a strongly sinuate posterior margin (Fig.

56) [N. Mindanao] … P. kalami sp. n.

Fore tibia less broad, its posterior margin evenly

andigentivicumedhia. satana nn 5

First genital segment of male with 2-5 spine-like

Processestontuberelessvenerallyernr. ern. en. 6

Remark: under strong magnification the spine-like

processes turn out to consist of a basal tubercle with

a small acute tuft of closely packed bristles on top.

First genital segment of male not as above ..... 11

Head apart from the silvery hairs at inner margins

of eyes also with a patch of silvery hairs medio-

caudally

Head mediocaudally at most with very few isola-

ted silvery hairs not forming a patch

Segment 1 of male hind tarsus distinctly shorter

than segment 2, ventral side of first genital seg-

ment of male anteriorly with a suboval cup sha-

ped structure (fig. 50) [W. Malaysia]

Rete ek Tan P. lundbladi Andersen

Segment 1 of male hind tarsus subequal to or

slightly longer than segment 2, segment 8 vent-

rally without cup shaped structure

First genital segment of male with one large me-

dian triangular process at base of ventral depres-

sion, in addition one slender median anda pair of

lateral process near its posterior margin [Suma-

Cerda eo P. hypodonta Lundblad

First genital segment of male with two small tu-

bercles at base and three near caudal margin of

ventral depression (fig. 9) [N. Borneo] ...............

Br ben alle lan Palani eek P. borneensis Andersen

Larger and relatively more slender species , d L

2.4, W 0.83 mm. Apical comb on fore tibia of

male about half as long as tibia (fig. 22), ventral

impression of first genital segment bordered by fi-

ve elongate and sharp processes (consisting of

densely packed bristles) (figs. 20, 21) [N. Sula-

Wes nn P. epimekta sp. n.

Shorter and relatively chubbier species, 4 L 2.0-

2.15, W 0.80-0.84 mm. Tibial comb of male

about one third the length of tibia (fig. 1), ventral

impression of first genital segment bordered by 4-

bishorterprocessesi(fio sto 90) VA 10

Ventral impression of first genital segment

bordered by five processes, two pairs laterally and

Jar

122

US:

14.

16.

170

NIESER: New species of Veliidae

a single mediocaudally (figs. 3, 8) [N. Sulawesi] .

ER P. aflia sp.n.

Ventral impression of first genital segment borde-

red by four processes, a mediocaudal one absent

(figs. 39, 40) [Sangihe 1.] .......... P. sangihe sp. n.

Trochanter of male fore leg distally with a short

stout process (fig 25), first genital segment vent-

rally with a large spoon-shaped process (fig. 26)

INAS ul amesi eee oe P. koutali sp. n.

Trochanter of male fore leg more or less rounded,

first genital segment ventrally at most with a cus-

hion-like plate, no distinct ventrally directed pro-

JECTION. RME i LR aas 12

Segment 1 of male hind tarsus with ventral row of

long (as long as segment, fig. 44) bristle-like hairs

Segment 1 of male hind tarsus without distinct

ventral row of long hairs which are as long as seg-

mente ts Etes MEERN ME oe aes 14

Tibial comb 3/4 the length of fore tibia (fig. 16),

length of antenna 2/3 body length (1.4/2.2)

[Karakelong I., SE Mindanao]

RAN ER P. argyropardala sp. n.

Comb on fore tibia of male 2/3 the length of tibia

(fig. 19), length of antenna slightly less than 2/3

body length (1.18/2.0) [Philippines]

zn ee bi a P. reiseni Polhemus

Smaller (length of d up to 2.2 mm) grey-blackish

species, with a distinctly contrasting transverse

orange patch anteriorly on pronotum ............ 15

Larger (length d 2.2mm or more), brownish spe-

cies, anterior orange pronotal mark not strongly

EOntrasting NE RE met 16

Comb on fore tibia of male about one third the

length of tibia, impression ventrally on first geni-

tal segment with a transverse rim in caudal part

(Giese) Sulawesi tr... P. afrofila sp. n.

Comb on fore tibia of male slightly more than

half the length of tibia, ventral impression on first

genital segment simple [Sri Lanka]

PR RE ER is. PA TIERE VE P. gnoma Polhemus

Second genital segment of male with a tuft of

bristles on each side; first genital segment ventral-

ly with a broad cushion reaching across the seg-

ment in apical half (figs. 30, 31). [Hind tarsus +

half as long as tibia] [N. Sulawesi]

ONE EEE Ei QAR. rt re re P. mystax sp. n.

Second genital segment of male without tufts of

bristles laterally; first genital segment ventrally

with a less strongly chitinized impression in api-

Canal (He TIC) een ee 1%

Length of antenna in male about 2/3 the length

of body, hind tarsus 2/3 the length of hind tibia

ISESSulamesi een P. pyrokrene sp. n.

Length of antenna in male less than half the body

length, hind tarsus about half the length of hind

Val

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

tibia. [Thailand, West Malaysia to Jawa] ............

RR nn P. feuerborni Lundblad

Pseudovelia aflia sp. n.

(figs. 1-6, 8)

Type material. — Holotype apterous à , INDONESIA,

Sulawesi Utara, near Danau Mooat, temporary

stream in hills (W of Desa Kankenturan), 21. Nov.

1985, leg. G. Zimmermann (RMNH). Paratypes 14d

72 apt, 18 29 macr., same data as holotype, dis-

tributed as follows: 1d (macr.) 19 (allotype) RMNH;

26 22 pere; 1d rees 1d) MBBJ; 46) 32 (12

macr.) NC; 4d 39 (19 macr.) zc; 16 ZMAN; 14

ZMUC.

Description. — Apterous specimens. Dimensions.

Length d 2.00-2.13, 2 2.40-2.52; width of head 3

0.50-0.52, 9 0.52-0.54; width of pronotum d 0.68-

0.72, 2 0.73-0.78; width (across metanotum/first

abdominal tergite) d 0.80-0.84, 2 0.92-0.98.

Colour and pilosity. Generally pale brownish with a

greyish tinge, suture between body sclerites blackish,

legs and antennae light brown. Patches of silvery hairs

along inner margins of eyes rather inconspicuous; lat-

erally on first abdominal tergite and mediocaudally

on tergites 2-6, variable and indistinct in some speci-

mens. Body clothed with rather short and sparse,

erect brownish pilosity.

Length of antennal segments I-IV, 0.40: 0.25:

0.24: 0.42. Anterior quarter of pronotum and swol-

len anterolateral angles smooth, posterior three quar-

ters coarsely punctate. Lateral margins distinctly in-

dented about halfway on anteroposterior line,

greatest width just anterior of indentation. Ventral

arolia bristle like. Middle tibia with about six long

erect hairs on inner side of apical half. Hind tibia with

slightly denser and thicker pilosity apically.

Male. Elongate, connexiva roughly horizontal, lat-

eral margins parallel to slightly diverging in anterior,

converging in posterior half. Length of antenna two

thirds of body length (1.3/2.1). Pronotum one and

three quarters as wide as its median length

(0.74/0.42), distinctly narrower than width across

metanotum. Anterior trochanter rounded. Apical

comb on fore tibia relatively short, + one third

(0.2/0.6) tibial length (fig. 1), consisting of about 90

teeth of equal width except for a small group of nar-

rower teeth in the middle. First segment of hind tar-

sus apart from the normal pilosity with about 7 long

(longer than diameter of segment) cilia ventrally.

Abdominal tergite 1 large, coarsely punctate, tergites

1 and 2 nearly flat, in lateral view level with or raised

slightly above connexiva. Pregenital tergite + one fifth

the dorsal median length of abdomen. First genital

segment with a large ventral impression with five

spine-like processes on its margin (figs. 3, 4, 8).

Female. Oval shaped, bulkier than male. Con-

nexiva slanting + 0.257 upward, their lateral margin

evenly curved with its greatest width on tergite 4.

Length of antenna slightly over half the length of

body (1.30/2.45). Pronotum twice as wide as long as

its median length (0.78/0.41), distinctly narrower

than width across metanotum. Abdominal tergites 1-

2, only slightly convex dorsally, in lateral view hardly

rising above the margin of connexiva. Caudal apices

of connexiva remaining considerably removed from

each other, leaving tergite 7 visible. Tergite 8 orient-

ed horizontally, fully visible in dorsal view. Pregenital

segment large, its sternite slightly more than one

fourth the median length of abdominal venter

(0.50/1.75). Genital segments relatively large and

protruding caudally, more or less U-shaped in caudal

view (figs. 5, 6).

Macropterous form. — Structurally essentially as

apterous form except for the usual modifications of

macropters in the subfamily. Length d 2.2, 9 2.3-

2.5; humeral width of pronotum d 1.00, 9 1.00-

1.10. Hemielytra light grey-brown with an, usually ill

defined, elongate sordid white stripe at base.

Venation rather indistinct, two elongate basal cells, of

which the one near hind (inner) margin seems two be

divided. In addition two shorter apical cells. Base of

abdomen with distinct lateral (near connexival su-

tures) keels on tergites 1-3 and a sorter, less distinct

sublateral pair on tergites 1 and 2.

Etymology. — Aflios (Greek adjective meaning

poor, meagre) refers to the comparatively poor pilos-

ity on body.

Comparative notes. — The male genitalia are some-

what similar to those of P. borneensis Andersen (figs.

8, 9) but P. aflia lacks the patch of silvery hairs me-

diocaudally on head, and the detailed structure of Ist

genital segment in d is different in that the rim of the

impression is thicker in P. aflia and the warts on

which the ventral groups of bristles are placed are less

strongly developed. Females of P. borneensis have ter-

gite 8 somewhat more produced caudally and abdom-

inal segment 7 somewhat less pilose laterally (figs. 6,

7). The male genitalia also remind those of P. buccu-

la Andersen, an aberrant species with ventral lobe of

head strongly produced caudally (fig. 46) and first

segment of hind tarsus very long. For comparison

with P. epimekta see under that species.

Pseudovelia afrofila sp. n.

(figs. 10-13)

Type material. — Holotype apterous 9, INDONESIA:

Sulawesi Utara, Sungai Girian (NE slope of G.

Klabat, E of Manado), mountain stream with rapids,

bottom boulders to coarse sand, specimens collected

NIESER: New species of Veliidae

Figs. 1-6. Pseudovelia aflia. Paratypes: 1-4 male; 1 fore leg, 2 paramere, 3 first genital segment (segment 8) in ventral view, 4

detail of ventral impression of segment 8; 5-6 apex of abdomen of female, 5 caudal, 6 lateral view. — 7 lateral view of apex of

abdomen of female of P. borneensis 8-9 lateral view of segment 8 of male, 8 P. aflia, 9 P. borneensis. — 10-13. Pseudovelia af-

rofila. 10-12 male allotype, 10 fore leg, 11 genital segments, lateral view, 12 segment 8, ventral view, 13 female paratype, ap-

ex of abdomen, lateral view. — 14-16 Pseudovelia argyropardala. Male paratypes: 14 segment 8 lateral view, 15 segment 8

ventral view, 16 fore tibia. — 17-19 P. reiseni male: 17 segment 8 lateral view, 18 segment 8 ventral view, 19 fore tibia.

Scale line for figs. 3 and 5: 0.1mm; figs. 10-13, 16 and 19: 0.5mm; figs. 1, 6, 8, 14, 15, 17 and 18: 0.25 mm; figs. 2 and 4:

0.05mm. Figs. 7 and 9 after Andersen (1983).

TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 138, 1995

on patch of foam behind boulder, 23. June 1994,

N9452, leg. N. Nieser (ZMAN). Paratype 16 (allo-

type) and 1 ®, same data as holotype, (NC). All apter-

ous.

Description. — Apterous specimens. Dimensions.

Length d 2.10, 9 2.35-2.40; width of head d 0.55,

2 0.58-0.60; width of pronotum d 0.72, 9 0.80-

0.85; width (across third abdominal tergite) & 0.90,

® 1.08-1.10. Colour and pilosity. Generally dull

dark grey, anterior third of pronotum pale orange,

one female with an indication of orange median line;

lateral halves of connexiva (dorsally and ventrally)

and distal rims of acetabula brownish; antennae and

legs pale yellowish to light brown, proximally lighter

than distally. Patches of silvery hairs along inner mar-

gins of eyes distinct, a narrow longitudinal stripe me-

dially on vertex shining black (due to lack of basic pi-

losity). Silvery hairs laterally on first abdominal

tergite and on posterior parts of remaining tergites in-

distinct. Body covered with velvety pilosity super-

posed with distinct erect brownish pilosity, giving the

specimens a hairy look more obvious dorsally than

ventrally.

Length of antennal segments I-IV, 0.41: 0.24: 0.29:

0.43. Anterior third of pronotum and anterolateral an-

gles smooth, posterior two thirds coarsely punctate.

Greatest width of pronotum about halfway anteropos-

terior line, lateral margins convex without pronounced

humeral angles or indentation. Ventral arolia bristle

like. Middle tibia with about 5 long (twice as long as

the width of tibia) hairs in apical half of inner side. First

segment of hind tarsus ventrally with a few longer cilia

twice as long as the width of segment.

Male. General shape elongate oval with a truncate

caudal apex, connexiva more or less horizontal, their

lateral margins convergent in posterior half. Length

of antenna 2/3 the length of body (1.4/2.1).

Pronotum less than twice as broad as its median

length (0.7/0.4). Anterior trochanter rounded. Fore

tibia with grasping comb one third the length of tibia

(0.2/0.6, fig. 10), consisting of about 80 coarse teeth.

Hind tibia with ciliation apically denser but not mod-

ified. First abdominal tergite large, flat dorsally, raised

about as high as the edge of connexiva. Pregenital seg-

ment about one fifth the length of abdomen. First

genital segment ventrally with a large impression with

strongly chitinized margin in posterior half, otherwise

little modified (fig. 11, 12). Second genital segment

lying within first, without special features.

Female. General shape broadly oval, connexiva

nearly horizontal, distinctly tapering in posterior 3/5

of abdomen. Length of antenna more than half the

length of body (1.4/2.4). Pronotum nearly twice as

wide as its median length (0.80/0.45), distinctly nar-

rower than width across metanotum. Abdominal ter-

gites 1-3, somewhat convex dorsally, in lateral view

first three rising above the margin of connexiva.

Caudal apices of connexiva remaining considerably

removed from each other, leaving tergite 7 visible,

tergite 8 slanting ventrally 0.16 7, tergites 8 and 9 ful-

ly visible in dorsal view. Segment 7 large, its tergite

two thirds as long as wide (0.22/0.33),its sternite

slightly more than one third the median length of ab-

dominal venter (0.42/1.10). Genital segments well

developed, basal part of gonocoxa retracted into ab-

domen (fig. 13).

Macropterous form unknown.

Etymology. — Afrofilos (aphrophilos) is a greek

composite adjective meaning ‘foam loving’.

Comparative notes. — The general shape (especially

in females) and colour of this species is not similar to

any species known from Sulawesi. It is apparently

similar to P. gnoma Polhemus from Sri Lanka, which

is unknown to me, the characteristics in the key will

suffice to distinguish the species.

Remarks. — The species was found together with P.

koutali in a patch of foam behind some boulders at an

edge of the stream. Andersen (1983) reports the ap-

parently related P. gnoma also from ‘in foam in river’

other species have, however, also been found on

patches of foam.

Pseudovelia argyropardala sp. n.

(figs. 14-16, 44, 48-49)

Type material. — Holotype apterous d , INDONESIA:

Sulawesi Utara, Pulau Karakelong, Sungai Ambela at

inlet of irrigation canal for desa Ambela, large open

river bed in disturbed rain forest, transition from

mountain to lowland stream, only a little water (sluic-

es in dam open), actual stream about 2m wide, 0.3-

0.5m wide, water turbid, 30 Nov. 1994, N9493, leg.

N. Nieser (RMNH). — Paratypes (adults only), same

data as holotype 68 119 apterous, 38 52 macrop-

terous, 8 lvIV-V, distributed as follows: 1 9 apt. (allo-

type), 18 12 macr. RMNH, 16 19 NHMW, 1d 12

NMSC, 18 19 PCHC, 1d apt., 12 macr. SEMG; P.

Karakelong, Sungai (at desa) Pampalu, about 200 m

upstream of mouth, open lowland stream through ag-

ricultural area, mostly some shade, bottom sand and

pebbles, 1. XII. 1994, N9495, leg. N. Nieser, 2d

19; P. Karakelong, Sungai Ambela at foot of water-

fall, mountain stream in primary rainforest,

Pseudovelia under overhanging rocks at edges of

pond, 2. XII. 1994, N9497, leg. N. Nieser, 48 129

apt, 1d 29 macr., 20 lv IV/V (re me 192 MBB),

VP Mur, Io Save, WL ze, WS WL AUS 1.

Karakelong, Sungai Sawang, lowland stream through

agricultural area, very turbid after rain, Pseudovelia on

patch of foam at edge, 3. XII. 1994, N9498, leg. N.

Nieser, 18 12 apt., 6 IvIV/V. Pulau Salibabu,

Lirung, narrow stream on very steep rocks draining,

small shallow pools with much plant debris connect-

ed by trickles of water, shaded by remnants of forest,

+ 350m asl., 24. XII. 1994, N9483, leg. N. Nieser

54 119 apt, 19 macr. — PHILIPPINES, Mindanao:

Sarangani (S. Cotabato) prov. Siquel River, pool (ar-

ea + 20 m’, 0.2 m deep) at edge of river bed, slightly

turbid stagnant water, under overhanging grassy

bank, 23. XI. 1993, N9351, leg. N. Nieser, 29 apt.,

2d 22 macr. (ld 12 macr. USCP); Lake Sebu area,

stream at Bakdolong, lowland stream aspect, sand/

mud bottom, Pseudovelia at edge, 3.X11.1993,

N9364, leg. N. Nieser, 19 apt; Stream at Bakdo-

long, mountain stream aspect, bottom with well sized

pebbles, Pseudovelia at edge, 3 & 4. XII. 1993,

N9366, leg N. Nieser, 5d 10% apt, 1d 29

macr.(16 2% apt to resp: NHMW, RMNH, USCP,

ZMAN, 1d 19 apt. & 1d 1% macr. JTPC); stream at

Bakdolong, upstream across the agricultural area, be-

ginning of hilly area, narrow stream in limestone,

shaded by shrubs and trees on the banks, Pseudovelia

from pools mostly under overhanging ridges or tree

roots, 10. XII. 1993, N9379, leg. N. Nieser, 298

379 apt., 2d 49 macr.; Lake Sebu area, just up-

stream of third waterfall, edge upstream of a large

boulder in mountain stream, slopes with remnants of

primary forest, 7. XII. 1993, N9371, leg. N. Nieser,

VNS Rape acid dal? ape EME MONS

apt. 1? macr. SEMC); Lake Sebu area, just down

stream of second waterfall at edge of spray area, shel-

tered edge downstream of boulders, 8. XII. 1993,

N9373, leg. N. Nieser, 18 19 apt.

Description. — Apterous specimens. General shape

elongate oval. Dimensions. Length d 2.18-2.40, 9

2.53-2.75; width of head & 0.56-0.59, £ 0.56-0.60;

width of pronotum d 0.77-0.80, 9 0.78-0.81; width

(d across posterior margin of thorax, 9 across ab-

dominal tergites 4-5) 4 0.88-0.92, 9 0.94-1.03.

Colour and pilosity. Generally dark grey to blackish

with striking silvery spots dorsally on abdomen.

Transverse band anteriorly, lateral margin and nar-

row median longitudinal stripe on pronotum and lat-

eral half of connexiva brick red. Legs pale yellowish,

bases and apices of segments usually darker, antennae

light brown, base of rostrum pale, apical segment

dark. Bands of silvery hairs along inner margins of

eyes distinct; patches of silvery hairs posterior margin

of metanotum, laterally on first abdominal tergite,

medially on tergites 2 and 3, mediocaudally on con-

nexiva 3-6, posterior part of tergite 6 and most of ter-

gite 7. Compared to other species in the region the

silvery patches are more strongly contrasting in this

species. Body clothed with long light brown pilosity

which is more erect and distinct dorsally than ventral-

NIESER: New species of Veliidae

ly, especially in lateral view the species appears hirsute

dorsally. Antennae and legs also rather hirsute.

Length of antennal segments I-IV, 0.38 : 0.30 :

0.32 : 0.44. Anterior quarter of pronotum slightly

swollen, smooth except for a row of pits along anteri-

or margin, coarse punctation on posterior three quar-

ters obscured by dense pilosity. Lateral margins of

pronotum smoothly curved, not or hardly indented,

greatest width about halfway. Ventral arolia bristle

like. Middle tibia with about six long erect hairs on

inner side of apical half, becoming distinctly shorter

apically. Denser and thicker pilosity apically not obvi-

ous.

Male. Connexiva roughly horizontal to slanting

0.257 upward, lateral margins nearly parallel in ante-

rior, converging in posterior half. Length of antenna

about two thirds of body length (1.45/2.2).

Pronotum less than twice as wide as its median length

(0.8/0.45 - 0.8/0.5), narrower than width across

metanotum (0.8/0.9). Anterior trochanter rounded.

Apical comb on fore tibia long, slightly over three

quarters (0.5/0.65) tibial length (fig. 16), consisting

of over 300 teeth of which the proximal and distal

groups are much finer than those from the central

interval. First segment of hind tarsus apart from the

normal pilosity with a group of about 20 long (about

as long as length of segment, fig. 44) cilia ventrally.

Abdominal tergite 1 large, smooth, sloping down-

ward posteriorly; tergites 2-7 flat, horizontal, in later-

al view concealed by the connexiva. Pregenital tergite

+ one fifth the dorsal median length of abdomen.

First genital segment with a pair of ventral impres-

sions, beset with cilia, dorsocaudal pilosity long (figs.

14915):

Female. Larger and bulkier than male but of same

general form. Connexiva slanting 0.3% upward to

nearly vertical (nearly horizontal in gravid speci-

mens), their lateral margins nearly parallel in anterior,

converging in posterior half. Length of antenna

slightly more than half the length of body (1.45/2.6).

Pronotum twice as wide as its median length

(0.9/0.45), distinctly narrower than width across

metanotum. Abdominal tergite 1 large, coarsely

punctate, + horizontal, in lateral view visible dorsally

of connexiva, tergites 2-3 gradually sloping down-

ward posteriorly, less distinctly punctate than tergite

1; tergites 4-8 flat, horizontal, in lateral view con-

cealed by the connexiva (except in gravid females).

Caudal apices of connexiva rounded, remaining con-

siderably removed from each other, leaving tergite 7

visible. Tergite 8 oriented horizontally, fully visible in

dorsal view. Pregenital segment large, its sternite

about one fourth the median length of abdominal

venter (0.35/1.4). Genital segments largely retracted

within abdomen, more or less U-shaped in caudal

view (figs. 48, 49).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Macropterous form. — Structurally essentially as

apterous form except for the usual modifications of

macropters in the subfamily. Length d 2.55-2.60, 9

2.60-2.70; humeral width of pronotum d 1.06-1.08,

? 1.12-1.15. Pronotum dark grey-brown with a

lighter T-shaped mark, anterior transverse leg broad,

posterior longitudinal leg narrow, disc of pronotum

distinctly punctate. Hemielytra grey-brown with ba-

sal angles largely white, apical cells brownish with

lighter centres and membrane greyish. Venation dis-

tinct, two elongate basal cells and two shorter apical

cells. Lateral keels at base of abdomen long, reaching

base of tergite 4, medial pair less distinct, reaching

caudal margin of tergite 2; silvery pilosity on abdo-

men less developed especially anteriorly.

Etymology. — Argyropardalos, a greek adjective mean-

ing ‘with silvery spots’, referring to the distinct patches of

silvery pilosity on abdomen of most specimens.

Comparative notes. — Size in combination with pi-

losity, greyish colour and distinct patches of silvery

hairs on abdomen sets this species apart from other

Sulawesi species. This species is very similar to P. re-

iseni Polhemus from Luzon and Mindanao males of

which are, however, smaller and have a relatively

shorter fore tibial comb of somewhat over 200 teeth

which are all coarse distally (in argyropardala, which

has about 300 teeth in tibial comb, there is a row of

very fine teeth near the distal spur) and the ventral

impression on first genital segment of male less dis-

tinctly paired and much smaller (see key and figs. 14-

19). Females show considerable overlap and only the

smaller females of P. reiseni (L about 2.4) are reliably

distinguishable from the larger females of P. argyro-

pardala (L about 2.7).

Pseudovelia epimekta sp. n.

(figs. 20-24)

Type material. — Holotype apterous 4 , INDONESIA,

Sulawesi Utara, Dumoga Bone N. P., Tumpah, ru-

hige Abschnitt am Moos (quiet stretch at marshland),

28. Oct. 1985, leg. G. Zimmermann (RMNH).

Paratypes 29, same data as d (19 NC, 19 allotype

ZC). All apterous.

Description. — Apterous specimens. Dimensions:

length d 2.40, ® 2.50-2.65; width of head d 0.53,

? 0.58-0.59; width of pronotum d 0.68, ® 0.72-

0.73; greatest width (across mesothorax) d 0.83, ©

0.86-0.89.

Colour. Generally dark grey to blackish, eyes very

dark brown. Head with grey stripes at inner margins

of eyes. Pronotum with strongly contrasting anterior

orange transverse band not interrupted in the middle,

disk of pronotum medium brown with a very narrow

median orange-like stripe. Connexiva somewhat

brownish, more distinctly so on ventral side. Anten-

nal segments 1-3 medium brown, segment 4 pale,

whitish. Segments 2-3 of rostrum largely yellowish

brown (in frontal view). Acetabula on both sides of

suture brownish, legs sordid white to light brownish,

apical part of femorae and basal part of tibiae medium

brown.

Silvery pilosity at lateral margins of abdominal ter-

gite 1, most of tergites 2-3,6, hind margin of tergite 5

and median part of tergite 7, quite variable especially

on 6 and 7. Sparse, scattered longer erect bristles dor-

sally. Venter with well developed medium long some-

what yellowish pilosity, slightly denser on genital seg-

ments. Tuft of bristles at end of metasternal scent

channels strongly developed. Pilosity of appendages

normal, first antennal segment with two distinct erect

bristles on anterior side.

Body elongate spindle-shaped, greatest width

across mesothorax, subequal to width across posterior

margin of abdominal segment 4 (depending on slant-

ing of connexiva this is narrower in 16 19 and very

little wider in 1 9). Head over three fourths as long as

wide (0.45/0.55), longer than median length of pro-

notum (0.45/0.40). Eyes small, width of an eye only

slightly more than one third the interocular space

(0.12/0.32). Length of antennal segments I-IV =

0.62: 0.31: 0.42: 0.64. First antennal segment dis-

tinctly longer than head, relatively slender, elongate

club-shaped and softly curved, with two distinct erect

bristles. Second slightly narrower, third and fourth

narrowest, subequal. Rostrum nearly reaching base of

metasternum. Pronotum more than half as long as

wide (0.4/0.7), covering meso and metanotum except

for lateroposterior angles of metanotum. Disk shal-

lowly and indistinctly pitted, with an indistinct trans-

verse impression halfway, somewhat more distinct in

the male than in the females. Anterior transverse

orange band of pronotum smooth, slightly swollen.

Propleura with some coarse punctures, most distinct-

ly a row at dorsal margin of acetabulum and another

near posterior margin. Fore femur somewhat flat-

tened on under surface in apical part. Arolia bristle-

like. Connexiva with lateral margins nearly straight,

subparallel on segment 1-4, converging in caudal

part, slanting upward about 0.167. Connexiva widest

on segment 4, about half as wide as corresponding

tergite. Median length of abdominal tergites 1-7 three

times the median length of pronotum (1.40/0.45) in

male, slightly more in female (1.60/0.45). Abdominal

tergite 1 large, raised above dorsal margin of connex-

iva in lateral view, somewhat more so in females than

in males, tergites 2-3 declivous, tergites 2-5 each

about half as long as tergite 1, tergites 6 and 7 pro-

gressively longer, especially in male. Abdominal vent-

er flattened in male, indistinctly so in female, with a

NIESER: New species of Veliidae

Figs. 20-24. Pseudovelia epimekta. 20-23 male holotype: 20 genital segments ventral view, 21 segment 8 lateral view, 22 fore

tibia, 23-24 apex of abdomen, 23 male, ventral view, 24 female, lateral view. — 25-28, Pseudovelia koutali, paratypes: 25-27

male 25 fore leg, 26-27 genital segments, 26 lateral view, 27 ventral view, 28 female apex of abdomen, lateral view. — 29-33,

Pseudovelia mystax. Paratypes: 29-31 male, 29 fore leg, 30-31 genital segments, 30 lateral view, 31 ventral view, 32-33 fema-

le apex of abdomen, 32 lateral view, 33 caudal view. Scales 0.5 mm. — 34-38 Pseudovelia pyrokrene. Paratypes, 34-36 male, 34

fore leg, 35 genital segments, lateral view, 36 segment 8 ventral view, 37-38 female apex of abdomen, 37 lateral view, 38 cau-

dal view. Scale line for figs. 20-25, 28-32, 34-35: 0.5mm; figs. 26-27, 36-38: 0.25mm.

Hil

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

narrow, shallow, mediolongitudinal groove on ster-

nites 5-7, sternites 6 and 7 progressively longer than

preceding ones.

Male. Length of antenna five sixth the length of

body (2.0/ 2.4). Comb on fore tibia long (fig. 22),

about half as long as tibia consisting of about 200

teeth which are distinctly stouter in apical part.

Otherwise legs without obvious characters, first seg-

ment of hind tarsi with a few longer hairs. Tergite 7

half as long as preceding abdominal tergites together.

Genital segments relatively large but segment 8 near-

ly fully retracted within abdomen. Sternite 8 with five

curved, sharply pointed processes, one, narrower, me-

diocaudal, one pair subcaudal and one pair halfway,

surrounding a ventral impression (figs. 20-21). Para-

meres very small, nearly vestigial.

Female. Length of antenna slightly over three quar-

ters the length of body (2.0/2.6). Abdominal tergite 8

horizontal, comparatively large, covering gonocoxae

in dorsal view, over half as long as tergite 7

(0.18/0.3), covering gonocoxae in dorsal view. Gono-

coxae large, well visible in ventral view. (figs. 23-24).

Macropterous form unknown.

Etymology. — Epimektes, a greek adjective meaning

elongate, referring to shape of body, especially of fe-

male.

Comparative notes. — This species belongs to the

hypodonta group of species (Andersen 1983). Seg-

ment 8 of male is most similar to that of P. borneen-

sis, especially in lateral view, the mediocaudal process

in P. borneensis ıs, however, stouter, more triangular.

P. borneensis is, moreover, smaller, of lighter colour

and has the first antennal segment relatively shorter

and stouter. The first genital segment of the male has

a similar form to that of P. aflia described above, but

it is longer (when isolated from body 0.55 in P. epi-

mekta, 0.44 in P. aflia) and has more solid ventral

warts with bristles (figs. 3, 8, 20, 21). P. epimekta is,

moreover, generally longer and relatively more slen-

der than ?. aflia with a distinctly longer first antennal

segment (0.6 and 0.4 respectively). P. aflia and P.

epimekta have a few silvery hairs at base of head but

these are not recognizable as a silvery patch.

Pseudovelia kalami sp. n.

(figs. 56-58)

Type material. — Holotype apterous d (only speci-

men known, right middle leg lacking, genitalia in mi-

crovial), PHILIPPINES; Mindanao, Surigao del Norte

prov., Oslao, narrow mountain stream through coco-

nut groove, bottom stones, pebbles and coarse sand,

water colourless, clear, sampled at quiet edges, 29

Mar. 1993, leg. N. Nieser, N9332 (RMNH).

Apterous specimen. — General shape elongate,

nearly parallel sided except anterior and posterior re-

gions. Dimensions. Length d 2.49, width (across

posterior half of mesothorax) 1.02, width of prono-

tum 0.96, width of head d 0.63.

Colour and pilosity. Generally blackish with strik-

ing silvery spots dorsally on abdomen. Transverse

band on anterior half of pronotum, and lateral rim of

connexiva dark orange brown. Legs, coxae trochant-

ers and basal part of femurs yellowish, remaining

parts brownish, antennae and rostrum also brownish.

Bands of silvery hairs along inner margins of eyes dis-

tinct, in addition a small spot of silvery hairs medio-

caudally on head; patches of silvery hairs: latero-

posterior angles of metanotum, laterally on first

abdominal tergite, mediocaudally on tergites 6 and 7,

mediocaudally on connexiva 3-5, posterior part of

tergite 6 and most of tergite 7. Compared to most

species in the region the silvery patches are more

strongly contrasting in this species. Body clothed

with long light brown pilosity which is more erect

and distinct dorsally than ventrally.

Length of antennal segments I : II : HI: IV, 0.55:

0.46 : 0.45 : 0.40. Length of antenna three quarters of

body length (1.86/2.49). Hind margin of ventral lobe

of head produced slightly backwards. Pronotum less

than twice as wide as its median length (0.96/0.6),

only slightly narrower than width across metanotum.

Anterior half of pronotum slightly swollen, smooth

except for some pits caudally, coarse punctation on

posterior half obscured by dense pilosity. Lateral mar-

gins of pronotum smoothly curved, not indentate,

greatest width about halfway.

Anterior trochanter rounded. Fore tibia proximally

and distally thickened, resulting in a sinuate posterior

margin, tibial comb runnig nearly the entire length of

tibia (fig. 56). Middle tibia without long erect hairs

on inner side. Denser and thicker pilosity apically not

obvious. First segment of hind tarsus apart from the

normal pilosity with a group of about 15 long (about

3 times as long as diameter of segment) cilia ventrally.

Ventral arolia bristle like.

Connexiva roughly horizontal, lateral margins par-

allel anteriorly, converging on segments 5-7.

Abdominal tergite 1 large, smooth, only slightly slop-

ing downward posteriorly; tergites 2-7 flat, horizon-

tal, in lateral view concealed by the connexiva. tergites

1 and 7 twice (0.3) as long as tergites 2-6 (0.15).

Abdominal venter with a large and broad median im-

pression. First genital segment with a large central im-

pression bordered by a very high rim caudally ending

in a pair of tufts of small bristles, surface of the im-

pression anteriorly and medially also with small bris-

tles (figs. 57-58).

Comparative notes. — General shape and colour

similar to P. argyropardala, which is slightly smaller.

The small but distinct spot of silvery hairs mediacau-

dally on the head reminds of P. pyrokrene and P. bor-

neensis. The form of the anterior tibia and the strong-

ly developed rim around the ventral depression of

first genital segment in male are, however, unique

within the genus. Females will probably be recogniz-

able by the relatively long antennal segments especial-

ly segment 2.

Etymology. — Kalami is a Greek noun, meaning

tibia, in apposition, referring to the characteristic fore

tibia of male.

Pseudovelia koutali sp. n.

(figs. 25-28)

Type material. — Holotype apterous d , INDONESIA,

Sulawesi Utara, Dumoga Bone River, downstream of

bridge, 22. Oct. 1985 leg. G. Zimmermann (RMNH).

Paratypes 11d 89, same data as type, distributed as

follows: 1 2, allotype, RMNH; 1d MBBJ; 26 29 PCHG;

46 29 NC; 36 39 zc; 16 ZMAN; Sulawesi Utara,

Sungai Girian (NE slope of G. Klabat, E of Manado),

mountain stream with rapids, bottom boulders to

coarse sand, specimens collected on patch of foam be-

hind boulder, 23. VI. 1994, N9452, leg. N. Nieser,

94 79 (16 19 prec, 16 12 MUDH, 16 12 NHMW,

1d 12 SEMC). All apterous.

Description. — Apterous specimens. Dimensions.

Length d 2.32-2.40, 9 2.80-2.92, width of head d

0.54-0.56, 2 0.60-0.62; width of pronotum d 0.78-

0.80, 2 0.80-0.83; width (across metanotum/first

abdominal tergite) & 0.83-0.88, 2 0.97-1.03.

Colour and pilosity. Generally dark brown, antennae,

T-shaped mark on pronotum, lateral margin of con-

nexiva medium brown, legs medium to light brown,

coxae, trochanters and basal parts of femurs yellow-

ish. Patches of silvery hairs along inner margins of

eyes, caudolaterally on first abdominal tergite, the

disks of tergites 2 and 3 and small patches medially on

tergites 4 and 5. Body mostly covered with quite

long, erect brownish pilosity.

Length of antennal segments I-IV 0.43 : 0.30 :

0.30 : 0.40. Anterior quarter of pronotum smooth,

posterior three quarters coarsely punctate.

Anterolateral angles of pronotum only slightly swol-

len. Ventral arolia bristle like. Middle tibia with

about 6 long (twice as long as the width of tibia) hairs

in apical half of inner side.

Male. General shape elongate, connexiva slanting

to a variable angle, usually about 0.57, their lateral

margins convergent in posterior half. Length of an-

tenna over half the length of body (1.4/2.4).

Pronotum broader than its median length (0.7/0.5)

with lateral angles about midway, very broadly

rounded. Anterior trochanter with a distinct ventral

NIESER: New species of Veliidae

tubercle (fig. 25). Fore tibia with grasping comb oc-

cupying about two thirds of the length of tibia

((0.35/0.58), consisting of about 150 teeth of equal

thickness except for a group of about 25 very fine

ones just proximally of the tibial spur. Hind tibia

straight, with ciliation along inner margin denser in

apical part, with a patch of slightly thicker cilia at ex-

treme apex. Hind tarsal segments of subequal length,

first tarsal segment ventrally with normal pilosity on-

ly. First abdominal tergite large, raised about as high

as the edge of connexiva. Pregenital segment large,

about one third the length of abdomen. Genital seg-

ments largely retracted within abdomen, first genital

segment ventrally with a large spoon-shaped append-

age (figs. 26-27).

Female. General shape bulkier than male, with

connexiva tapering more strongly. Length of antenna

half the length of body (1.4/2.8) or very slightly less.

Pronotum wider than long (0.8/0.6), greatest width

relatively more posteriorly than in male and more dis-

tinctly narrower than width across metanotum.

Anterior trochanter without tubercle and tibia with-

out comb. Pilosity of hind tibia less conspicuous than

in male, basal segment of hind tarsus distinctly short-

er than apical (0.2/0.3). First abdominal tergite large,

convexly swollen dorsally, in lateral view distinctly

rising above the margin of connexiva. Connexiva ver-

tical, strongly converging posteriorly with apical pi-

losity meeting across midline and covering tergite 7.

Pregenital segment large, about two fifth the length of

abdomen. Genital segments small, retracted into ab-

domen with a distinct tuft of darker bristles, situated

dorsally on the vertically oriented tergite 8, pointing

caudally (fig. 28).

Etymology. — Koutali, a Greek noun meaning

spoon, refers to the spoon shaped ventral appendage

on first genital segment of male.

Comparative notes. — The male differs from other

species in the genus by the tubercle on fore trochant-

er and the characteristic spoon shaped appendage

ventrally on first genital segment. The female is char-

acterised by the short pregenital segment and the cau-

dal tuft of dark bristles. With the key to oriental spe-

cies by Andersen (1983) this species runs to P. gnoma

Polhemus from Sri Lanka, which is a smaller species

(6 2.1-2.2, 2 2.6-2.8) with different structure of

first genital segment of male. The male of Microvelia

magnifica Lundblad which might, judging from the

figures by Lundblad (1933), also be a Pseudovelia or a

related genus different from Microvelia s. str. also has

a ventral tongue on segment 8, but chis structure is

sharper apically (ventrally) and the anterior trochant-

er is rounded. The female of M. magnifica has the last

segment of connexiva meeting over the tip of abdo-

men.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Pseudovelia mystax sp. n.

(figs. 29-33)

Type material. — Holotype apterous d, Sulawesi

Utara Danaus Aleam 22 Novanml 8D Meg IG:

Zimmermann (ZMAN). — Paratypes, 155 419, same

data as holotype, distributed as follows: 22 (includ-

ing allotype) ZMAN; 2? BPUH; 26 29 PCHG; 16 29

JTPC; 18 22 MBBJ; 36 82 NC; 16 29 RMNH; 6d

209 ze; 1d 19 ZMUC. All apterous.

Description. — Apterous specimens. Dimensions.

Length d 2.20-2.26, 2 2.70-2.87, width (across

metanotum/first abdominal tergite) d 0.80-0.85, ?

0.95-1.003. Colour and pilosity. Generally brick red

to light castaneous, sutures between body sclerites

mostly blackish, coxae, trochanters and most part of

femurs yellowish. Patches of silvery hairs along inner

margins of eyes rather inconspicuous, more distinct

laterally on first abdominal tergite and lining the su-

tures between tergites. Body clothed with quite long,

erect brownish pilosity, more dense laterally and on

venter than on dorsum.

Length of antennal segments I-IV 0.49: 0.32: 0.33:

0.46. Anterior quarter of pronotum and swollen ante-

rolateral angles smooth, posterior three quarters

coarsely punctate. Greatest width of pronotum about

halfway anteroposterior line, lateral margins convex

without pronounced humeral angles or indentation.

Ventral arolia bristle like. Middle tibia with about 5

long (twice as long as the width of tibia) hairs in api-

cal half of inner side.

Male. General shape elongate with a truncate cau-

dal apex, connexiva usually more or less horizontal,

their lateral margins convergent in posterior third.

Length of antenna slightly less than 3/4 the length of

body (1.6/2.25). Pronotum twice as broad than its

median length (0.8/0.4). Anterior trochanter round-

ed. Fore tibia with grasping comb occupying some-

what more than half the length of tibia ((0.4/0.7, fig.

29), consisting of over 250 fine teeth except at the

spur where they are distinctly coarser. Hind tibia with

ciliation along inner margin apically denser but not

modified. First segment of hind tarsus ventrally with

normal pilosity only. First abdominal tergite large, to-

gether with second convex dorsally raised about as

high as the edge of connexiva. Pregenital segment

about one fifth the length of abdomen. First genital

segment ventrally strongly chitinized in posterior

half, otherwise little modified (figs. 30-31). Second

genital segment lying within first with a pair of tufts

(about 6 each) of laterally pointing whisker-like bris-

tles which are visible even when the segment is re-

tracted.

Female. General shape bulkier than male, with

connexiva usually more or less vertical, tapering only

slightly (but more distinctly than in male) posterior-

ly. Length of antenna more than half the length of

body (1.55/2.75). Pronotum twice as wide as its me-

dian length (0.82/0.42), distinctly narrower than

width across metanotum. Abdominal tergites 1-3,

convex dorsally, in lateral view 1 distinctly, 2 just ris-

ing above the margin of connexiva. Caudal apices of

connexiva remaining considerably removed from

each other, leaving tergite 7 visible, tergite 8 oriented

horizontally, fully visible in dorsal view. Pregenital

segment large, its sternite slightly less than one third

the median length of abdominal venter. Genital seg-

ments small, retracted into abdomen, more or less tri-

angular in caudal view (figs. 32-33).

Macropterous form unknown.

Etymology. — Mystax, a noun in apposition mean-

ing moustache or upper lip in Doric, referring to the

‘caudal whiskers’ of the male.

Comparative notes. — This species at first sight re-

sembles P. aflia and P. pyrokrene because of similar

general colour and shape. The bristles on segment 9

of the male separate P. mystax from other Pseudovelia

in tropical Asia. The apex of abdomen in caudal view

of the female is more broadly triangular than in other

species although the difference with P. aflia is very

slight.

Pseudovelia pyrokrene sp. n.

(figs. 34-38)

Type material. — Holotype 3, INDONESIA, Sulawesi

Tenggara, Kecamatan Mowewe, Sungai Mowewe,

lowland stream, 28 Feb. 1989, N8922, leg. N. Nieser

(ZMAN). Paratypes: same data als holotype 38 129

distributed as follows: 19 (allotype) ZMAN, 19 MBB},

19 BPUH, 26 49 NC, 1d 29 PCHC, 29 RMNH, 19

ZC. Road Kolaka-Kendari km 20, mountain stream,

under overhanging rock, 3. Mar. 1989, N8934, leg.

N. Nieser 1 9 ; Sungai Mokowu (+ 3°49’S 121°40’E),

30.198918; same CMOS PCM PIE

van Tol (RMNH). All apterous.

Additional specimens 1 lvV, same data as holotype;

P. Buton, lst mountain stream along road Bau-Bau

to Bunga, 8. III. 1989, N8935, leg. N. Nieser, 1?

apt.

Description. — Apterous specimens. Dimensions.

Length d 2.30-2.44, 9 2.71-2.90, width (across

metanotum/first abdominal tergite) d 0.79-0.80, 9

0.91-0.93. Colour and pilosity. Generally brick red/-

light castaneous to dark brown, sutures between body

sclerites mostly blackish, rostrum, basal part of first

antennal segment, coxae, trochanters and basal half of

femurs lighter, yellowish to light brown. Patches of

silvery hairs, distinctly contrasting, along inner mar-

gins of eyes lateral and caudal margin of first abdom-

inal tergite, most of tergites 2, 3, 6 and 7 less distinct

on tergites 4 and 5. Body clothed with quite sparse

but distinct long pilosity, in addition to the dense

short pilosity covering body.

General shape elongate. Length of antennal seg-

ments I-IV 0.44 : 0.30 : 0.34 : 0.50. Anterior quarter

of pronotum and swollen anterolateral angles

smooth, posterior three quarters coarsely punctate.

Greatest width of pronotum about halfway antero-

posterior line, lateral margins convex without pro-

nounced humeral angles or indentation. Ventral aro-

lia bristle-like. Middle tibia with about 5 long (twice

as long as the width of tibia) hairs in apical half of in-

ner side. First abdominal tergite coarsely punctate.

Male. Connexiva slightly slanting upward lateral-

ly, their lateral margins moderately convex, inner

margins hardly convergent posteriorly. Length of an-

tenna about 2/3 the length of body (1.6/2.35).

Pronotum one and a third times as broad than its me-

dian length (0.74/0.55), slightly narrower than the

width across metanotum. Anterior trochanter with a

nearly perpendicular angle apically (fig. 34). Fore tib-

ia with grasping comb occupying half the length of

tibia ((0.3/0.6), consisting of nearly 150 coarse teeth

except a few proximal ones and a row of 20 just before

the spur, which are narrow. Hind tibia virtually

straight, with ciliation along inner margin apically

denser but a dark patch of small spines absent. First

segment of hind tarsus ventrally with normal pilosity

only. First abdominal tergite large, slightly convex

dorsally raised just above the edge of connexiva.

Pregenital segment about one fifth the length of ab-

domen. First genital segment ventrally with a large

subcircular impression with a chitinized margin in

posterior half, otherwise little modified (figs. 35-36).

Second genital segment lying mostly within first,

without structural peculiarities.

Female. Connexiva vertical in most specimens, dis-

tinctly converging posteriorly. Length of antenna

more than half the length of body (1.58/2.79).

Pronotum 1.4 as wide as its median length

(0.8/0.56), only slightly narrower than width across

metanotum. Abdominal tergites 1 large, somewhat

convex dorsally, in lateral view raised slightly above

the margin of connexiva, tergites 2 and 3 declining,

remainder of abdominal dorsum horizontal. Caudal

apices of connexiva remaining considerably removed

from each other, leaving tergite 7 visible, tergite 8

oriented horizontally, fully visible in dorsal view.

Pregenital segment large, its sternite slightly less than

half the median length of abdominal venter. Genital

segments small, retracted into abdomen, more or less

U-shaped in caudal view (figs. 37-38).

Macropterous form unknown.

NIESER: New species of Veliidae

Etymology. — Pyrokrene, a greek noun in apposi-

tion, refers to the similarity with P. feuerborni.

Comparative notes. — The ventral impression of

first genital segment of male is of similar shape, but

relatively larger, as in P. feuerborni Lundblad, in addi-

tion the male of P. feuerborni has a slightly curved

hind tibia with a group of small spines apically which

is lacking in P. pyrokrene which has hind tibia virtual-

ly straight and the fore tibial comb of P. feuerborni

consists of 45 coarse proximal and a ‘hardly countable

number’ of fine teeth in the distal part (lauter winzi-

gen, kaum zählbaren Dornen, Lundblad 1934: 335).

P. feuerborni is only known from macropterous spec-

imens but in sofar comparison with apterous speci-

mens gives clues, it appears quite related to P. pyro-

krene. In general shape the female reminds of P.

koutali but lacks the tuft of caudally directed dark

bristles on tergite 8.

Pseudovelia sangihe sp. n. (figs. 39-43)

Type material. — Holotype apterous male: INDONE-

SIA: Sulawesi Utara, Pulau Sangihe, Laine, Sungai

Laine at waterfall, at edge of pothole between plants

(Hydrilla) and floating plant debris, shade, ‘Pseu-

dovelia especially at spots with some foam’,

12.X1.1994, N9463, leg. N. Nieser (ZMAN).

Paratypes (adults only), same data as holotype 85 49

apt., 4d 22 macr., 22 larvae III/V distributed as fol-

lows; 12 apt. (allotype) 18 12 macr. ZMAN, 1d

apt., MBBJ, 1d apt. NMSC, Id 19 apt, 16 macr.

RMNH, 1d apt. SEMG; P. Sangihe, Lelepu, Sungai Le-

lepu, mainly lowland stream aspect, shaded pools

under overhanging vegetation, no foam, 13.X1.1994,

N9464, 16 59 apt, 16 macr., 2 lvV (19 MBBJ, 14

19 MUDH, 19 NHMW, 1d macr., 19 SEMC, 19

ZMUC); P. Sangihe, Desa Utaurano, Sungai Apanu-

kang, pothole, at edge between vegetation, no foam,

14.X1.1994, N9465A, 38 69 apt, 19 macr. [S.

Apanukang is an upstream tributary of S. Laine near

Naha, which is different from S. Laine at Laine]; P.

Sangihe, Sungai Masalihe, mountain stream in rather

deep (c. 20m) gorge, Pseudovelia from puddles at

edge of stream bed, 16.XI.1994, N9468, 4d 49

apt.( 14 19 JTPC, 1d NHMW, 19 SEMC, 1d 12 Zc,

1d 19 ZMUC); P. Sangihe, Sungai Kendahe, bridge

at Poto, mountain stream, pools at edge of stream

bed, 17.XI.1994, N9471, 3d 49 apt; P. Sangihe,

smaller stream near N9471, 17.XI.1994, N9473, 64

19 apt., 16 macr., 2 lvV; P. Sangihe, mountain

stream at Bowokulu, rather small stream with rocky

bottom and large boulders, parts with strong current

alternating with virtually stagnant pools, Pseudovelia

at edges under overhanging rocks, 19.X1.1994,

N9477, 26 89 apt. (1d 19 PCHC).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 39-43. Pseudovelia sangihe. Paratypes: 39-43 male, 39-40 segment 8, 39 lateral view, 40 detail of rim of ventral impres-

sion on segment 8 ventrolateral view, 41 fore leg, 42-43 female apex of abdomen, 42 lateral view, 43 caudal view. — 44

Pseudovelia argyropardala. Male paratype hind tarsus. Scale 0.25 mm. — 45 Tarsal segment (P. sexualis) illustrating blade like

ventral arolium. — 46-47 Head of Pseudovelia in lateral view: 46 P. buccula illustrating caudally prolonged ventral lobe, 47 P.

lundbladi with normal ventral lobe. — 48-49 Pseudovelia argyropardala female paratype apex of abdomen, 48 lateral view, 49

caudal view. — 50 Pseudovelia lundbladi male segment 8 in ventral view. — 51-55 Xiphovelia skoteina. Male paratypes: 51 mid-

dle tarsus, 52 fore leg, 53 genital segments lateral view, 54-55 segment 9, 54 lateral view, 55 caudal view. — 56-58 Pseudovelia

kalami holotype male; 56 fore tibia and tarsus, 57-58 segment 8, 57 lateral view, 58 ventral view.

Scale lines for figs. 41, 48, 51: 0.5mm; 39-40, 42-58: 0.25 mm.

Description. — Apterous specimens. Dimensions.

Length d 2.12-2.15, 2 2.36-2.45; width (across ab-

dominal tergites 4-5) & 0.72-0.76, 2 0.92-0.96;

width of head & 0.50-0.51, 2 0.52-0.56. Colour and

pilosity. Dorsally pale orange to pale brick red, head

darker, brownish, anterior third of pronotum lighter;

lateral and ventral parts lighter than dorsum. Coxae

and basal 3/4 of femurs yellowish, remainder of ap-

pendages darker, brownish; sutures between body

sclerites and spiracles blackish, especially on abdo-

men. Bands of silvery hairs along inner margins of

eyes distinct; patches of silvery hairs laterally on first

abdominal tergite, covering most of tergites 2 and 3

and mediocaudally on tergite 6. Body clothed with

rather short light brown pilosity which is more dis-

tinct ventrally than dorsally. Antennae and legs with

usual pilosity.

Length of antennal segments I-IV, 0.43 : 0.28 :

0.31 : 0.47. Anterior third of pronotum smooth,

coarse punctation on posterior two thirds sparse and

rather indistinct. Lateral margins of pronotum

smoothly curved, not or hardly indented, greatest

width on posterior third. Ventral arolia bristle like.

Middle tibia with about seven long erect hairs on in-

ner side of apical half, becoming distinctly shorter ap-

ically. Hind tibia with slightly denser and thicker pi-

losity apically.

Male. Elongate, connexiva roughly horizontal to

slanting slightly upward, lateral margins diverging in

anterior, converging in posterior half. Length of an-

tenna slightly more than two thirds of body length

(1.5/2.1). Pronotum one and a half times as wide as

its median length (0.6/0.4), narrower than width

across metanotum. Anterior trochanter rounded.

Apical comb on fore tibia relatively short, + one third

(0.2/0.6) tibial length (fig. 41), consisting of about

140 teeth of equal width. First segment of hind tarsus

apart from the normal pilosity with 2-3 long (longer

than diameter of segment) cilia ventrally. Abdominal

tergite 1 large, smooth, sloping downward posterior-

ly; tergites 2-7 flat, horizontal, in lateral view con-

cealed by the connexiva. Pregenital tergite + one fifth

the dorsal median length of abdomen. First genital

segment with a large ventral impression, anteriorly

and posteriorly with a pair of spine-like structures,

consisting of closely packed bristles (figs. 39, 40).

Female. More broadly oval than male. Connexiva

nearly horizontal to slanting 0.167 upward, their lat-

eral margin evenly curved with its greatest width on

tergite 4-5. Length of antenna slightly less than two

thirds the length of body (1.5/2.4). Pronotum slight-

ly over one and a half as wide as its median length

(0.65/0.4), distinctly narrower than width across

metanotum. Abdominal tergite 1 large, smooth, slop-

ing downward posteriorly; tergites 2-7 flat, horizon-

tal, in lateral view concealed by the connexiva (except

in gravid females). Caudal apices of connexiva round-

ed, remaining considerably removed from each other,

leaving tergite 7 visible. Tergite 8 oriented horizontal-

ly, fully visible in dorsal view. Pregenital segment

large, its sternite slightly more than one fourth the

median length of abdominal venter (0.3/1.1). Genital

segments relatively large and protruding caudally,

more or less U-shaped in caudal view (fig. 42, 43).

Macropterous form. — Structurally essentially as

apterous form except for the usual modifications of

macropters in the subfamily. Length d 2.15-2.36, 9

2.30-2.35; humeral width of pronotum ¢ 0.85-0.90,

? 0.90-0.92. Pronotum brown with a lighter T-

shaped mark, anterior transverse leg broad, posterior

NIESER: New species of Veliidae

longitudinal leg narrow, disc of pronotum indistinct-

ly punctate. Hemielytra grey-brown with basal cells

largely sordid white and apical cells brownish with

lighter centres and membrane greyish. Venation dis-

tinct, two elongate basal cells and two shorter apical

cells. Lateral keels on basal part of abdomen reaching

to the caudal margin of tergite 4, medial keels reach-

ing onto basal part of tergite 2.

Etymology. — Sangihe is the Sangirese name for the

island where this species was found (Sangir in bahasa

Indonesia).

Comparative notes. — (See key). The male genitalia

are of a similar shape as in P. aflia, P. borneensis and P.

epimekta. Of these P. borneensis has an additional me-

diocaudal ventral projection on first genital segment

(fig. 9); P.epimekta has the first genital segment and

first antennal segment distinctly longer and P. aflia has

the ventral spine-like groups of bristles less developed.

Pseudovelia borneensis Andersen

(figs. 7, 9)

Pseudovelia borneensis. — Andersen 1983: 264, figs. 35-38.

Material. — BRUNEI DARUSSALAM: Labi Road, Kota Teraja,

mountain stream, on foam upstream of log, 15. IV. 1993,

N9343A, 1d 69 apt, 3d 69 macr.(19 apt. 12 macr.

BMKB, 19 macr. NMSC, 19 macr. NHMW, 12 apt. RMNH,

12 apt. ZMAN); Kuala Belalong, small stream behind Field

Study Centre, 16. IV. 1993, N9345, leg. N. Nieser, 18 2?

apt.

Distribution. — Northern Borneo (Sarawak,

Brunei, Sabah).

Remarks. — P. borneensis was described from mac-

ropterous specimens. The apterous form agrees struc-

turally with the macropterous except for absence of

wings and reduction of thorax. Dimensions of apter-

ous form: L d 2.35, 2 2.43-2.52; width of head &

0.55, 2 0.56-0.60; width of pronotum d 0.70, 9

0.72-0.77; width d 0.81, 2 0.82-0.88. The connex-

iva of apterous females are nearly vertical whereas in

macropters they slant laterally. The pattern of silvery

hairs on head and pronotum is identical in both

forms, the patch medially on hind margin of head

may be rather insignificant in some specimens, both

apters and macropters. Apters have patches of silvery

hairs laterally on abdominal tergite 1, nearly the

whole of tergites 2 and 3 and the posterior 2/3 of ter-

gites 6 and 7.

Pseudovelia reiseni Polhemus

(figs. 17-19)

Pseudovelia reiseni Polhemus 1976: 272-273

Material. — PHILIPPINES: Luzon, Laguna, Los Banos,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Table 1 Measurements of leg segments of Microveliinae described in this paper.

femur tibia tarsl tars2

Pseudovelia aflia

d fore leg 0.61 0.57 0.30

& middle leg 0.72 0.73 0.17 0.28

d hind leg 0.81 0.88 0.20 0.27

9 fore leg 0.65 0.53 0.32

? middle leg 0.73 0.71 0.18 0.29

9 hind leg 0.86 0.90 0.19 0.27

Pseudovelia afrofila

d fore leg 0.70 0.60 0.29

d middle leg 0.82 0.74 0.12 0.30

3 hind leg 0.88 0.92 0.16 0.28

9 fore leg 0.69 0.54 0.30

9 middle leg 0.78 0.90 0.14 0.26

2 hind leg 0.82 0.90 0.14 0.26

Pseudovelia argyropardala

3 fore leg 0.63 0.61 0.31

3 middle leg 0.83 0.80 0.15 0.30

3 hind leg 0.84 0.88 0.19 0.32

9 fore leg 0.66 0.60 0.34

® middle leg 0.82 0.81 0.15 0.30

Pseudovelia epimekta

d fore leg 0.85 0.82 0.40

d middle leg 0.96 1.00 0.25 0.30

8 hind leg 1.12 1.30 0.41 0.30

9 fore leg 0.88 0.83 0.40

2 middle leg 0.99 1.34 0.25 0.27

2 hind leg 1.21 1.34 0.42 0.31

Pseudovelia kalami*

d fore leg 0.80 0.75 0.30

d middle leg 1.03 1.04 0.20 0.38

3 hind leg 0.98 1.01 0.33 0.30

Pseudovelia koutali

3 fore leg 0.68 0.59 0.30

3 middle leg 0.82 0.82 0.20 0.30

d hind leg 0.86 0.87 0.24 0.26

9 fore leg 0.69 0.63 0.35

® middle leg 0.93 0.89 0.20 0.30

2 hind leg 0.98 1.06 0.20 0.30

Pseudovelia mystax

d fore leg 0.77 0.68 0.38

3 middle leg 0.93 0.94 0.19 0.33

3 hind leg 0.99 1.11 0.22 0.32

9 fore leg 0.81 0.70 0.38

? middle leg 1.01 0.98 0.20 0.36

2 hind leg 1.08 1.19 0.21 0.33

Pseudovelia pyrokrene

d fore leg 0.70 0.60 0.31

3 middle leg 0.89 0.83 0.19 0.30

3 hind leg 0.91 0.92 0.34 0.30

2 fore leg 0.71 0.59 0.33

? middle leg 0.87 0.82 0.19 0.28

? hind leg 0.98 0.99 0.23 0.31

femur

Pseudovelia sangihe

d fore leg 0.65

3 middle leg 0.83

d hind leg 0.87

9 fore leg 0.61

9 middle leg 0.78

9 hind leg 0.88

Xiphovelia skoteina

d fore leg 0.49

3 middle leg 0.56

d hind leg 0.54

? fore leg 0.52

? middle leg 0.61

9 hind leg 0.59

* Measurements based on single specimen.

stream from Tampalit Falls, 15. XI.1993, leg. H. Zettel, 3d

(25 NHMW, 1d NC); Luzon, Infanta, small pool (10x1.5m,

up to 0.2 deep) at edge of river bed near mouth of Agos

River, many gelatinous algae, 4. IV. 1993, N9335, leg. N.

Nieser, 18 19. Mindanao, Sarangani (S. Cotabato) prov.

Siquel River, pool (area + 20 m’, 0.2 m deep) at edge of riv-

er bed, slightly turbid stagnant water, under overhanging

grassy bank, 23. XI. 1993, N9351, leg. N. Nieser, 3d; Lake

Sebu area, just down stream of second waterfall at edge of

spray area, sheltered edge downstream of boulders, 8. XII.

1993, N9373, leg. N. Nieser, 88 19 (1d BMKB); stream,

just downstream of dam at Lopo, under low overhanging

grassy bank, 9. XII. 1933, N9377B, leg. N. Nieser, 36 19

(28 19 macr.; 1d apt., 1d macr. USCP); irrigation canal at

Lopo, rather fast current, 9. XII. 1993, N9378, leg. N.

Nieser, 1 2 macr. Apterous unless otherwise indicated.

Distribution. — Only known from Luzon and

Mindanao.

Genus Xiphovelia Lundblad

Xiphovelia was erected by Lundblad (1933) to ac-

commodate his X. ensis from Jawa. In addition there

are a few species known from the Far Fast (Esaki &

Miyamoto 1959) and one from Sri Lanka (Polhemus

1979). As these belong to the smaller Microveliinae,

the paucity of described species is probably due to

poor collecting. In specialisation of male genital seg-

ments they show some affınity to Pseudovelia (small

with small parameres) although the genital segments

are much smaller. On the other hand the structure of

antenna with a short stout first segment, slightly nar-

rower second and distinctly narrower third and

fourth and the thoracical structure are quite different.

In this genus the middle tarsus has a blade like ventr-

al arolium and meso- meta- notum and first abdomi-

nal tergite are fused medially, in the description me-

dial length of mesonotal area refers to the median

NIESER: New species of Veliidae

tibia tarsl tars2

0.58 0.28

0.79 0.17 0.30

0.95 0.21 0.26

0.57 0.29

0.78 0.15 0.29

0.96 0.18 0.25

0.39 0.20

0.51 0.18 0.26

0.64 0.13 0.19

0.39 0.24

0.53 0.19 0.28

0.69 0.13 0.22

length of these three tergites together. As stated by

Andersen (1982) character states of genera of

Microveliinae have not yet sufficiently been studied

to get a clear picture about their relationship.

Xiphovelia skoteina sp. n.

(figs. 51-55)

Type material. — Holotype apterous d INDONESIA:

Sulawesi Utara, Lakes I, Quellbereich (source area),

PPA Bungalow, Nov. 1985, leg. G. Zimmermann

(RMNH). (The PPA bungalow is just N of the W tip of

Danau Mooat). Paratypes, 516 16% all apterous,

same data as holotype, distributed as follows: 24 19

(allotype) RMNH; 24 BPUH; 2d 29 PCHG; 26 19

TEC 2S) IO IMBBT IS OM2P NC: 208 7.22 2673.5

19 ZMAN; 2d 19 ZMUC. Dumoga Bone N. P.

Tumpah River Beach, 19. Oct. 1985, leg. G.

Zimmermann, 4d (2d NC, 26 Zc).

Description. — Apterous specimens. Dimensions.

Length d 1.80-1.90, 2 2.00-2.18, width (across

metanotum/first abdominal tergite) & 0.73-0.78, ®

0.90-0.99. Colour and pilosity. Generally dull dark

ash-grey (anthracite), eyes shining, base of first anten-

nal segment, segments 2 and 3 of rostrum , coxae, tro-

chanters and basal part of femurs yellowish, remain-

der of appendages castaneous. Pronotum with an

uninterrupted lighter band, about as wide as head, in

most specimens only slightly contrasting with back-

ground. Body covered with well developed pilosity,

yellowish, appressed, relatively fine and short on

body, longer, more erect and somewhat darker on ap-

pendages, eyes bare.

Head slightly less than twice as wide as its median

length (0.5/0.3). Length of antennal segments I-IV,

0.19: 0.19: 0.22: 0.31. Width of third antennal seg-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

ment two third that of fourth (0.02/0.03). Body wall

with shallow punctures, posteriorly more shallow and

hence less distinct than anteriorly. Pronotum short,

about five times as wide as long. median length of

mesonotal area one fourth the median length of body

(8 0.45, 9 0.52). Ventral arolia of fore and hind tar-

sus bristle like. Ventral arolium of middle tarsus leaf-

like (fig. 52). Middle and hind trochanter of subequal

length. Middle tibia with about seven long hairs

(apart from normal pilosity) in apical half of inner

side, their length compared to the width of tibia

(which is stout in both sexes) varying from slightly

longer more distally to slightly shorter apically.

Male. General shape elongate oval, greatest width

at posterior margin of metanotum, connexiva more

or less horizontal, their lateral margins only slightly

convergent posteriorly. Apical part of genital seg-

ments in dorsal view visible as a small caudal button

(in most alcohol specimens genital segments expand-

ed). Length of antenna slightly less than half the

length of body (0.89/1.86). Measurements of leg seg-

ments, table 1. Anterior trochanter with an ventro-

apical knob (fig. 51). Dorsal margin of fore femur

straight in apical three fourth. Dorsal (‘outer’) margin

of fore tibia convex, grasping comb about half as long

as tibia, more distinct in expanded apical part.

Genital segments (figs. 53-55) in caudal view with

comparatively low lateral ‘ears’.

Female. General shape broadly oval, connexiva

nearly vertical in most specimens, especially in poste-

rior half where they are distinctly converging. Legs

generally more slender than male, measurements of

leg segments table 1. Anterior trochanter rounded,

anterior tibia with an apical tuft os bristles, mimick-

ing the apex of tibial comb of male. Abdominal ter-

gite 7 somewhat longer than tergite 6, tergite 8 hori-

zontal. Sternite 7 large, its median length one third

the median length of abdominal venter. Genital seg-

ments retracted into abdomen.

Etymology. — Skoteinos, a Greek adjective meaning

dark, sombre, refers to the general colour of the spe-

cies.

Comparative notes. — In the key to Xiphovelia by

Esaki & Miyamoto (1959) this species runs to X. bon-

inensis Esaki & Miyamoto, due to the lack of shiny

(‘silvery’) pilosity. Males of X. skoteina differ from all

species described so far by the ventroapical tubercle of

fore trochanter. Females of X. boninensis have the

connexiva reflexed over the abdomen in caudal half,

whereas in X. skoteina these are at most vertical.

Moreover in X. skoteina the length of middle and

hind trochanter is subequal while in other species the

middle trochanter is distinctly longer than hind tro-

chanter.

DISCUSSION

Andersen (1983) deals with ten species of

Pseudovelia from the Oriental Region (in addition

seven taxa from the E. Palaearctic are listed). So in

this paper the number of Oriental species is nearly

doubled. In addition Dr. Zettel (NHMW) writes me

that he has several apparently undescribed species

from the Philippines. Apparently this is a, still under-

collected, species rich genus in the area. They are usu-

ally found in shaded places (including overhanging

banks or rocks) at the edges of streams or, more rare-

ly, lakes. They are quite often found at places where

some foam is assembled (probably because food is

trapped in the foam).

Considering primarily the development of the

male genital segments the genus Pseudovelia seems to

split into two groups. The first group is the hypodon-

ta-group of Andersen (1983), which is primarily

characterized by 4-5 spine-like processes along the

margin of the ventral impression of the first genital

segment of male. On closer examination, these pro-

cesses consist of a basal chitinous stump on which a

small, acute narrowly packed tuft of bristles is placed.

They also agree more or less in general colour pattern

and body shape. The hypodonta-group consists of P.

aflia, P. borneensis, P. buccula, P. epimekta, P. hypo-

donta and P. sangihe. Associated with this group is P.

koutali whereas P. lundbladi which has anteriorly a

broad ventral process which, although much shorter,

reminds somewhat of the process in P. koutali and

caudally three processes of the same type as found in

the hypodonta-group may be considered a link

between P. koutali and the hypodonta group. The sec-

ond group which could be named feuerborni-group

has the ventral impression on first genital segment of

male bordered by a more or less distinct chitinous rim

which can be fringed with hairs whereas the impres-

sion itself may also be clothed with hairs. This group

contains: P. afrofila, P. argyropardala, P. feuerborni, P.

gnoma, P. longitarsa, P. mystax, P. pyrokrene, P. reise-

ni, P. sexualis and according to Andersen (1983) P.

tjurpensis. Whereas the hypodonta-group is compara-

tively homogeneous, the feuerborni-group can be di-

vided in four pairs of similar species with P. mystax

with its rather simple first genital segment and whisk-

ers on the second as the odd one out. The first pair is

formed by P. longitarsa and P. sexualis which have in

common blade-like ventral arolia and strongly hairy

eyes. The second pair consists of P. argyropardala and

P. reiseni, which have in common a ventral impres-

sion of first genital segment without strong rim and

sublateral patches of hairs inside the impression. In

addition the males have the ‘swimming hairs’ on first

segment of hind tarsi more strongly developed than

in other species and both are dark grey with very con-

trasting patches of silvery hairs which are arranged in

roughly the same pattern. These two are also geo-

graphically more restricted than the other pairs, P.

Karakelong is closer to the Philippines than to the

main islands of Indonesia. P. feuerborni and P. pyro-

krene have the ventral impression on first genital seg-

ment somewhat similar to P.argyropardala/reisent (al-

though the chitinous rim is somewhat better

developed) but have the brick red colour of most

Pseudovelia and distinctly less developed ‘swimming

hairs’ on first segment of hind tarsi of male. Finally

P. afrofila and P. gnoma are both smaller and more

oval than the others and are generally of a dark grey

colour with a contrasting transverse rectangular

orange mark anteriorly on pronotum. P. kalami fi-

nally is quite aberrant, apart from the fore tibia the

antennal structure and the large depression in male

venter are different from other species. This species

might turn out to represent a distinct subgenus or

closely related genus.

ACKNOWLEDGEMENTS

Thanks are due to Drs: J. P. Duffels (ZMAN), J. van

Tol (RMNH), H. Zettel (NHMW) and G. Zimmer-

mann (ZC), for the loan of specimens. And Dr. J. T.

Polhemus (JTPC) for discussion of the identity of

some specimens.

REFERENCES

Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera,

Gerromorpha). — Entomonograph 3: 1-455.

NIESER: New species of Veliidae

Andersen, N. M., 1983. The Old World Microveliinae

(Hemiptera: Veliidae) I. The status of Pseudovelia

Hoberlandt and Perivelia Poisson, with a review of Oriental

species. — Entomologica Scandinavica 14: 253-268.

Arnett, jr., R. H., Samuelson, G. A., & Nishida, G. M.,

1993. The insect and spider collections of the world. —

Flora & Fauna Handbook 11: i-v, 1-308.

Esaki, T., & Miyamoto, S., 1959. Veliidae of Japan and its

adjacent territory (Hemiptera-Heteroptera) II. Xiphovelia

Lundblad. — Sieboldia 2: 91-108, pls.10-13.

Lundblad, O., 1933. Zur Kenntnis der aquatilen und semi-

aquatilen Hemipteren von Sumatra, Java und Bali. — Archiv

fiir Hydrobiologie Supplement 12: 1-195, 263-488.

Nieser, N. & Chen, P. P., 1991. Naucoridae, Nepidae and

Notonectidae, mainly from Sulawesi and Pulau Buton

(Indonesia). — Tijdschrift voor Entomologie 134: 47-67.

Nieser, N. &2 Chen PR 91993) Ihe Rhagovelia

(Heteroptera: Veliidae) of Sulawesi (Indonesia). —

Tijdschrift voor Entomologie 136: 259-281.

Polhemus, J. T., Pseudovelia reiseni Polhemus n. sp. p. 272-

273, fig. 2. — In J. T. Polhemus & W. K. Reisen, 1976.

Aquatic Hemiptera of the Philippines. — Kalikasan,

Philippine Journal of Biology 5: 259-294.

Polhemus, J. T., 1979. Results of the Austrian-Ceylonese

Hydrobiological Mission 1970. XIX: Aquatic and semi-

aquatic Hemiptera of Sri Lanka from the Austrian Indo-

Pacific Expedition, 1970-71. — Bulletin of the Fisheries

Research Station, Sri Lanka 29: 89-113.

Zimmermann, G., 1984. Heteroptera aus dem Nepal-

Himalaya. Geovelia n. gen. eine Gattung terrestrischer

Microveliinae (Insecta: Gerromorpha: Veliinae). —

Senckenbergiana Biologia 65: 65-74.

Received: 18 January 1995

Accepted: 7 March 1995

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

JOHN D. OSWALD

Department of Entomology, National Museum of Natural History, Smithsonian Institution,

Washington D. C.

REVISION OF THE SOUTHEAST ASIAN SILKY

LACEWING GENUS BALMES (NEUROPTERA:

BSIXCHIORSIDAE)

Oswald, J. D., 1995. Revision of the southeast Asian silky lacewing genus Balmes (Neuroptera:

Psychopsidae). — Tijdschrift voor Entomologie 138: 89-101, figs. 1-28, tab. 1. [ISSN 0040-

7496]. Published 15 June 1995.

The southeast Asian psychopsid genus Balmes is comprehensively revised for the first time. Four

species are recognized: birmanus, formosus, notabilis, and terissinus. Each species is redescribed,

illustrated, and keyed. Balmes notabilis is removed from the synonymy of birmanus and re-es-

tablished as a valid species. The four species are phylogenetically (cladistically) related as fol-

lows: (formosus + (terissinus + (notabilis + birmanus))).

Dr. John D. Oswald. Present address: Department of Entomology, 412 Minnie Belle Heep

Building, Texas A&M University, College Station, TX 77843-2475, United States of America.

Key words. — Oriental region, phylogeny, taxonomy, key.

The family Psychopsidae, silky lacewings, is a

monophyletic (=holophyletic) clade containing five

genera and 26 extant species (Oswald 1993). The dis-

tribution of extant psychopsids is relictual, being re-

stricted to southern Africa, Australia, and southeast

Asia. Fossil taxa attributed to this family are currently

known from North America, Europe, Asia, and

Australia. Psychopsids are of special interest because

of their highly disjunct present geographic distribu-

tion, their unusual female ovipositional habits, and

their phylogenetic position as the basalmost clade of

the neuropterous superfamily Myrmeleontoidea, i.e.,

as the sister-group to the Nymphidae + Nemopteri-

dae + Myrmeleontidae + Ascalaphidae (Mansell

1992; Oswald, unpublished data).

Since the publication of my earlier monograph of

this family (Oswald 1993), I have had the opportu-

nity to examine additional material of the little-

known Oriental genus Balmes Naväs. The recent tax-

onomic history of this genus can be summarized as

follows. Kimmins (1939) recognized three Balmes

species: birmanus (as ‘birmana [sic], with terissinus

and notabilis as synonyms), formosus (as ‘formosana

[sic]), and gallardi. New ([1989]) demoted Balmes to

a junior subjective synonym of the otherwise

Australian endemic genus Psychopsis, and treated bir-

manus, formosus, and gallardi as valid species of

Psychopsis. Oswald (1993) (1) re-erected the genus

Balmes on the basis of evidence supporting its mono-

phyly and the existence of a sister-group relationship

berween Balmesand Psychopsis, (2) established the va-

lidity of B. terissinus and removed it from the synony-

my of birmanus, and (3) noted the existence of a

fourth, at that time apparently undescribed, Balmes

species in China (the ‘bnsl’ of Oswald 1993).

The species ‘bnsl’ has since proven to be conspecif-

ic with notabilis, which is herein recognized as a valid

species and removed from synonymy with birmanus.

Thus, four valid species of Balmes are treated here: B.

birmanus (McLachlan), B. formosus (Kuwayama), B.

notabilis Naväs and B. terissinus Naväs. The current

revision contains redescriptions and illustrations of

each species, a key to distinguish among the species,

and a phylogenetic (cladistic) analysis of interspecific

relationships within Balmes. The latter extends the

prior analysis of Oswald (1993) through the incorpo-

ration of new data for the Taiwanese species B. formo-

sus.

MATERIALS AND METHODS

Material and collection acronyms. — This study is

based upon approximately 40 adult Balmes specimens

in the following collections: BMNH, The Natural

History Museum, London, England; CUIC, Cornell

University Insect Collection, Ithaca, NY, U.S.A;

EIHU, Insect Collection, Hokkaido University,

Sapporo, Japan; MCZC, Museum of Comparative

Zoology, Cambridge, MA, U.S.A; MNHP, Muséum

National d'Histoire Naturelle, Paris, France; USNM,

National Museum of Natural History, Washington,

DC, U.S.A.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Illustrations. — Line drawings were executed with the

aid of a drawing tube attached to a dissecting micro-

scope. Terminalic illustrations were made from cleared,

and generally stained, preparations temporarily mount-

ed in glycerin. Setae are not shown. In illustrations of the

gonarcus/mediuncus/9th gonocoxite complex, mem-

branes attached along the antextragonarcal commissure

and the venter of the 9th gonocoxites are shown

stretched in a manner calculated to best convey their

lines of attachment, not necessarily as zn situ.

Terminology. — General entomological terminolo-

gy follows Nichols (1989). Terms for terminalic

structures follow Oswald (1993).

Annotations. — The following annotations are used

in the synonymical listings: Dst, distribution; FT, fe-

male terminalia; FW, forewing; H, habitus; Lst, list

or listed; MT, male terminalia; Nom, nomenclature;

OD, original publication/description; RD, redescrip-

tion; Tax, taxonomy; W, wing. An asterisk (*) follow-

ing an annotation indicates a figure (e.g., FW*, fo-

rewing figure).

Miscellaneous. — Species are treated in alphabetical

order. Forewing lengths were measured from the

proximal margin of the tegula to the wing apex (4.1

mm). Unless otherwise cited, reported flight periods

are the earliest and latest dates of collection of adults

as indicated by label data. Bracketed collection local-

ity names and latitude and longitude coordinates

found in the material examined sections have been

taken principally from the Official Standard Names

Gazetteers of individual southeast Asian countries,

which were compiled by the U.S. Defense Mapping

Agency Topographic Center for the U.S. Board on

Geographic Names.

SYSTEMATIC PART

Genus Balmes Navás

Balmes Navás, 1910: 85 (Type species: Balmes terissinus

Navás, 1910: 85, by monotypy): Navás 1917 (Tax);

Kimmins 1939 (Tax); New [1989] (Nom); Oswald 1993

(RD, Tax).

Diagnosis. — The only extant genus of psychopsids

known from southeastern Asia. Distinguished from

the Australian genus Psychopsis (except P. gallardı) by

the absence in Balmes of a prominent dark macula sit-

uated distally on each hind wing (fig. 3). Based on il-

lustrations contained in New ([1989]), males of P.

B. birmanus

B. formosanus

B. notabilis

B. terissinus

Fig. 1. Geographic distributions of Balmes species and cladogram of interspecific relationships. Only accurately located re-

cords have been plotted.

gallardi (unique holotype in BMNH, not examined)

may be distinguished from males of Balmes species by

(1) the presence in gallardi of an elongate [not short]

Ith gonocoxite costa, and (2) the absence in gallardi

of the male 9th gonocoxite superprocesses modified

into a transverse tumulus.

Description. — Head: Ocelli absent, but vertex

bearing 2 well-developed ocellar/cranial pulvinae.

Forewing: length: 10.8 - ca. 21 mm; coloration:

brown mottling on a hyaline ground; forewing costal

gradate series absent or reduced (i.e, <6 crossveins),

except in formosus where it is well developed (>20

crossveins).

Hind wing: hyaline, without a dark distal macula.

Male terminalia: 8th sternite (fig. 6): without a

posteromedian lobe; 9th tergite (fig. 6): without free

posteroventral processes; Ith sternite (figs. 6, 7): apex

rounded (fig. 7) or emarginate (fig. 13), not narrow

and parallel-sided in ventral view; gonarcus (figs. 9,

10): intragonarcus present but narrow relative to ex-

tragonarcus; extragonarcus composed of a posteriorly

protruded dorsal plate, the extragonopons (egps), and

a pair of lateral narrow to broad extrahemigonarcus

(ehgs); mediuncus (med) weakly decurved, not recur-

ved; 9th gonocoxites (figs. 10, 11): fused medially in-

to a transverse sclerite which articulates with postero-

ventral angles of hemigonarcus; ventral costae (vc)

absent or reduced to short thickenings; superproces-

ses (spp) present but fused into a transverse elevation

on dorsal surface of conjoined gonocoxites (elevation

produced as an attenuate process in notabilis); ventro-

lateral lobes (vll) sometimes present; miscellaneous:

gonosaccal membrane bearing one (fig. 8) or two

(figs. 21, 22) sets of bilaterally paired spiculate lobes;

subanale (sa) present (figs. 9, 10).

Female terminalia: Posterior margin of 7th sternite

medially depressed and emarginate (figs. 26, 27); 7th

and 8th sternites fused, but path of fusion marked by

a suture; copulatory fovea approximately hemispheri-

cal, overhung posteriorly by anteromedian margin of

8th sternite (figs. 26, 27); 9th gonocoxites: without a

longitudinal row of stiff setae below each gonocoxal

costa, without a compact aggregation of setae borne

adjacent to insertion of stylus, suprastylar setae >50%

cochleariform; bursa without lateral corniform diver-

ticulae; spermatheca sigmoid in lateral view, lacking

distinct hollow ventrolateral lobes; two bursal acces-

sory glands present, but form and insertion of ducts

on bursa varies interspecifically.

Distribution (fig. 1). — Southeast Asia (reported here

from Taiwan, southern China, northeastern Burma,

northern Laos, and northern Vietnam). The three

southeast Asian species of Balmes appear to form a

complex of allopatric or parapatric species. The few el-

evation records available for Balmes species (all for

mainland species) range between 600 and 1825 me-

OSWALD: Revision of Balmes

ters, suggesting that Balmes species are montane. This

conclusion is also supported by the apparent geograph-

ic restriction of mainland Balmes species to the upland

areas of the countries in which they occur.

Biology and immature stages. — Unknown.

Species (4). — birmanus. Burma, China [Yunnan

Prov.]; formosus. Taiwan; notabilis China [Yunnan

Prov.], Laos, Vietnam; terissinus China [Sichuan

Prov.].

Classification. — The following sequenced classifi-

cation is recommended for this genus:

Genus Balmes Naväs

B. formosus (Kuwayama, 1927)

B. terissinus Navás, 1910

B. notabilis Navás, 1912

B. birmanus (McLachlan, 1891)

Etymology. — From the surname of Catalan philos-

opher Jaime Luciano Balmes [1810-1848], see Navás

1910: 85. Gender: Masculine, implied from the orig-

inal combination Balmes terissinus, Art. 30d.

Key to adult Balmes species

Notes: The male of formosus and the female of

notabilis are unknown.

1. Forewing (fig. 3): costal gradate series absent (<6

crossveins/wing), intraradial area crossed by 2 gra-

date series, innermost series not bordered by fus-

cous maculae (light brown shading or mottling

may be present) [continental southeast Asia]....... 2

— Forewing (fig. 2); costal gradate series present

(>20 crossveins/wing), intraradial area crossed by

3 well-developed gradate series, innermost series

bordered by fuscous maculae [Taiwan and conti-

nentallsouthease Asia] ee B. formosus

2. Forewing (figs. 4, 5): membrane immaculate or

evenly patterned with light brown mottling,

strongly contrasting darker brown maculae ab-

sent; Male Terminalia: ventrolateral lobes of male

Ith gonocoxites present (figs. 11, 18) [continen-

talksoutheastAstal m ee a 3

— Forewing (fig. 3): membrane marked with 2

prominent rows of brown maculae (one centered

on the ‘vena triplica’, the other on the mediocu-

bital and intracubital spaces), each row with 2 or

3 maculae that strongly contrast with the light

brown to hyaline ground colour of the membrane

(fig. 3); hind margin of wing also often with a

row of contrasting maculae; Male Terminalia:

ventrolateral lobes of male 9th gonocoxites absent

(figs. 24, 25) [China: Sichuan Province] ............

En HER die EB B. terissinus

3. Male 9th sternite: apex rounded (fig. 7); Male 9th

gonocoxites: ventrolateral lobes digitate and

bluntly pointed distally, apices not membrane

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 2-5. Balmes spp., wings. 2, B. formosus (paratype); 3, B. terissinus 4, B. notabilis 5, B. birmanus. Scale bars = 5 mm.

margined (fig. 11); superprocesses fused into a

broad transverse prominence (fig. 11) [Burma,

China: Yunnan Province] ............... B. birmanus

— Male 9th sternite: apex conspicuously emarginate

(fig. 13); Male 9th gonocoxites: ventrolateral

lobes plate like and broadly rounded distally, api-

ces membrane margined (fig. 18); superprocesses

fused and projecting medially as a short process

(figs. 16, 17) (China: Yunnan Province, Laos,

Vietnam eeeh Mens nes B. notabilis

Balmes birmanus (McLachlan)

(figs. 5-11)

Psychopsis birmana McLachlan, 1891: 320 (OD). — Krüger

1922 (Lst); New [1989] (RD, Dst, MT*, FT*, W*).

Balmes birmanus (-a [sicl). — Navás 1930 (Lst, Dst);

Kimmins 1939 (Lst, Dst); Oswald 1993 (Lst).

Diagnosis. — Distinguished from all other Balmes

OSWALD: Revision of Balmes

species by the following combination of characters:

(1) forewing intraradial area traversed by two [not

three] gradate series (fig. 5), (2) apex of male 9th ster-

nite rounded [not conspicuously emarginate] (fig. 7),

and (3) ventrolateral lobes of male 9th gonocoxites

present [not absent] (fig. 11).

Description. — Forewing (fig. 5): length: 11.2 -

15.7 mm (mean = 12.5 mm, n = 9 wings); markings:

light brown mottling on a hyaline ground, markings

somewhat darker along ‘vena triplica’ and along cubi-

tal and mediocubital spaces, distal margin often with

small brown patches bounded by narrow hyaline arcs

Figs. 6-11. Balmes birmanus, male terminalia. 6, abdominal apex, lateral; 7, 9th sternite, ventral; 8, ventral spiculate lobe of

gonosaccus, lateral; 9, medial portion of gonarcus, dorsal; 10, gonarcus / 9th gonocoxite complex, lateral; 11, fused 9th go-

nocoxites, posterodorsal. Abbreviations: 7s, 8s, 9s, sternites; 7t, 8t, It, tergites; Igcx, Ith gonocoxite(s); cc, cercal callus; ect,

ectoproct; ehgs, extrahemigonarcus; egps, extragonopons; mas, mediuncal accessory sclerite; med, mediuncus; sa, subanale;

spp, superprocessus; vc, ventral costa; vll, ventrolateral lobe.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

1 mm 12-13

1mm 14-18

Figs. 12-18. Balmes notabilis, male terminalia. 12, abdominal apex, lateral; 13, 9th sternite, ventral; 14, ventral spiculate lobe

of gonosaccus, lateral; 15, medial portion of gonarcus, dorsal; 16, gonarcus / 9th gonocoxite complex, lateral; 17, medial pro-

cess of 9th gonocoxites, posterior; 18, fused 9th gonocoxites, posterodorsal.

(these lacking or inconspicuous in rubbed wings); co-

stal gradate series absent (1-2 adventitious costal

crossveins occasionally present); intraradial area tra-

versed by two gradate series.

Male terminalia (figs. 6-11): 9th sternite: posterior

margin rounded, not sagittally emarginate; gonarcus:

intragonarcus narrow dorsally, broadened toward an-

teroventral angles, anterior and ventral margins

marked internally by thickened costae; extrahemigo-

narcus broad; midline and posterior margin of extra-

gonopons marked by a T-shaped costa internally,

posterior margin transverse; mediuncus: length ap-

proximately twice average width, slightly constricted

basally; apex rounded or obtusely angulate with a nar-

row sagittal emargination; 9th gonocoxites: superpro-

cesses united sagittally to form a low transverse eleva-

tion; elevation bilobed in posterior view due to a shal-

low median emargination; ventrolateral lobes present

as a pair of free, attenuated, and medially convergent

lobes; spiculate gonosaccal lobes: one pair present on

ventral surface of bursa.

Female terminalia: 7th sternite: parafoveal lobes

weakly developed, not prominently projecting; 8th

sternite: reduced, transverse, expanded medially, at-

tenuate laterally, anterior margin not emarginate sa-

gittally; bursal accessory gland ducts: fused proximal-

ly, attached to sagittodorsal surface of bursa as a single

common duct.

Distribution (fig. 1). — Mainland southeast Asia

(reported here from northeastern Burma and the ad-

jacent portions of Yunnan Province, China).

Flight period. — 14 March - April (no day record-

ed).

Primary type. — Psychopsis birmana. Holotype ?

(BMNH), examined. Type locality: ‘Birmah’ [=BUR-

MA, precise locality unknown). Verbatim label data:

‘Type.’ [pink rectangle], Type [white circle with red

border], ‘Birmah’ [script, white rectangle],

‘McLachlan Coll. / B.M. 1938-674 [blue rectangle],

‘Psychopsis / birmana ML.’ [McLachlan script, white

rectangle], ‘Holotype / Psychopsis / birmana det. /

J.D. Oswald 1994 [red rectangle]. Condition: parts

of the following missing — both antennae, left fore-

and hind legs and right foreleg. Minor insect pest

damage. Pinned through thorax, wings spread.

Terminalia macerated in KOH by Oswald in 1994,

stained with Chlorazol Black, and placed in a glyce-

rin-filled microvial pinned below specimen. This is

the single specimen upon which the description of

birmana was based (McLachlan 1891: 321).

Other material examined (69, 3% = 9). — BURMA:

Mandalay divisioni 18, 29, Maymyo [22°02°N

96°28’E], iv.1912, Mackwood; 14.iii.[19]01, Bar-

row; no date, Scott (BMNH). Shan state 26, Kolaw, S.

Shan States, iv.1916, 1215m (4000’), Mackwood

(BMNH); 19, Loimwe [Loi Mwe, 21°11’N 99°46’E],

iv, 1520-1825m (5000-6000°), Kingford (MCZC).

CHINA: Yunnan province. 13, bet[ween]. Tengyueh

[=Tengchong, 25°02°N 98°28’E] and Nan Tien,

‘1909-10’, Brown (CUIC); 24 , Yun Hsien [=Yunxian,

24°25’N 100°06 E], iv.1942, Jellison (USNM).

Etymology. — Unexplained, probably derived from

the type locality “Birmah’ [=Burma].

Balmes formosus (Kuwayama)

(figs. 2, 27)

Psychopsis (Orientichopsis) formosa Kuwayama, 1927: 123

(OD, H*).

Balmes formosana [sic]. — Kimmins 1939 (Lst, Dst).

Psychopsis formosa. — New [1989] (RD, Dst, FT*, W*).

Balmes formosa [sic]. — Oswald 1993 (Lst).

Diagnosis. — Distinguished from all other Balmes

species by the presence of three [not two] gradate se-

ries traversing the intraradial area of the forewing (fig.

2).

Description. — Forewing (fig. 2): length: ca. 21 mm

(n = 1 wing); markings: predominantly hyaline, cen-

ter of disk with two fuscous maculae, one (proximal)

nearly circular and one (distal) elongate and contin-

ued anteriorly across ‘vena triplica’, an additional

small fuscous dot in middle of intraradial space just

beyond middle gradate series, also with scattered pal-

OSWALD: Revision of Balmes

er brown markings, particularly along cubital and

mediocubital spaces; costal gradate series with more

than 20 crossveins; intraradial area traversed by three

gradate series.

Male: Unknown.

Female terminalia (fig. 27): 7th sternite: parafoveal

lobes strongly developed, prominently projecting; 8th

sternite: a small cordate plate, anterior margin sagit-

tally emarginate; bursal accessory gland ducts: two

free ducts inserted bilaterally symmetrically on dorsal

surface of bursa at a pair of widely separated (more

than two duct diameters) points.

Distribution (fig. 1). — Taiwan.

Flight period. — August.

Primary type. — Holotype ® [not d] (EIHU), not

examined. Type locality: TAIWAN: ‘Rengechi, near

Horisha, Taichiu-district, Formosa’ [Horisha = TAI-

WAN: Nan-t'ou hsien: Pu-li, 23°58’N 120°57’E].

Material examined (19 paratype). — TAIWAN:

Chia-i hsien 19, ‘Kagi [=Chia-i, 23°29N

120°27’E], viii.1921, Hirayama (EIHU).

Etymology. — Unexplained, probably derived ei-

ther from Latin formosus, beautiful, or the island

name Formosa (=Taiwan). Name incorrectly treated

by Oswald (1993) as a noun in apposition, based on

the island name Formosa. Since ‘formosa’ is a valid

Latin adjective, the termination of a species-group

name based on this word must match the gender of

the genus-group name with which it is combined,

Art. 31b.

Comments. — Balmes formosus is known only from

the two females (not males as stated by Kuwayama) of

the type series.

Balmes notabilis Navas, stat. n.

(figs. 4, 12-18)

Balmes notabilis Navas, 1912: 197 (OD). — Navas 1917

(RD).

‘bnsl’. — Oswald 1993 (Lst).

Diagnosis. — Distinguished from all other Balmes

species by the following combination of characters:

(1) forewing intraradial area traversed by two [not

three] gradate series (fig. 4), (2) apex of male 9th ster-

nite conspicuously emarginate [not rounded] (fig.

Description. — Forewing (fig. 4): length 12.5 - 14.0

mm (mean = 13.2 mm, n = 4 wings); markings: very

similar to birmanus, light brown mottling on a hya-

line ground, markings somewhat darker along ‘vena

triplica’ and along cubital and mediocubital spaces,

hyaline-bounded brown patches of distal margin (see

birmanus description) obscure to absent; costal gra-

date series absent; intraradial area traversed by two

gradate series.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 19-27. Balmes terissinus, male terminalia. 19, abdominal apex, lateral; 20, 9th sternite, ventral; 21, dorsal spiculate lobe

of gonosaccus, lateral; 22, ventral spiculate lobe of gonosaccus, lateral; 23, medial portion of gonarcus, dorsal; 24, gonarcus /

9th gonocoxite complex, lateral; 25, fused 9th gonocoxites, posterodorsal. 26-27, Balmes spp., female 7th (medial portion)

and 8th sternites, ventral. 26, B. terissinus, 27, B. formosus.

Male terminalia (figs. 12-18): 9th sternite: posteri-

or margin prominently emarginate medially; gonar-

cus: intragonarcus narrow throughout, broadest at

anteroventral angle; extragonopons with a strong sag-

ittal costa internally, posterior margin with a pair of

rounded lobes divided by a broad medial excavation;

extrahemigonarcus composed of a broad proximal re-

gion and a narrower distal region, the two regions be-

ing continuous anteroventrally but separated distal to

this region by an approximately 60° angle in the pos-

terior margin of the hemigonarcus; mediuncus: apical

2/3 to 3/4 slender and nearly parallel sided, base

flared and broader; apex bifid (condition usually not

obvious because paired apical lobes usually lie parallel

with their medial margins adpressed); 9th gonocox-

ites: superprocesses united and produced medially to

form a short digitate process; apex of process distinct-

ly forked (as shown in fig. 17) to only slightly emargi-

nate; anterior margin of transverse gonocoxal bar lat-

eral to median process usually with a pair of low,

setose, convexities; ventrolateral lobes present as a

pair of shallowly concavo-convex, membrane-mar-

gined, spathulate lobes; spiculate gonosaccal lobes:

one pair present on ventral surface of bursa.

Female: Unknown.

Distribution (fig. 1). — Mainland southeast Asia

(reported here from southern China [Yunnan

Province], northern Laos and northern Vietnam).

Flight period. — 22 April - 2 June.

Primary type. — Lectotype d (BMNH), designated

by New ([1989]) by inference of holotype, examined.

Type locality: PEOPLE’S REPUBLIC OF CHINA: probably

Yunnan Province, see discussion below. Verbatim la-

bel data: ‘Type’ [white circle with red border], ‘ma-

munf[?], yuman [=? Yunnan] / H. E. Hobson / 1900-

192° [white rectangle, text in somewhat illegible

script], ‘Balmes | notabilis | Navläs].. [blue-lined

white notebook paper, rectangle with corners clipped,

in Naväs (?) script], “Lectotype / Balmes | notabilis

det. / J.D. Oswald 1994 [red rectangle]. Condition:

parts of left fore- and midlegs and right midleg miss-

ing, wings somewhat tattered. Stage mounted, wings

spread, probably formerly extracted from alcohol.

Terminalia macerated in KOH and placed in a glyce-

rin-filled microvial pinned below specimen.

Naväs (1912) cited two specimens in the type series

of notabilis, one in the British Museum and one in his

personal collection. The BMNH specimen cited by

New ([1989]:845) as the ‘holotype’ must, conse-

quently, be considered a lectotype, Art. 74b. The

specimen stated to have been in Naväs’ collection has

not been traced. The precise type locality of notabilis

is uncertain. In the original description Naväs cited a

single locality, ‘China, Junam’ (referred to by Naväs

1917 as ‘China: Yu-nam’), but this site does not

match the text of the lectotype’s locality label. It is

possible that this locality was taken from the unseen

specimen retained in Naväs’ collection. Attempts to

identify Junam’ or ‘Yu-nam’ in several Chinese gaz-

etteers have failed. In the present work I interpret the

word ‘yuman’ on the lectotype locality label to be a

variant or erroneous spelling of ‘Yunnan’, in reference

to the Chinese province of Yunnan, where notabilis

has been confirmed to occur. The significance of the

word ‘mamun[?]’ is unknown. It could not be traced

in the available gazetteers.

OSWALD: Revision of Balmes

Other material examined (34). — CHINA (Main-

land): Yunnan Prov: 18, Kunming [25°04’N

102°41’E], 2.vi.1941 (USNM). LAOS: Xieng khouang

Id, Xieng Khouang [19°20’N 103°22’E],

22.iv.1919, Salvaza (BMNH). VIETNAM: Hoang lien

sone» MChapaz ESP D22ENF10350EN

Tonkin, v.-vi.1916, Salvaza (BMNH).

Etymology. — Unexplained, probably derived from

Latin, notabilis, noteworthy.

Comments. — Based on its distinctive male termi-

nalia, B. notabilis is here recognized as a valid species

and is resurrected from the synonymy of birmanus,

where it was placed by Kimmins (1939) and subse-

quent authors.

Balmes terissinus Navás

(figs. 3, 19-26)

Balmes terissinus Navás, 1910: 85 (OD, FW*). — Navás 1917

(RD); Oswald 1993 (Lst).

Diagnosis. — Distinguished from all other Balmes

species by the following combination of characters:

(1) forewing intraradial area traversed by two [not

three] gradate series (fig. 3), (2) apex of male 9th ster-

nite rounded [not conspicuously emarginate] (fig.

20), and (3) ventrolateral lobes of male 9th gonocox-

ites absent [not present] (fig. 25).

Description. — Forewing (fig. 3): length 10.8 - 15.7

mm (mean = 13.1 mm, n = 22 wings); markings: pat-

tern similar to birmanus with brown mottling on a

hyaline ground, but markings along ‘vena triplica’

and cubital and mediocubital spaces usually strongly

contrasting with adjacent membrane, hyaline-bound-

ed brown patches of distal margin (see birmanus de-

scription) obscure to absent; costal gradate series ab-

sent (1 or 2 adventitious costal crossveins occasionally

present); intraradial area traversed by two gradate se-

ries.

Male terminalia (figs. 19-25): 9th sternite: posteri-

or margin rounded, not sagittally emarginate; gonar-

cus: intragonarcus expressed as a narrow internal cos-

ta following course of antextragonarcal commissure;

extragonopons not prominently produced, midline

marked internally by a thickened costa which divides

posteriorly, distomedial margin of extragonopons

narrowly emarginate; extrahemigonarcus broad dor-

sally, tapering toward articulation with 9th gonocox-

ites; postextragonarcal commissure often poorly de-

fined; mediuncus: length 1 1/2 to 2 times width,

nearly parallel-sided proximally, attenuated distally;

apex bifid with a narrow but distinct separation

between the tines; 9th gonocoxites: superprocesses

united sagittally to form a low transverse elevation;

ventrolateral lobes absent; spiculate gonosaccal lobes:

two pairs present — an inconspicuous pair on dorsal

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

surface of bursa adjacent to ventral margin of fused

9th gonocoxites, and a second larger pair on ventral

surface.

Female terminalia (fig. 26): 7th sternite: parafoveal

lobes weakly developed, not prominently projecting;

8th sternite: reduced, transverse, expanded medially,

attenuate laterally, anterior margin not emarginate sa-

gittally; bursal accessory gland ducts: two free ducts

inserted bilaterally symmetrically on dorsal surface of

bursa at a pair of closely adjacent (less than two duct

diameters) points.

Distribution (fig. 1). — Mainland southeast Asia

(reported here only from Sichuan Province, China

[some imprecise records may also refer to far northern

Yunnan Province]).

Flight period. — April (no date) - 21 July.

Primary type. — Holotype d ? (MNHP?), not exam-

ined. Type locality: ‘Tibet, Ta-Tsien-lou’ [=CHINA:

Sichuan province Lucheng (=Tatsienlu, Tatsientu,

Kangting, and Kangding, 30°03’N 102°02’E)].

The presumptive holotype of terissinus, stated by

Navás to reside in the Paris Museum, was not avail-

able for examination. The forewing maculation pat-

terns of the material attributed here to this species

closely match the maculation pattern shown in

Naväs’ (1910:86, fig. 24) figure of the forewing of te-

rissinus. Particularly apparent are: (1) the strong con-

trast between the wing’s dark macula and its light

ground colour, and (2) the well-developed row of ma-

cula along the hind margin of the forewing. The iden-

tification of this material is also supported by distrib-

utional evidence. All of the specimens attributed here

to terissinus were collected in China in either Sichuan

province or, possibly for some specimens, far north-

Table 1. Coded character state data for Balmes formosus.

Characters (left numbers) and states (right numbers) are

identical to those given by Oswald (1993:28-40), except as

noted in the text under ‘Phylogenetic Analysis: Characters’.

Matrix symbols: 0, 1, 2, 4 character state numbers; ?, mis-

sing data (i.e., missing female data [characters 7 and 49] and

unknown male data [characters 10-39]).

1 1 16.= 8 DIE 46 1

2 0 AR DR 47 1

3 0 18 DONNE 48 1

4 0 ID 8 34 ? 49 ?

5 1 20 3 op) 8 50 1

6 1 Al ie 3) 18 51 1

7 à DD 8 SA DO)

8 1 28 DO: a (1

9 0 24 ? SO. SAN?

10 ? DD 8 40 1 55 1

11 ? Zom: 41 0 56 "I

12 2 ZI 8 42 1 0

13 2 DS 8 43 wl DS 0

14 ? DD) 44 4 > O IO

15 g SV 8 45 0 60 1

ern Yunnan province. The combined distribution of

this material and the type locality of terissinus (from

Sichuan province) form a compact range which is al-

lopatric relative to the ranges of other Balmes species.

Material examined (18d, 39, 52). — CHINA: Sz-

chuan {=Szechwan] province 1?, Suifu [=Yibin,

28°46N 104°34’E], 30.v.1925, 610m (2000,

Graham (MCZC); 176 ,4?, Yunnan border S of Suifu,

iv.[19]29, Graham (MCZC, USNM); 19, Chengtu

[=Chengdu, 30°40’N 104°04’E], 21.v.1929, Parish?

(cuIC); 1d, 19, Chengtu, v.1934, Graham (MCZC,

USNM); 19, Mt. Omei [=Emei Shan, 29°32’N

103°21’E], 21.vii.[19]35, 1215m (4000), Graham

(USNM).

Etymology. — Derived from the surname of J.

Terisse, collector of the holotype (see Naväs

1910:85).

PHYLOGENETIC ANALYSIS

Overview. — A cladistic analysis was used to esti-

mate relative interspecific relationships within the ge-

nus Balmes and the position of Balmes within the

family Psychopsidae.

Computational methods. — Cladograms were gen-

erated by application of the ‘ie’ (implicit enumera-

tion) tree calculation option of HENNIG86 (Farris

1988) to the input data file described under Data be-

low. The ‘ie’ tree calculation option guarantees iden-

tification of the minimal length tree(s) for a given in-

put matrix.

Characters. — The characters and character states

used in the present analysis are identical to those used

in the earlier analysis of Oswald (1993), except for the

addition of a fifth state to character 44, see below. See

Oswald (1993) for detailed definitions and discus-

sions of characters and their states.

Character 44. Female 8th sternite. (state 4 [new

state]) present, a small cordate plate, emarginate ante-

romedially. Oswald (1993) attributed four unordered

states to this character. Because the female 8th ster-

nite state found in formosus does not fit well into any

of those states, the preceding state is added here to ac-

commodate formosus into the present analysis. This

state is an autapomorphy of formosus.

Data. — The input data matrix contained numeri-

cally-coded morphological character data derived

from: (1) Oswald (1993: Appendix 2), for 21 previ-

ously character-scored psychopsid species and the hy-

pothetical ancestor used to root the tree, and (2) a

row of newly coded data (Table 1) for B. formosus.

Characters 10-39 in table 1 pertain to the unknown

male of formosus and were coded as unknown data

(‘?’). The following two additional formosus characters

were coded as unknown: [1] Shape of forewing hu-

meral plate (character 7) — the humeral plates of the

OSWALD: Revision of Balmes

Southeast Asia

Zygophlebiinae Psychopsinae

Silveira Balmes

Cabralis Psychopsis © £ 6 ¢

Zygophlebius È È N È

(Sold piedi E È

10,1-2

56,1-0

27,0-1

Lineage Symbol Key

24,0-1 the associated state change is:

unique within the Psychopsidae

unique, but later reversed,

within the Psychopsidae

of uncertain position on the

cladogram

parallelled elsewhere within the

Psychopsidae

a reversal parallelled elsewhere

within the Psychopsidae

NEO gps

Fig. 28. Cladogram showing relative interspecific relationships within Balmes, and the position of Balmes relative to the other

higher taxa of extant psychopsids. Character data are mapped only for Balmes and immediately adjacent lineages. Character

state changes are shown in the following format [left to right]: (1) number of occurrences on the cladogram, including those

on lineages not fully illustrated (see Phylogenetic Analysis Results), (2) lineage symbol, (3) character number, (4) state numbers

[ancestral state - derived state]. Male terminalic characters 21, 24, and 39 cannot be unambiguously mapped within Balmes

because the male of formosus is unknown.

available paratype of formosus are obscured, conse-

quently, this character could not be coded, and [2]

Distal apodemes of female 9th tergite (character 49) —

no distal apodemes could be located in formosus, how-

ever, because the membranous region where these ap-

odemes are located is poorly preserved in the available

paratype, the apparent absence of these structures

may be an artifact, particularly since they are fre-

quently difficult to observe even in well-preserved

material. In the analysis of Oswald (1993), distal ap-

odemes were shown to be a synapomorphy of the

Psychopsidae, and it is strongly suspected that they

will subsequently be found to be present in formosus.

To indicate the present unreliability of the coding of

this character in formosus, it has been coded as an un-

known datum in the present analysis.

Results. — A single most parsimonious tree (length

= 105, c.i. = 0.83, r.i. = 0.91) was found. This tree is

shown in fig. 28 in a reduced form that illustrates

character data only for lineages within Balmes and

those that immediately surround its attachment site

to the more general tree. The branching patterns and

character distributions of species within Psychopsis

and the Zygophlebiinae are identical to those present-

ed by Oswald (1993: 43, fig. 53). The tree shown in

fig. 28 places Balmes as the sister-group to the genus

Psychopsis and resolves relative interspecific relation-

ships within Balmes as follows: (formosus + (terissinus

+ (notabilis + birmanus))).

General discussion. — Oswald (1993) depicted the

southeast Asian species clade terissinus + notabilis [as

‘bnsl’] + birmanus as the monophyletic sister-group

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

of the Australian genus Psychopsis, and advocated tax-

onomic recognition of Balmes at generic rank.

Additionally, the Taiwanese species formosus (not ex-

amined for that analysis) was tentatively attributed to

Balmes. The finding here that formosus represents the

sister-group to the clade terissinus + notabilis + birma-

nus confirms the appropriateness of placing formosus

within Balmes.

The monophyly of Balmes is supported in the cur-

rent analysis by a single character: the fusion of the fe-

male 7th and 8th sternites (character 43; indepen-

dently derived in Psychopsis illidgei). However,

because the male of formosus is currently unknown,

this weak level of support may be misleading. Three

male terminalic characters (21, 24, 39) contain state

transformations that are synapomorphic for the three

non-formosus Balmes species; any or all of the derived

states of these transformations could be subsequently

identified in the male of formosus and lead to more ro-

bust support for the Balmes clade.

Character discussions. — [Character 6] Forewing,

crossveins of costal gradate series: (0) absent or few

[mean $ 5 crossveins/wing]; (1) numerous [mean > 5

crossveins/wing]. The previous analysis of Oswald

(1993) identified two equally parsimonious three-

step optimizations of this character, either (1) the

presence of a well-developed gradate series is plesio-

morphic within the Psychopsidae and secondarily re-

duced in Zygophlebius pseudosilveira Oswald [‘zns1° of

Oswald 1993] and Balmes, or (2) the absence of well-

developed gradate series is plesiomorphic within the

Psychopsidae and developed independently in

Psychopsis and the Zygophlebiinae, but subsequently

lost in the zygophlebiine species Zygophlebius pseudo-

silveira. The presence of a well-developed gradate se-

ries in Balmes formosus, and its inferred position here

as the sister-group to the three Balmes species treated

by Oswald (1993), now renders the first optimization

more parsimonious than the second. The current cla-

dogram unambiguously supports the first optimiza-

tion as the better interpretation of costal gradate series

evolution within the Psychopsidae, with the minor

modification that the loss of the costal gradate series

in Balmes is now seen as a synapomorphy of only the

three Indochinese Balmes species, rather than as a syn-

apomorphy of the entire genus.

[Character 39] Male gonosaccal membrane,

rounded spiculate lobes: (0) absent; (1) present. The

presence of a pair of spiculate lobes on the venter of

the eversible male gonosaccus is an interesting feature

of the male terminalia of the Indochinese species of

Balmes. A similar pair of lobes has been developed in-

dependently in Psychopsis insolens. These lobes prob-

ably function during copulation and insemination to

anchor the everted male gonosaccus within the female

bursa (Oswald 1993). Because the male of B. formosus

100

is unknown, it is currently uncertain whether the

presence of these lobes constitutes a synapomorphy of

Balmes in its entirety, or only of its mainland species.

A second, smaller, pair of lobes is present on the dor-

sal surface of the gonosaccus in B. terissinus. Because

terissinus is the sister-group of the one-pair-lobed

clade notabilis + birmanus, and the male of formosus is

unknown, it is presently impossible to infer the prop-

er cladogram positions of, and polarities among, the

three states (a) no lobes, (b) one pair of lobes, and (c)

two pairs of lobes. Discovery of the male of formosus

should resolve these questions.

[Character 56] Female bursa, number of inserted

bursal accessory gland ducts: (0) 1 unpaired duct; (1)

1 pair of ducts; (2) 2 pairs of ducts. One pair of bur-

sal accessory glands with ducts that insert at a pair a

widely separated, bilaterally symmetrical, points on

the dorsal/dorsolateral surface(s) of the female bursa

is plesiomorphic in the Psychopsidae (Oswald 1993;

state (1) above). This plesiomorphic condition under-

goes significant alteration in the Indochinese species

of Balmes. Balmes formosus exhibits the plesiomorphic

condition of two widely separated [>>2 times single

duct diameter] ducts. In B. terissinus two separate

ducts are present, but they are inserted very close to-

gether [<2 times single duct diameter]. In B. birma-

nus the ducts are partially fused distally, resulting in

only one, common, duct being inserted on the dor-

sum of the bursa. The intermediate state of terissinus

was not treated separately here, to maintain consis-

tency with the analysis of Oswald (1933); however,

were it to be considered as a separate state, the narro-

wing of the interduct insertion distance would map

on fig. 28 as another synapomorphy of the clade teris-

sinus + notabilis + birmanus. Since the female of nota-

bilis is unknown, it is uncertain whether the fusion of

the bursal accessory gland ducts should be considered

a synapomorphy of notabilis + birmanus, or simply an

autapomorphy of birmanus. The distal fusion of bur-

sal accessory gland ducts is an unusual condition,

which, to my knowledge, has not been reported in

any other neuropteran.

BIOGEOGRAPHY

The placement of formosus within Balmes is consis-

tent with the biogeographic hypothesis advanced by

Oswald (1993), which suggested that southeast Asian

psychopsids constituted a monophyletic group that

could be traced to a common ancestor that reached

Asia either by active dispersal from Australia or by

transport on an ancient rift fragment derived from the

northern margin of Gondwanaland in the vicinity of

present-day Australia. Additionally, the basal position

of formosus within Balmes correlates with the marked

disjunction between the known distributions of formo-

sus (Taiwan) and the mainland Balmes clade composed

of terissinus, notabilis, and birmanus (upland areas in

Burma, Laos, Vietnam, and south central China).

Although the precise significance of this disjunction is

unclear, its explanation is probably rooted in the his-

torical biogeography of southeastern Asia.

FUTURE RESEARCH

The genus Balmes is the most poorly known of the

five extant genera of the family Psychopsidae.

Distributional records for this genus are sparse and

additional collecting is needed to more fully docu-

ment the ranges of its species. It will be especially

interesting to see if the apparent allopatry of the

Indochinese species is corroborated by future collec-

tions. The adult female of notabilis and the adult male

of formosus, remain unknown. The discovery of the

male of formosus is particularly desirable.

The immature stages of Balmes species are com-

pletely unknown. Their discovery would be of con-

siderable interest because it would make possible

comparisons among the larvae of psychopsids from

southern Africa, Australia, and southeast Asia, the

three disjunct regions of the world in which living

silky lacewings are currently found.

Because the neuropterid fauna of southeast Asia is

poorly known, additional undescribed psychopsid

species may exist in this region. Any new living spe-

cies discovered in this region would be expected to fall

within, or as sister-groups to, the Balmes clade.

ACKNOWLEDGEMENTS

I thank the following curators, collection manag-

ers, and their staffs, for loaning specimens for this

study: Stephen Brooks and Peter C. Barnard (BMNH);

James K. Liebherr (CUIC); Sadao Takagi (EIHU);

David G. Furth (MCZC); Oliver S. Flint, Jr. (USNM).

The computer-generated map of southeast Asia was

produced by Dan Cole (USNM). Photographs were

produced by the Office of Photographic Services,

Smithsonian Institution. O. S. Flint, Jr. and G.

Miller kindly reviewed earlier drafts of the manu-

script. The research for this paper was supported in

part by two Smithsonian Postdoctoral Fellowships

and was undertaken while the author was in residence

OSWALD: Revision of Balmes

in the Department of Entomology at the National

Museum of Natural History (Smithsonian

Institution), Washington, DC, U. S. A.

REFERENCES

Farris, J. S, 1988. Hennig86 Reference. Version 1.5. —

Privately published.

Kimmins, D. E, 1939. A review of the genera of the

Psychopsidae (Neuroptera), with a description of a new

species. — Annals and Magazine of Natural History (11)4:

144-153.

Krüger, L, 1922. Psychopsidae. Beiträge zu einer

Monographie der Neuropteren-Familie der Psycho-

psiden. — Stettiner Entomologische Zeitung 83: 17-48.

Kuwayama, S, 1927. On a new species of Psychopsidae from

Formosa. — Insecta Matsumurana 1: 123-126.

Mansell, M. W, 1992. The systematic position of

Nemopteridae (Insecta: Neuroptera: Myrmeleontoidea).

— In: Canard, M., H. Aspöck & M. W. Mansell, eds.,

Current Research in Neuropterology. — Proceedings of

the Fourth International Symposium on Neuropterology,

pp. 233-241.

McLachlan, R, 1891. An Asiatic Psychopsis (Ps. birmana, n.

sp.). — Entomologist’s Monthly Magazine 27: 320-321.

Naväs, L, 1910. Hemeröbidos (Ins. Neur.) nuevos con la

clave de las tribus y géneros de la familia. — Broteria

(Zoologica) 9: 69-90.

Navas, L, 1912. Insectos neurépteros nuevos o poco conoci-

dos. — Memorias de la Real Academia de Ciencias y Artes

de Barcelona (3)10: 135-202.

Naväs, L, 1917. Ensayo monogräfico de la familia de los

Sicöpsidos (Ins. Neur.). — In: Asociación Española para el

Progreso de las Ciencias, Congreso de Valladolid (Sth,

held October 1915) 6: 181-210.

Navás, L, 1930. Névroptères et insectes voisins. Chine et

pays environnants. Première [I] série. — Notes d’Entomo-

logie Chinoise 1(6): 1-12.

New, T. R, 1989. The Psychopsidae (Insecta: Neuroptera)

of Australia and the Oriental Region. — Invertebrate

Taxonomy 2 (1988): 841-883.

Nichols, S. W, 1989. The Torre-Bueno glossary of entomol-

ogy. Revised Edition, compiled by Stephen W. Nichols.

— New York Entomological Society, New York, 840 pp.

Oswald, J. D, 1993. Phylogeny, taxonomy and biogeogra-

phy of extant silky lacewings (Insecta: Neuroptera:

Psychopsidae). — Memoirs of the American Entomolo-

gical Society 40: 1-65.

Received: 27 September 1994

Accepted: 8 November 1994

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Louis M. ROTH

Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, U.S.A.

REVISION OF THE COCKROACH GENUS

HOMOPTEROIDEA SHELFORD

(BLATTARIA, POLYPHAGIDAE)

Roth, L.M., 1995. Revision of the cockroach genus Homopteroidea Shelford (Blattaria,

Polyphagidae). — Tijdschrift voor Entomologie 138: 103-116, figs. 1-55. [ISSN 0040-7496].

Published 15 June 1995.

Eight species of Homopteroidea occur in Malaysia, Mentawai I., Philippines, Sabah, Sarawak,

Sulawesi, and Sumatra. Five species are redescribed, one is transferred from Ctenoneura, an-

other is resurrected from synonymy, and two new taxa are described. Diagnostic characters are

given for the genus and a key is presented to distinguish the adults.

Dr. L. M. Roth, 81 Brush Hill Road, P.O. Box 540, Sherborn, MA 01770, U.S.A.

Key words. — Homopteroidea; Blattaria, Polyphagidae; cockroaches; taxonomy; redescriptions,

new species.

Princis (1963: 104) listed four species of Homo-

pteroidea from Malaysia and Indonesia. In the present

paper eight taxa are considered to be valid: one species

is transferred from Ctenoneura, another is resurrected

from synonymy, five are redescribed, and two new

taxa are described.

The following museums and their curators or col-

lection managers kindly loaned me specimens: HECO

— Hope Entomological Collections, University of

Oxford, England; Dr. George C. McGavin & Mr. I.

Lansbury; MZZB — Museum Zoologi Bogoriense,

Bogor, Indonesia; MCZC — Museum of Comparative

Zoology, Harvard University, Cambridge, MA,

U.S.A.; RMNH — National Museum of Natural His-

tory (Rijksmuseum van Natuurlijke Historie), Lei-

den, The Netherlands; Mr. J. van Tol; ZILS — Zoo-

logical Institute, Lund, Sweden; Dr. Roy Danielsson.

SYSTEMATIC PART

Genus Homopteroidea Shelford

Homopteroidea Shelford, 1906: 274. — Type species:

Homopteroidea nigra Shelford, by monotypy.

Fulmekia Karny, 1926: 152. — Type species: Homopteroidea

nodipennis (Karny) = Fulmekia nodipennis Karny, by mo-

notypy.

Diagnosis. — Eyes reduced, wide apart, lateral, loca-

ted behind the antennal sockets (fig. 19). Tegmina

and wings usually fully developed extending beyond

end of abdomen, or tegmina reduced and hind wings

vestigial (one species, fig. 53); venation and pigmen-

tation of right and left tegmina usually differ in fully

developed winged forms, the major veins thickened

or raised, the region of the presutural vein (Hanitsch’s

terminology) branches of the right tegmen colourless

hyaline (fig. 3, right, a; except in aberrans). Hind

wing without an intercalary vein between the radius

and media, cubitus vein with two (fig. 34) or three

(fig. 11), branches whose distal regions usually are

connected by cross veins (fig. 4; except in aberrans).

Front femur Type C,, the piliform spinules are prac-

tically contiguous and appear as a dense fringe (fig.

22); pulvilli absent, tarsal claws symmetrical, distinct-

ly toothed on their proximal halves, arolia usually ab-

sent (fig. 5), or if present small and ‘fleshy’ whitish

(fig. 6). Male: Abdomen unspecialized; the supraanal

plate usually with a large white or yellowish medial

nonsetose macula (fig. 13; absent in male aberrans

and all females; I do not consider this a tergal gland).

I have examined a pair of Homopteroidea nodipennis

that were collected joined end to end in copula with

the distal half of the male’s supraanal plate bent ante-

riorly and the female’s terminalia were inserted and

held by the male’s phallomeres in his genital cham-

ber. Apparently the unsclerotized membranous zone

on the male’s supraanal plate is more flexible than the

heavily sclerotized areas and allows the plate to be

bent so that the female’s genital segments can be in-

serted and grasped properly; some pinned specimens

have the distal half of the supraanal plate bent up-

wards even though they are not in copula. Subgenital

plate symmetrical with a pair of similar widely spaced

styli (fig. 14); the anterior margin of the plate often is

straight and usually lacks lateral apodemes (fig. 30).

103

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Female subgenital plate incised longitudinally (fig.

50).

Distribution checklist by species of

Homopteroidea

(Records for specimens from localities which I ha-

ve not seen are taken from the literature and are

shown in brackets.)

aberrans Hanitsch: Mentawai; Sabah; Sumatra … 115

(BIA HAD SDs UB Sumatranen en 112

brachyptera.sp..n. Sumatra. init 114

maculata Hanitsch: Mentawai; Philippines; Sabah.

Havas Sumatra mer ee: 112

minor Hanitsch: Sabah; Sumatra ....... 114

nigra Shelford: [Java]; Malaysia; Mentawai; Sabah;

Sarawak; Sulawesi; [Sumatra] … 109

nodipennis Karny: Malay Peninsula; Sabah; Sarawak;

Sumatra ie 106

shelfordi Hanitsch: Sabah; Sarawak. [Borneo;

Malacca Sumatra) MN 104

Geographical distribution checklist of

Homopteroidea species

(Specimens from localities I have not seen are taken

from the literature and are shown in brackets.)

Borneo: shelfordi. — Java: [maculata; nigra). —

Malacca: [shelfordi]. — Malayasia: nigra. — Malay

Peninsula: nodipennis. — Mentawai L.: aberrans; macu-

lata; nigra. — Philippine L.: maculata. — Sabah: aber-

rans; maculata; minor; nigra; nodipennis; shelfordi. —

Sarawak: nigra; nodipennis; shelfordi. — Sulawesi: nig-

ra. — Sumatra: aberrans; biramiata; brachyptera; mi-

nor; nodipennis; (maculata; nigra; shelfordi]

Key to species of Homopteroidea

Tegmina and wings fully developed extending

beyondendiofabdomen men ann 2

— Tegmina reduced, not reaching end of abdomen,

hind wings vestigial (fig. 53) ............ brachyptera

2. Left and right tegmina similar in colour and scle-

rotization. Hind wing branches of the cubitus

vein not connected by cross veins (fig. 1E in Roth

Ideen alia ln. aberrans

— Right tegmen with a clear hyaline presutural zone

which is coloured in the left one (fig. 3). Hind

wing branches of the cubitus vein connected by

erossiyeinsültig 14) fn 3

3. Tegmina with a pale hyaline macula on their

proximallhalyesilhie! 28)... maculata

— Tegmina without pale maculae ....................... 4

4. Hind wing with two cubitus branches (fig. 34)

— Hind wing with three cubitus branches (fig. 4)

eed 6

5. Hyaline presutural zone with four or five oblique

branches (fig. 43). Arolia absent … … … … minor

— Hyaline presutural zone with more than five

branches (fig. 34). Arolia present (fig. 33)

EEDE SE AE O biramiata

6. Pronotum dark reddish brown with the lateral

border regions lighter (fig. 20). Antennae brow-

nish or black with as many as 15 white terminal

antennomeres. Male genitalia as in fig. 25 … … …

DONA PAR LE PR Er EE RE NINNI nigra

— Pronotum with the distal lateral border regions

whitish or pale (figs. 1, 7). Antennae without

whitesterminalsseementse re 7

7. Arolia absent (fig. 5). Male genitalia as in fig. 2

arri ER Ao IE a an shelfordi

— Arolia present (fig. 6). Male genitalia as in fig. 8

LARA der ARRE RR waa dc + CERN Rae he oe nodipennis

REDESCRIPTIONS, AND DESCRIPTIONS OF NEW

SPECIES

Homopteroidea shelfordi Hanitsch

(figs. 1-5)

Homopteroidea shelfordi Hanitsch 1925: 99, fig. 12 (4 & 9)

(in part, 26 from Mt. Murud only); 1929: 266, fig. 6;

1932a: 52; 1932b: 6; 1933a: 328; 1933b: 235; Bruijning

1948: 146; Princis 1963: 104.

Fulmekia nodipennis (nec Karny). Princis, 1963: 105.

Type material. — Lectotype d (terminalia slide

276) (here designated; Princis labelled this specimen

lectotype, 1963 but apparently never published the

designation), Mt. Murud, Sarawak, 6500 ft., Dr. E.

Mjôberg, Sarawak Museum dd 1925; Type Orth.

206'/, in HECO. Paralectotypes. Sarawak. HECO: same

data as lectotype 16 (fragmented), Type Orth.

206°/,. Additional material. — SABAH. HECO: ® para-

lectotype of Homopteroidea minor Hanitsch, Type

Orth. 389°/,, B.N. Borneo, Mt. Kinabalu, Kenokok,

3300 ft., 22.iv.1929.

Two other paralectotypes in HECO with the follo-

wing data are not shelfordi: one with the same data as

the lectotype, Type Orth. 206°/,, has on the back of

the type label ‘Homopteroidea hanitschi sp. n., K.

Princis, 1963’, but the description was never publis-

Figs. 1-11. Homopteroidea spp. — 1-4. H. shelfordi Hanitsch, male lectotype: 1, pronotum; 2, genitalia (dorsal); 3, left and

right tegmina (a = presutural vein in right tegmen); 4, left hind wing; 5, tarsal claws. — 6-11. H. nodipennis (Karny) males: 6,

tarsal claws; 7, pronotum; 8, genitalia (dorsal); 9, left tegmen; 10, right tegmen; 11, right wing. (6, 9-11, from Singgalang,

Sumatra; 7, 8, from Sipitang, Sabah).

104

ROTH: Revision of Homopteroidea

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

hed. It is a new species, namely biramiata, described

below. The other specimen (Type Orth. 206°/,) is

Homopteroidea nodipennis (Karny) a species which

Princis incorrectly synonymized with shelfordi.

Redescription. — Male: Head slightly exposed, eyes

reduced, lateral, widely separated, interocular space

greater than the distance between antennal sockets,

ocellar spots well developed, elliptical. Pronotum

suboval, hind margin straight (fig. 1). Tegmina and

wings fully developed, extending beyond end of ab-

domen, left tegmen with seven oblique branches in

the presutural zone, the right one with seven or eight

branches (fig. 3). Hind wing cubitus vein with three

branches distally connected by cross veins (fig. 4).

Front femur Type C,, pulvilli and arolia absent, tarsal

claws symmetrical, serrated on their proximal halves

(fig. 5). Abdominal terga unspecialized, supraanal

plate hind margin convexly rounded, right and left

paraprocts similar simple plates. Genitalia as in fig. 2.

Female (based on the paralectotype of

Homopteroidea minor Hanitsch): Left tegmen dama-

ged, right tegmen presutural vein with seven oblique

branches. Cubitus vein of hind wing with three cur-

ved branches distally connected by cross veins. Front

femur Type C,, pulvilli and arolia absent, tarsal claws

serrated. Supraanal plate trigonal, apex broadly, shal-

lowly concave.

Colour. — Head blackish, clypeus, labrum, and

mandibles pale, ocellar spots white, maxillary palpi

black. Pronotum dark brown with about two thirds

of the lateral borders pale, hyaline (fig. 1). Tegmina

reddish brown, presutural area of the right one clear

hyaline (fig. 3). Hind wing infuscated (fig. 4).

Adomen brown, male supraanal plate with a medial

pale macula.

Measurements (mm) (? in parentheses). — Length,

4.7 (4.0); pronotum length X width, 1.6 X 2.0 (1.3

X 1.7); tegmen length, 6.4 (4.7); interocular width,

0.9 (0.7).

Comments. — So far shelfordi is found only on Mt.

Murud [3°52N 115°30’E], and Mt. Kinabalu

[6°05°N 116°33’E]. Most of the other records

(Hanitsch’s and Bruijning’s) apparently are nodipen-

nis. The two species are very close and are distinguis-

hed by the presence or absence of arolia, and differen-

ces in their male genitalia.

The female specimen from Mt. Kinabalu is smaller

than the male and is similar in size to specimens of

Homopteroidea minor which probably led Hanitsch to

designate it a syntype of that species.

Homopteroidea nodipennis (Karny)

(figs. 6-11)

Fulmekia nodipennis Karny, 1926: 158, figs. 151-155.

106

Syntypes 1d, 29, ‘Maryland’ in Sumatra, OK.

Lichtfang, v.1925, leg. Dr. L. Fulmek, No. 2; in MZZB

[not examined].

Homopteoidea nodipennis (Karny). — Princis 1963: 105 (in-

correctly listed as a synonym of Homopteroidea shelfordi

Hanitsch).

Material examined. — SABAH. ZILS: The following were

collected by S. Adebratt: Malaysia, Sabah, Sipitang,

Mendolong, T5/R, 1 (abdomen missing), 11.v.1988; 16

(terminalia slide 426), 19, 28.iv.1988, 16, 2.11.1989,

T1B/W4, 19, 10.11.1988, T6/R, 19, 14.11.1989, T3/W5,

1d, 20.11.1989, 19, 24.11.1989. One specimen retained in

MCZC. RMNH: Malaysia, Sabah, Long Pa Sia airstrip along S.

Pa Sia, 115°43’E 4°25’N, 1090 m, 1%, 14.x.1986, J.

Huisman; Beaufort, 105 km S. of Long Pa Sia area, Sg.

Ritan, 115°42’E 4°24’N, 1160 m, undisturbed evergreen

tropical rainforest, 1, 7.iv.1987, J. van Tol & J. Huisman.

— SARAWAK. RMNH: Borneo, Sarawak, 16km N. of Bario,

Long Rapun, Sg. Dapur, ML, 115°35’E 3°53’N, 1200 m,

16, 19-20.11.1987, J. Huisman. — [Indonesian] BORNEO.

HECO: O. Borneo, Pajau R., 19, Mjöberg. — SUMATRA.

RMNH: The following were collected by E. Jacobson: Fort de

Kock (Sumatra), 920 m, 2d, 22, 1925; Air Njuruk

Dempu, 1400 m, 16, 49, viii.1916; Gun. Teleman, Sum.,

1d, 12, 1917; Sungai Kumbang, Sum., 1d (terminalia sli-

de 208), viii.1915. Two retained in MCZC. HECO: Gunung

Singgalang (Sumatra’s Westkust), 1800 m, d & © in copu-

la, 58, 12, 1925, E. Jacobson. MALAY PENINSULA. HECO:

paralectotype of Homopteroidea shelfordi Hanitsch (Type

Orth. 206°/,), Perak, Larut Hills, 4000-5000 ft, capt. ii.-

iii.1905, & pres. by R. Shelford.

Redescription. — Male: Head globose, exposed, ey-

es wide apart, lateral, behind the antennal sockets.

Pronotum subparabolic the dark portions slightly rai-

sed (fig. 7). Tegmina and wings fully developed ex-

tending beyond end of abdomen; right and left teg-

mina with six to eight oblique presutural branches

(fig. 9-10). Hind wing with three subcostal veins, cu-

bitus vein with three branches distally connected by

cross veins (fig. 11). Front femur Type C;; pulvilli ab-

sent, tarsal claws symmetrical, serrated on proximal

halves, very small arolia present (fig. 6). Abdominal

terga unspecialized; supraanal plate with a large hya-

line whitish median macula, hind margin convexly

rounded, right and left paraprocts similar simple pla-

tes. Subgenital plate symmetrical with small, similar,

cylindical widely separated styli, interstylar margin

rounded. Cerci with a long terminal spine. Genitalia

as in fig. 8.

Female: Left tegmen presutural zone with four to

six oblique branches, right one with seven (one bran-

ched) or eight. Supraanal plate hind margin trigonal,

apex rounded. Subgenital plate divided mediolongi-

tudinally, distal margin in lateral view truncate.

Colour. — Head dark reddish brown, ocellar spots

whitish, distal part of clypeus yellowish, palpi dark

reddish brown to blackish; proximal segments of an-

tennae brown, remainder blackish. Pronotum with

ROTH: Revision of Homopteroidea

0.25 mm

Figs. 12-18. Homopteroidea nigra Shelford, males from Sipitang, Sabah. — 12, pronotum; 13, supraanal plate (dorsal); 14, sub-

genital plate (ventral); 15, left hind wing; 16, left tegmen; 17, right tegmen; 18, tarsal claws.

107

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 19-26. Homopteroidea nigra Shelford, male from Perak. — 19, head; 20, pronotum; 21, subgenital plate (ventral); 22,

front femur (anterior surface); 23, supraanal plate and paraprocts (ventral); 24, styli and interstylar margin of subgenital pla-

te (dorsal); 25, genitalia (dorsal). 26, left and right tegmina, and left hind wing.

raised portions dark brown, about two thirds of the

distal lateral zones yellowish white (in some females,

white) (fig. 7). Tegmina brown, raised veins on dorsal

surface dull yellowish, on ventral surface white, pre-

108

sutural zone on left tegmen brown, on right tegmen

clear hyaline. Hind wing with subcostal and apical

areas darkened, some veins distally white. Abdomen

brown, supraanal plate with a clear hyaline macula.

| 0.25 mm

Cerci brown. Legs brown, tarsi lighter.

Measurements (mm) (2 in parentheses). — Length,

4.6-5.5 (4.5-5.3); pronotum length X width, 1.4-1.6

x 1.6-1.9 (1.3-1.6 X 1.6-1.8; tegmen length, 5.3-

6.5 (5.0-6.7); interocular width, 0.7 (0.7-0.8).

Comments. — This species is very similar to shelfor-

di and Princis incorrectly synonymized it with that

taxon. Karny’s description of nodipennis is very com-

plete and the specimens I have identified as this spe-

cies agree closely with his description. One of the cri-

tical characters he mentioned (p. 160) was the

presence of ‘verkiimmerten Haftläppchen’ (rudimen-

tary arolia), a structure absent in shelfordi.

Homopteroidea nigra Shelford

(figs. 12-26)

Homopteroidea nigra Shelford, 1906: 274, pl. 16, figs. 13, 14

(2); Hanitsch 1915: 127; 1923: 466; 1928: 37, 43;

1932a: 52, 80; 1932b: 6; 1933a: 303, 328; 1933b: 235;

ROTH: Revision of Homopteroidea

Figs. 27-30. Homopteroidea

maculata Hanitsch: 27, ma-

le pronotum; 28-29, female

left and right tegmina, and

left hind wing; 30, subgeni-

tal plate and genitalia (dor-

sal). (27, 30, from Sipora;

28, 29, from holotype).

Hebard 1929: 96; Bruijning 1948: 44, 146; Princis 1950:

162.

Type material. — Holotype 2, Kuching, N.W.

Borneo [SARAWAK], 9.x.1899, pres. 1905 by the

Sarawak Museum; Type Orth. 205 in HECO [the spe-

cimen is fragmented with parts mounted on cards]. —

Additional material: MENTAWE[A]I. HECO: Sipora,

1d, 19, 1 (abdomen missing), v.-vi.1894, E.

Modigliani, Mus. Civico Genova. MALAYSIA. HECO:

Perak, F.M.S., Batang Padang, Jor Camp, 1800 ft.

Id (wing slide 272, terminalia slide 273),

31.v.1923, H.M. Pendlebury. — SULAWESI (Celebes).

HECO: Macassar, 1d, 1896, Doherty, M. Burr

Collection, pres. 1903 by M.B. — SABAH. ZILS:

Malaysia, Sipitang, Mendolong, T2A/W4, 16, 1

(abdomen missing), 8.xii.1987, T1B/W4, 1d (termi-

nalia slide 425), 15.iv. 1988, 19, 1 (abdomen mis-

sing), 20.11.1988, 1 (abdomen missing), 8.111.1989,

T4/R, 18. 13.v.1988, T5/R, 25, 28.iv.1988, 16,

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 31-36. Homopteroidea biramiata sp. n.. — 31-34. female: 31, pronotum; 32, supraanal plate (dorsal); 33, tarsal claws and

arolium; 34, left and right tegmina, and right hind wing; 35, ootheca (lateral); 36, male genitalia (dorsal). (31-33, from ho-

lotype; 34, 35, paratype from Fort de Kock, Sumatra; 36, paralectotype of Homopteroidea shelfordi from Mt. Murud,

Sumatra).

3.v. 1988, 2 (abdomens missing), 11.v.1988. Two re-

tained in MCZC.

Description. — Male (previously undescribed):

Head globose, eyes reduced, widely separated, lateral,

interocular space greater than the distance between

antennal sockets, ocellar spots transverse (fig. 19);

fifth maxillary palpomere swollen, about as long as

the fourth, each shorter than the third. Pronotum

suboval (figs. 12, 20). Tegmina and wings fully deve-

loped extending beyond end of abdomen; right teg-

men with seven or eight oblique branches in the clear

hyaline presutural zone (figs. 17, 26, top). Hind wing

110

cubitus vein with three branches that are distally joi-

ned by cross veins (figs. 15, 26, bottom). Front femur

Type C,, with a dense row of almost contiguous pili-

form spinuli (fig. 22); pulvilli and arolia absent, tarsal

claws symmetrical basal half serrated (fig. 18).

Abdominal terga unspecialized, supraanal plate hind

margin convexly rounded, apex weakly indented (fig.

13), right and left paraprocts similar simple plates

(fig. 23). Subgenital plate with a pair of small, similar,

cylindrical widely separated styli, interstylar margin

convexly rounded (figs. 14, 24). Cerci with a large

terminal spine. Genitalia with complex phallomeres

as in fig. 25.

ROTH: Revision of Homopteroidea

Figs. 37-43. Homopteroidea minor Hanitsch, male from Sipitang, Sabah. — 37, pronotum; 38, supraanal plate (dorsal); 39, cer-

cus; 40, right hind wing; 41, subgenital plate (ventral); 42, 43. left and right tegmina.

Female: Similar to male except for the subgenital

plate which is deeply, longitudinally incised, the mar-

gins of the incision contiguous, the distal margin in

profile truncate (as in figs. 50, 51).

Colour. — Head shiny black, clypeus, labrum, and

mandibles light brown, ocellar spots yellowish white

or white (fig. 19); antennae dark brownish or black

with about 15 terminal white segments (Shelford de-

scribed the antenna as being fuscous; the fragmented

holotype lacks antennae and they probably lacked the

distal antennomeres when it was originally described;

several specimens from Sabah had some white anten-

nomeres and one which appeared to have undamaged

antennae had 15 white segments but in most speci-

mens segments are missing and these have fewer or no

white antennomeres.); maxillary and labial palpi

black. Pronotum with raised portion black or dark

brown, broad lateral zones lighter reddish brown

(figs. 12, 20). Tegmina dark reddish brown except for

the clear zone in the presutural area of the right wing

cover (figs. 16, 17, 26 top). Hind wing with costal

veins and apical region infuscated (figs. 15, 26 bot-

tom). Abdomen dark brown, supraanal plate with a

large clear hyaline area (figs. 13, 23; absent in female).

Legs with femora brownish, tibiae and tarsi pale.

Measurements (mm) (® in parentheses). — Length,

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

5.2-6.1 (6.3); pronotum length X width, 1.8-2.0 X

2.1-2.3 (1.9-2.0 X 2.3); tegmen length, 5.6-6.3 (5.5-

5.8); interocular width, 0.9 (0.9-1.0).

Homopteroidea maculata Hanitsch

(figs. 27-30)

Homopteroidea maculata Hanitsch, 1929: 266, figs. 7-9 (2);

1932a: 81; Bruijning 1948: 147; Princis 1963: 105.

Material examined. — Holotype, 9, Lubuksikaping (SU-

MATRA’s Westkust), 450 m, 1926, E, Jacobson; Type Orth.

363 in HECO. — Additional material: SABAH. ZILS: Malaysia,

Sabah, Sipitang, Mendolong, T5/R, 19, 28.iv.1988, 19

(abdomen missing), 11.v.1988, S. Adebratt. — PHILIPPINE IS-

LANDS. HECO: Mt. Makiling, Luzon, 19, Baker. — MENTA-

wE[A]I Islands. HECO: Sipora, 14 (terminalia slide 275), v.-

vi.1894, Modigliani, Mus. Civ. Genova.

Redescription. — Female: Head slightly exposed, ey-

es wide apart, reduced, lateral, interocular space greater

than the distance between antennal sockets. Pronotum

suboval, the midregion shallowly cucullate, hind mar-

gin straight. Tegmina and wings fully developed exten-

ding well beyond end of abdomen, left tegmen with

four or five branches of the presutural vein, the right

one with six or seven rami (fig. 28). Hind wing cubitus

vein with three branches connected distally by cross

veins (fig. 29). Front femur Type C,, pulvilli and arolia

absent, tarsal claws symmetrical, their proximal halves

serrated. Supraanal plate with hind margin convexly

rounded, entire. Subgenital plate divided medially,

hind margin in profile truncate.

Male (previously undescribed). — Pronotum (fig.

27), tegmina and wings as in female (fig. 28, 29).

Abdominal terga unspecialized. Supraanal plate with

a hyaline area medially, hind margin convexly roun-

ded, entire, right and left paraprocts similar plates.

Subgenital plate with hind margin rounded, styli si-

milar, small, widely separated, interstylar margin

weakly curved (fig. 30). Genitalia as in fig. 30.

Colour. — Head dark brown, labrum and lower half

of clypeus lighter; antennae dark brown; maxillary

palpomeres grayish. Pronotal disk with cucullate por-

tion dark brown, about two thirds of the lateral re-

gions lighter or whitish (fig. 27). Left tegmen brown

with a brownish yellow hyaline macula on anterior

half; right tegmen similarly coloured except that the

presutural vein area is colourless-hyaline (fig. 28).

Abdomen brown. Legs brown, tarsi lighter.

Measurements (mm). — Length, 4.5 (5.0-5.2); pro-

notum length X width, 1.5 X 1.6 (1.4-1.6 X 1.6-

2.0); tegmen length, 4.5 (4.0-5.6); interocular width,

0.8 (0.7-0.8).

Comments. — The species is readily recognized by

the hyaline mark on the proximal half of each teg-

men.

112

Homopteroidea biramiata sp. n.

(figs. 31-36)

Type material. — Holotype, 2, Fort de Kock (SU-

MATRA), 920 m, E. Jacobson (det. by Bruijning as

Homopteroidea shelfordi Hanitsch); in RMNH. [I selec-

ted a female as the holotype because the one male is in

very poor condition]. — Paratypes. SARAWAK. HECO:

Mt. Murud, Sarawak, 6500 ft., 16 (terminalia slide

277) paralectotype of Homopteroidea shelfordi

Hanitsch (Type Orth., 206°/,). — SUMATRA. HECO:

Fort de Kock (Sumatra), 920 m, 1% (carrying an

ootheca in the vertical position), 1925, E. Jacobson.

Description. — Male: Head hidden, eyes reduced,

wide apart, lateral, ocellar spots very small, interocu-

lar width greater than the distance between antennal

sockets. Pronotum suboval, hind margin weakly

curved. Tegmina and wings fully developed exten-

ding beyond end of abdomen, the left tegmen with

about seven oblique branches in the presutural zone,

a similar number in the clear presutural zone of the

right wing cover. Hind wing with two curved bran-

ches of the cubitus vein, distally joined by cross veins.

Front femur Type C,, pulvilli absent, tarsal claws

symmetrical, serrated, arolia present. Abdominal ter-

ga unspecialized. Supraanal plate with hind margin

convexly rounded, apex very shallowly indented,

right and left paraprocts similar plates. Subgenital

plate with a pair of widely separated styli, interstylar

margin convex. Genitalia as in fig. 36.

Female: Head slightly exposed, eyes reduced, later-

al, wide apart, interocular width greater than the dis-

tance between antennal sockets, ocellar spots absent.

Pronotum suboval, hind margin straight, (fig. 31).

Tegmina and wings fully developed extending be-

yond end of abdomen both with seven or eight obli-

que branches in the presutural zones (fig. 34 top &

middle). Hind wing with two branches of the cubitus

vein, distally connected by cross veins (fig. 34 bot-

tom). Front femur Type C,, pulvilli absent, tarsal

claws symmetrical, serrated on their proximal halves,

arolia present (fig. 33). Supraanal plate subtrigonal,

apex rounded (fig. 32). Subgenital plate divided lon-

gitudinally the distal margin truncate in profile. Cerci

with a terminal spine (fig. 32).

Colour. — Head reddish brown, clypeus, labrum,

and mandibles yellowish brown, maxillary palpi

blackish; antennae brown. Pronotum dark reddish

brown, the lateral proximal two thirds whitish (fig.

31). Tegmina brownish hyaline, the presutural zone

of the right one clear hyaline (fig. 34 middle). Hind

wing infuscated darkest in the costal and apical zones

(fig. 34 bottom; d wing darker than the 9).

Abdomen brown, male supraanal plate with a hyaline

macula. Cerci and legs brown, tarsi lighter.

ROTH: Revision of Homopteroidea

Figs. 44-52. Homopteroidea minor Hanitsch. 44-47, male lectotype. — 44, left and right (damaged) tegmina; 45, right hind

wing; 46, genitalia (dorsal); 47, pronotum. — 48-52, female from Sipitang, Sabah: 48, supraanal plate (dorsal); 49, left and

right tegmina; 50, subgenital plate (ventral); 51, subgenital plate (lateral); 52, pronotum.

Ootheca. — The ootheca carried by the female (fig. keel. The serrations contain respiratory tubes that al-

35) measures 1.3 mm high and 2 mm long. It is white low air to reach the eggs. There is a yellowish (yolk-

and has about 40 variably shaped serrations in the like) swelling internally that distends part of the later-

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

al walls of the egg case but there are no longitudinal

lines demarking the egg cells. The shape of the serra-

tions resemble those of the polyphagid Latindia sp.

but in that species the teeth are more uniform (Roth

1971: figs. 12, 13). In most cockroach oothecae that

have toothed or serrated keels, each serration leads to

a single egg but this is not true in Homopteroidea or

Latindia spp.; the latter has about 60 serrations but

there are usually only eight eggs (Roth 1971: 128).

Measurements (mm) (@ in parentheses). — Length,

? (4.7-5.4); pronotum length X width, 1.6 X 1.9

(1.5 X 1.8); tegmen length, ca 7 (5.7-5.8); interocu-

lar width, 0.8 (0.7-0.8).

Etymology. — The specific name refers to the two

branches of the cubitus vein of the hind wing.

Comments. — The hind wings of Homopteroidea

biramiata (fig. 34, bottom), and minor (fig. 40) have

two cubitus branches (other species have three), but

their tegmina differ in the number of oblique

branches in the clear presutural zones (seven in bira-

miata, fig. 34, middle; five in minor, fig. 43).

Homopteroidea minor Hanitsch

(figs. 37-52)

Homopteroidea minor Hanitsch, 1933a: 303, 328 (d lecto-

type only; 2 paralectotype = Homopteroidea shelfordi).

Material examined. — Lectotype (here designated), d

(terminalia slide 274), Mt. Kinabalu, B.N. Borneo [SABAH],

Kenokok, 3300 ft., 22.iv.1929; Type Orth. 389 /,, in HECO.

— Additional material: SABAH. ZILS: The following were col-

lected by S. Adebratt: Malaysia, Sipatang, Mendolong, 16,

T4/R, 19, T6/R, 14. i.1989, T3/W5, 16, 19,

17.11.1989, T5/R, 19, 1 (abdomen missing), 28.iv. 1988,

19, 1.v.1988. One retained in MCZC. — SUMATRA. HECO: Si-

Rambé (Sumatra's Westkust, 1d, xii.90-iii.91, E.

Modigliani, Mus. Civ. Genova.

Redescription. — Male: Head largely exposed, eyes

lateral, very far apart, interocular space greater than

the distance between the ocellar spots or antennal

sockets. Pronotum suboval (figs. 37, 47). Tegmina

and wings fully developed extending beyond end of

abdomen; left tegmen with three or four oblique

branches in the presutural zone (figs. 42, 44 top), the

right tegmen with four or five rami in that area (figs.

43, 44 bottom). Hind wings with about three costal

veins (only their bases distinct), median vein with a

branch that reaches the anterior margin, cubitus vein

with two branches distally joined by cross veins (figs.

40, 45). Front femur Type C;; pulvilli and arolia ab-

sent, tarsal claws symmetrical, distinctly toothed.

Abdominal terga unspecialized. Supraanal plate

ample, the distal region turned upwards its apex

weakly indented (fig. 38). Subgenital plate symmetri-

cal with a pair of small, similar, widely separated sty-

li, interstylar margin not produced (fig. 41). Cerci

114

with a long terminal spine (fig. 39). Complex genital

phallomeres as in fig. 46.

Female: Pronotum as in fig. 52. Tegmina (fig. 49)

and hind wing are similar to the male. The supraanal

plate lacks a medial hyaline spot (fig. 48) and the asty-

lar subgenital plate is longitudinally divided with its

distal margin truncate in lateral view (figs. 50, 51).

Colour. — Brown. Head dark brown, clypeus and la-

brum lighter, maxillary palpi grayish. Pronotum with

broad lateral regions whitish subhyaline (darker in lec-

totype), the remainder dark brown (figs. 37, 47, 52).

Left tegmen entirely dark brown or reddish brown

(figs. 42, 44 top, 49 top), right tegmen with the presu-

tural region clear hyaline, the remainder dark brown or

reddish brown (figs. 43, 44 bottom, 49 bottom). Hind

wing with the apical region, and the costal vein zone

infuscated (figs. 40, 45). Abdomen brown, supraanal

plate with a white or yellowish macula across the mid-

dle (fig. 38). Legs and cerci lighter brown.

Measurements (mm) (2 in parentheses). — Length,

3.8-4.5 (3.8-4.6); pronotum length X width, 1.3-1.4

x 1.6-1.7 (1.3-1.4 X 1.7); tegmen length, 3.5-4.2

(4.1-4.4); interocular width, 0.7 (0.7).

Comments. — Hanitsch’s ‘description’ of this spe-

cies was as follows: ‘The two specimens [d and 9]

quite agree in colouring and structure with A. nigra

Shelford, but measure only 5 mm. in total length, as

against 7 mm. of the type of nigra. In the absence of

more material I must leave it uncertain as to whether

or not they represent a new species.’ The differences

in male genitalia, tegmen and wing structure (fewer

cubitus branches in minor) and size, leave no doubt

that both taxa are valid.

Homopteroidea brachyptera sp. n.

(figs. 53, 54)

Type material. — Holotype, d (right tegmen on sli-

de 271, genitalia on slide 278), SUMATRA, Si-Rambé,

xii.1890-iii.1891, E. Modigliani; in HECO. — Para-

type. SUMATRA. HECO: Wai Lima, 19.

Description. — Male: Head globose, slightly expo-

sed, eyes reduced, wide apart, interocular space greater

than distance between antennal sockets. Pronotum

suboval, widest behind the middle (fig. 53). All but one

hind leg missing: pulvilli absent, tarsal claws symmetri-

cal, proximal half of ventral margin serrated, arolia ab-

sent. Tegmina reduced, widely separated reaching to

about the sixth segment, venation indistinct; hind

wing vestigial (fig. 53). Supranal plate hind margin

convexly rounded (fig. 53), right and left paraprocts si-

milar simple plates. Genitalia as in fig. 54.

Female: The specimen is in very poor condition

and mounted on a card. Tegmina are reduced and

hind wings are vestigial. Only one hind leg is present

ROTH: Revision of Homopteroidea

Figs. 53-55. Homopteroidea spp. — 53-54. Homopteroidea brachyptera sp. n., holotype male: 53, habitus (dotted line on the

right tegmen shows the outline of the underlying vestigial wing); 54, genitalia (dorsal). — 55. Homopteroide aberrans

(Hanitsch), male genitalia (dorsal), from Wai Lima, Sumatra.

and it lacks pulvilli and an arolium and the tarsal

claws are serrated. Supraanal plate trigonal, apex

rounded.

Colour. — Head reddish brown. Pronotum with rai-

sed portion dark brown, lateral regions pale (fig. 53).

Both tegmina similar, light brown hyaline, veins indis-

tinct, right tegmen without a clear hyaline presutural

zone (fig. 53). Abdominal terga and sterna brown, ma-

le supraanal plate with a medial clear zone (fig. 53).

Measurements (mm) ( in parentheses). — Length,

4.1; pronotum length X width, 1.5 X 1.8 (1.2

X 1.8); tegmen length, 1.9 (2.1).

Etymology. — The specific name refers to the redu-

ced tegmina and wings.

Comments. — This is the only species of

Homopteroidea whose tegmina are reduced and wings

are vestigial.

Homopteroidea aberrans (Hanitsch) comb. n.

(fig. 55)

Ctenoneura aberrans Hanitsch. — Roth 1993: 86, fig. 1 (re-

described ® and described d).

Material examined. — SABAH. ZILS: Malaysia, Sabah,

Sipitang, Mendolong, T5/R, 1d, 6.v.1988, T4/R. 16,

14.iii.1989, AIL, 14, 8.iv.1988, 1d, 25.iv.1988, ld,

26.iv.1988, 1d, 5.v. 1988, S. Adebratt. One specimen re-

tained in MCZC. — SUMATRA. HECO: Wai Lima, Z. Sumatra,

Lampongs, 24,29, xi.-xii.1921, Karny & Siebers; Sumatra

Binjei, Deli Hoa, Kapala Roessu, 19, viii.1922, Toxopeus.

MCZC: Wai Lima, Sumatra, 16 (terminalia slide 275).

Measurements (mm) ( in parentheses). — Length,

4.0-4.3 (4.7); pronotum length X width, 1.0-1.2 X

1.6-1.8 (1.2 X 1.6); tegmen length, 4.0-4.5 (4.4); in-

terocular width, 0.7 (0.8).

Comments. — This species was originally in

Ctenoneura, and I considered it to be atypical from

other members of that genus because its tarsal claws

are serrated and its subgenital plate is symmetrical

with two small similar styles. These are diagnostic

characters of Homopteroidea and I am therefore

transferring aberrans to that genus. However, it may

be considered ‘aberrant’ in this genus because the

presutural zone of the right tegmen is not clear hya-

line, the curved branches of the cubitus vein are not

connected by cross veins, and the male supraanal

plate lacks a hyaline macula (Roth 1993: figs. 1C,

115

“TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

E). The male genital phallomeres are shown in fig.

DÌ,

ACKNOWLEDGEMENTS

I thank the Australian Biological Resources Survey

(ABRS) for partial support, and the museums, cura-

tors, and collection managers indicated in the intro-

duction, who loaned me specimens.

REFERENCES

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glands, ovarioles, and tergal glands of Blattaria. — Annals

of the Entomological Society of America 64: 127-141.

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Received: 15 August 1994

Revised version accepted: 18 January 1995

Louis M. ROTH

Museum of Comparative Zoology, Harvard University, Cambridge, MA, U.S.A.

DESCRIPTION OF A NEW SPECIES OF CTENONEURA

HANITSCH FROM SABAH (BLATTARIA,

POLYPHAGIDAE)

Roth, L. M., 1995. Description of a new species of Ctenoneura from Sabah (Blattaria,

Polyphagidae). — Tijdschrift voor Entomologie 138: 117-119, figs. 1-8, table 1. [ISSN 0040-

7496]. Published 15 June 1995.

A new species of Ctenoneura Hanitsch from Sabah is described. New collection records are giv-

en for two known species.

Dr. L. M. Roth, 81 Brush Hill Road, P.O. Box 540, Sherborn, MA 01770, U.S.A.

Key words. — Ctenoneura; Blattaria, Polyphagidae; cockroaches; new species.

Princis (1963: 101; 1971: 1138) listed 15 species

of Ctenoneura. I (Roth 1993: 83) added 12 new taxa

in revising the genus and also provided a key to dis-

tinguish the adults; in a recent paper (Roth 1995), I

transferred Ctenoneura aberrans Hanitsch to

Homopteroidea. The genus is principally Malaysian,

Indonesian, and Asian.

While my 1993 revision was in press, Dr. Roy

Danielsson sent me a number of specimens of

Ctenoneura from the Zoological Institute, Lund,

Sweden (ZILS), of which at least one is new and is de-

scribed below. Mr. Willem Hogenes of the Zoolo-

gisch Museum, Universiteit van Amsterdam, The

Netherlands (ZMAN) sent me some material. New

records of two known species are also presented be-

low.

SYSTEMATIC PART

Genus Ctenoneura Hanitsch

Ctenoneura Hanitsch, 1925: 100. — Roth 1993: 83 (revi-

sion).

Rediagnosis. — Tegmina and wings fully developed

extending beyond end of abdomen. Right and left

tegmina generally similar in venation and sclerotiza-

tion, major veins not thickened, often densely reticu-

late with numerous cross veins, discoidal sector

oblique (fig. 7). Hind wing usually with an interca-

lary vein (between the radial and media veins), cubit-

us vein with three to eight branches (usually more

than three) that generally are parallel and reach, or al-

most reach the wing margin, and with numerous

small cross veins connecting them (fig. 8). Front fe-

mur Type C,, pulvilli absent, tarsal claws symmetri-

cal, simple, arolia present (small or subobsolete) or

absent. Male: Abdominal terga unspecialized.

Subgenital plate asymmetrical, usually with one sty-

lus, or none. Female: subgenital plate valvular.

Ctenoneura Hanitsch is very close to Homopteroi-

dea. The two genera can be separated by differences

shown in table 1.

Ctenoneura sipitanga sp. n.

(figs. 1-8)

Type material. — Holotype d, Malaysia, SABAH,

Sipitang, Mendolong, T1B/W4, 15.iv.1988, S. Ade-

bratt; in ZILS. — Paratypes. Sabah. ZILS: same locality

and collector as holotype, T6/R, 1d, 8.iii.1989,

T1B/W4, 14, 15.iv.1988 (retained in the Museum

of Comparative Zoology, Harvard University), T4/R,

1d, 3.v.1988.

Description. — Male: Head well exposed, eyes large,

bulbous, (fig. 1), interocular width slightly less than

the distance between antennal sockets, ocellar spots

very small. Pronotum suboval (fig. 1). Tegmina and

wings fully developed extending well beyond end of

the abdomen, the former with nonthickened veins

and oblique discoidal sectors (fig. 7). Hind wing with

an elongated subcostal vein that terminates, along

with about four costal veins, in a swelling; intercalary

vein present, simple, media vein bifurcate distad, cub-

itus vein with five curved, about equally spaced, par-

allel, simple branches (fig. 8). Front femur Type C;;

pulvilli absent from all legs, tarsal claws simple, sym-

metrical, arolia minute. Abdominal terga unspecial-

ized. Supraanal plate transverse, short, with midre-

gion of the hind margin convexly rounded, entire

117

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 1-8. Ctenoneura sipitanga sp. n., males from Sipitang, Sabah: 1. pronotum; 2. supraanal plate (dorsal); 3. cercus; 4. sub-

genital plate (end view); 5. subgenital plate (left lateral); 6. subgenital plate (right lateral); 7. left tegmen; 8. right hind wing.

(fig. 2). Subgenital plate asymmetrical, strongly con-

vex, hind margin on the left side curved dorsad and

forming an apically bifurcate spicular process; the

right side has three dissimilar processes, two of them

contiguous at the mid margin, and the third closer to

the cercus; styli absent (figs. 4-6). Cerci without a ter-

minal spine (fig. 3).

Colour. — Head reddish brown; antennae light

brown. Pronotum with a dark reddish brown macula

that extends from the anterior to the posterior mar-

gins, the wide lateral regions yellowish subopaque or

opaque (fig. 1). Right and left tegmina similar, their

humeral and proximal part of the costal vein zones

118

yellowish, the remainder reddish brown, hyaline (fig.

7). Hind wings with a yellowish tinge. Abdomen, cer-

ci, and legs light brown, subgenital plate yellow its

hind margin that forms the bifurcate process on the

left side very dark brown (figs. 4, 5).

Female: Unknown.

Measurements (mm). — Length, 5.0-6.5; prono-

tum length X width, 1.6-1.7 X 2.1-2.2; tegmen

length, 6.1-6.5; interocular width, 0.5.

Etymology. — The species is named after the local-

ity, Sipitang [5°05’N 115°33’E].

Comments. — Most of the males of Ctenoneura are

determined by differences in the shapes of their sub-

ROTH: À new species of Ctenoneura

Table 1. Generic differences between Ctenoneura and Homopteroidea

Character Ctenoneura

Tegmina Right tegmen without a

clear presutural zone.

Hind wing Intercalary vein present;

cubitus vein with 3 to 8 branches.

Tarsal claws symmetrical, simple.

Asymmetrical; with one

left stylus or none.

Subgenital

plate (3)

genital plates (key, #7 Roth, 1993: 84). The very dark

apically bifurcate process on the left hind margin of

the subgenital plate (figs. 4, 5) distinguishes sipitanga

from all other known Ctenoneura.

Ctenoneura scutica Roth

Ctenoneura scutica Roth, 1993: 102, fig. 16 (4).

Material examined.- SABAH. ZILS: Malaysia, Sabah,

Sipitang, Mendolong, P11, 1d, 10.iii.1989, S. Adebratt.

Comments. — The species is known only from Sabah.

Ctenoneura major Hanitsch

Ctenoneura major Hanitsch: Roth, 1993: 96, fig. I (9 & 2).

Material examined. — SABAH. ZMAN: Borneo,

Sabah, Kinabalu Nat. Park, sample Sab. 37, montane

forest, understory at light, headquarters area Kiau

View trail, 1560 m, 1d, 18.xi.1989, M.J. & J. P.

Duffels.

Comments. — This male is slightly smaller than the

holotype which is from Mt. Murud, Sarawak. Its

measurements (mm) are as follows: Length, 7.7; pro-

notum length X width, 2.1 X 2.8; tegmen length,

10.2; interocular width, 0.6.

Homopteroidea

Right tegmen with a

clear presutural zone.

Intercalary vein absent; cubitus vein with

2 or 3 branches.

symmetrical, serrated.

Symmetrical; with a

pair of small, similar styli.

ACKNOWLEDGEMENTS

I thank the Australian Biological Resources Survey

(ABRS) for partial support, and Dr. Roy Danielsson,

and Mr. Willem Hogenes who sent me the speci-

mens.

REFERENCES

Hanitsch, R., 1925. On a collection of Blattidae from

northern Sarawak, chiefly Mt. Murud and Mt. Dulit. —

Sarawak Museum Journal 3(1): 75-106.

Princis, K., 1963. Blattariae: Subordo Polyphagoidea, Fam.

Polyphagidae. /n Beier (ed): Orthopterorum Catalogus

4: 77- 172. ’s-Gravenhage.

Princis, K., 1971. Blattariae: Subordo Epilamproidea, Fam.

Ectobiidae. /n Beier (ed.): Orthopterorum Catalogus 14:

1039-1224. ’s-Gravenhage.

Roth, L. M., 1993. Revision of the cockroach genus

Ctenoneura Hanitsch (Blattaria, Polyphagidae). —

Tijdschrift voor Entomologie 136: 83-109.

Roth, L. M., 1995. Revision of the cockroach genus

Homopteroidea Shelford, with redescriptions, and de-

scriptions of two new species (Blattaria, Polyphagidae). —

Tijdschrift voor Entomologie 138: 103-116.

Received: 18 January 1995

Accepted: 29 March 1995

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Institute for Systematics and Population biology (Zoological Museum), Amsterdam

NEW AND RARE SPHECIDAE (HYMENOPTERA)

FROM WEST AFRICA

Simon Thomas, R. T., 1995. New and rare Sphecidae (Hymenoptera) from West Africa. —

Tijdschrift voor Entomologie 138: 121-130, figs. 1-14, tabs. 1-3. [ISSN 0040-7496]. Published

15 June 1995.

Eight new species of Sphecidae from Senegal, Cöte d’Ivoire, Nigeria, and Niger are described:

Liris (Leptolarra) rufula sp. n., L. (Leptolarra) senegalensis sp. n., Nitela (s.s.) miekae sp. n.,

Miscophus rufigaster sp. n., M. senegalensis sp. n., M. wieringi sp. n., M. eburneus sp. n., and M.

pseudochrysis sp. n. In addition a description is presented of the unknown female Miscophus sal-

litus Andrade.

Dr. R. T. Simon Thomas, Institute for Systematics and Population Biology (Zoological

Museum), University of Amsterdam, PO Box 4766, 1009 AT Amsterdam, The Netherlands.

Key words. — Hymenoptera, Sphecidae, West Africa, new species.

The Sphecidae were obtained during a research

program in northern Senegal on the effects of chemi-

cal locust and grasshopper control on other insects.

The area in which the research was executed was ho-

mogenous savannah with sparse trees and shrubs

(Everts 1990). Collecting was carried out on seven

plots (A-G), each with five malaise traps (1-5). The

plots, 2 x 2 and 2 x 3 km, were situated 25-35 km

south east of Richard Toll (fig. 1). The insects in the

malaise traps were collected once a week, five times

before and five times after spraying. Plot C, for refer-

ence, was not sprayed.

By counting the number of different insect species or

groups before and after spraying the effect of the insec-

ticides could be measured (Evers 1990). This paper de-

scribes new species of Sphecidae found in the material.

Through the courtesy of J. W. Evers and K. W. R.

Zwart (LUW) I also had the opportunity of studying

other Sphecidae from Côte d'Ivoire, Nigeria, and

Niger, which are described here too.

Abbreviations of depositories

ZMAN, Institute for Systematics and Population

Biology (Zoological Museum), University of

Amsterdam, The Netherlands ; LUW, Laboratory of

Entomology, University of Wageningen, The

Netherlands; RMNH, National Museum of Natural

History (formerly: Rijksmuseum van Natuurlijke

Historie), Leiden, The Netherlands.

Abbreviations and symbols

Ocellar measurements (after Andrade 1960): POL -

Distance between the posterior ocelli; OOL -Distance

between a posterior ocellus and the corresponding eye;

SOL -Distance between the anterior ocellus and a pos-

terior one; VOL -Distance between a posterior ocellus

and an imaginary line interconnecting the posterior

corners of the eyes (fig. 2). The microsculpture terms

used in this paper are after Eady (1968).

SYSTEMATIC PART

Liris Fabricius, 1804

This is a large cosmopolitan genus comprising over

260 species, for the greater part found in the tropics.

About 70 species are found in the Ethiopian Region.

Both described species can be assigned to the subge-

nus Leptolarra Cameron, 1900, by having a transverse

row of stout setae on the apex of the pygidial plate,

the surface of pygidium covered with appressed hairs,

and claws untoothed.

Liris (Leptolarra) rufula sp. n.

(fig. 3)

Type material. — Holotype female: Senegal, 25-35

km sud de Richard Toll, piege malaise El, 10.

viii.1989, leg. H. v. d. Valk c.s. (LUW). — Paratypes:

same locality and collector as holotype, piege malaise

I, 2 Dey IS Mi piegeMmalaise AS MOMIE

viii.1989 (LUW); piège malaise C2, 19 8.viii.1989,

piège malaise C3, 1d 20.ix.1989 (ZMAN); piège mal-

aise C3, 19 13.ix.1989 (RMNH).

Description

Female 17 mm. Black; shining apical part of cly-

peus ferruginous. Palpi yellow ferruginous. Mandible

121

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

15°30’W

MAURITANIA

Sam Sam è LY

Lac de

Guiers A

Preis Legend

>= Lake, river

—— Road

[A] Plots with

malaise traps

ferruginous with dark brown apex. Scape, pronotal

lobe, tegula, and propodeal spiracle ferruginous.

Pedicel and flagellomeres I-VIII blackish brown with

ferruginous apical ring. Gaster bright ferruginous.

Apical margin of clypeus faintly undulated.

Mandible monodont with broad, deep emargination

on externoventral margin. Clypeus with central lon-

gitudinal shining spot and shining rim along apex.

Clypeus two and a half times wider than long. All fla-

gellomeres three times as long as wide at apex.

Flagellomere X not flattened. Vertex dull with fine

points. Frons and genae with silvery pubescence. Eyes

with scattered erect hairs. Inner eye margins converge

Fig. 1. Map of Senegal, the plots

| in the research area are indicat-

ed.

16°30’N

SENEGAL

P) e Keur Mor Ibra:

° Mo Poudjié

to vertex (from 40 to 17). Midocellus round.

Pronotum with scarce silvery pubescence, sides of

pronotum faintly obliquely striated. Mesonotum,

scutellum, and metanotum shining with very fine

points. Propodeum wider at the base than long (53 :

43). Dorsum and posterior face of propodeum with

strong transversal striation. Propodeum with some

pubescence along lateral sides. Side of propodeum

with oblique striation.

Gaster with very short silvery pubescence. Pygidium

narrow, shining, with large points, points separated

from each other by twice their own diameter. Pygidial

plate a little more than twice as long as wide, ending

Table 1. Differences in colour and characteristics of the females of the species Liris (Leptolarra) rufula sp. n., L. (Leptolarra)

senegalensis sp. n. and L. (Leptolarra) croesus (F. Smith).

L. (L.) rufula sp. n.

Scape ferruginous

Antennae blackish brown

Flagellomere X round

Pronotal lobe brown, no pubescence

Mesonotum dull and finely punctulated

Lateral part of propodeum strigose with silvery

pubescence

Gaster bright ferruginous

Pygidial plate narrow (38 : 17)

Apex of pygidium 6 satae in one row

Body slender

122

L. (L.) senegalensis sp. n.

brown with silver pubescence

with goldish pubescence

strigose with silvery

pubescence

dark ferruginous

broad (37 : 26)

10 satae in 2 rows

slender

L. (L.) croesus

(F. Smith)

ferruginous black

partly black, partly ferruginous black

flat round

black, no pubescence

with goldish pubescence

punctulated without

pubescence, short ridges

along edges

black

broad (38 : 28)

6 satae in one row

robust

blunt with a row of five stout setae at apex (fig. 3), with

some short reclining hairs and a few upright thin hairs.

Wings hyaline with ferruginous veins. Apex of

forewing beyond the nervation softly fumigated.

Hindwing hyaline.

Legs including coxae ferruginous. Spines of tibiae

II and III black.

Claws untoothed; spines of tarses I dark brown and

pointed at top. Metatarse I with five spines, last spine

not longer than next tarsus. Largest spine of tibia III

0.6 length of metatarse III.

No difference in structure or colour of holotype

and paratypes was found; the sizes of the paratypes are

15 and 10 mm. The number of setae at apex of pyg-

idium varies from 5 to 3.

Male 8-9 mm. Colour and structure largely the

same as in holotype. On exterior side of flagellum a

low carina runs from flagellomeres III-X. No differ-

ence in structure or colour in second male.

Etymology. — The name of the species refers to its

remarkable reddish coloration of gaster and legs.

Comparative notes. — See under L. (Leptolarra) se-

negalensis sp. n.

Liris (Leptotarra) senegalensis sp. n.

(fig. 4)

Type material. — Holotype female, Senegal, 25-35

km sud de Richard Toll, piège malaise El, 2.ix.1989,

leg. H. v. d. Valk c.s. (LUW). — Paratypes: same local-

ity and collector as holotype; piège malaise C3, 19

28.viii.1989 (LUW); piège malaise Gl, 19

WOO SN Mpièse malaise G25) 19 el G:ix- 1989

(ZMAN); piège malaise G3, 19 23.ix.1989 (RMNH).

Description

Female 17 mm. Black; mandibles ferruginous with

apical 0.4 black. Palpi, clypeus, scape, pedicel, and

flagellomere I ferruginous. Flagellomere II ferrugi-

nous with black stripe externally. Flagellomeres III-X

internally ferruginous, externally blackish. Genae and

vertex with short silvery pubescence. Gaster dark fer-

ruginous, apical halves of tergites with very short sil-

very pubescence.

Apical margin of median lobe of clypeus shining

and broadly arched. Clypeus nearly two and a half

times as wide as long (48 : 19). Head dull with tiny

points and some depressed white hairs. Frons and ge-

nae with silvery pubescence covering the underlying

structure. Flagellomere I three times as long as wide at

apex. Flagellomeres III-IV two times as long as wide

at apex. Flagellomeres V-IX one and a half times as

long as wide at apex. Flagellomere X externally flat-

tened. Eyes with scattered erect hairs. Inner eye mar-

SIMON THOMAS: Sphecidae from West Africa

gins converge to vertex (from 43 to 16). Mandible

monodont with broad deep emargination on externo-

ventral margin.

Pronotum with scarce silvery pubescence, apical

half of pronotal lobe ferruginous with rounded top.

Thorax with light yellow pubescence, shining, with

tiny points. Propodeum dull with fine granulation.

Side of propodeum with oblique striation.

Sternites II-V with a few long apical setae.

Pygidium wide, 1.3 times as long as wide at the base

(fig. 4). Dorsal side of pygidium punctulated covered

with short depressed goldish hairs. Apical rim round-

ed with ten setae in two rows.

Forewing yellowish with yellow veins. Apical part

beyond first transverse submarginal vein grey with

brown veins. Hindwing yellowish with yellow veins.

All legs including coxae ferruginous. Claws un-

toothed, apical half black. Spines of tibiae II and III

black. Largest spine of tibia II as long as metatarse

III. Spines of frontleg ferruginous. Metatarse I with

five spatulated spines; apical spine nearly 0.4 longer

than next tarsus. All pecten of the tarsi longer than

next tarsus.

The paratypes do not differ in structure or colour.

Their sizes are between 14 and 15 mm. The apical se-

tae of the pygidium range from 7-12.

Male unknown.

Etymology. — The species is named after the coun-

try in which the material was found (Senegal).

Comparative notes. — Although Liris (Leptolarra)

rufula sp. n. and L. (Leptolarra) senegalensis sp. n. are

much alike. L. senegalensis resembles in structure and

colour L. (Leptolarra) croesus (F. Smith), the first spe-

cies being slender in general appearance and the sec-

ond species more robust. L. rufula differs from all the

other species of Liris (Leptolarra) in having a bright

ferruginous gaster and legs. The light coloration

could be an adaption to strong solar radiation and re-

flection from the sandy soil (Lomholdt 1985). The

differences of the three species are summarized in ta-

ble 1.

Nitela Latreille, 1809

Nitela is a cosmopolitian genus. The Ethiopian

Region has the largest share of the 43 species. The

subgenus Nitela is mostly found in the Old World

and Australia. The described species belong to the

subgenus Nitela Latreille in having bare eyes and

frontoclypeal carine absent or if present not lamelli-

form.

Nitela (s.s.) miekae sp. n.

Type material. — Holotype female, Senegal,

123

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 2-4, 6-14: 2, dorsal view of head of Myscophus, showing the ocellar measurements; 3, Liris rufula sp. n., female, pygidi-

um; 4, Liris senegalensis sp. n., female, pygidium; 6, Miscophus rufigaster sp. n., female, head, anterior view; 7, Miscophus se-

negalensis sp. n., female, head, anterior view; 8-9, Miscophus wieringi sp. n., female - 8, head, anterior view; 9, forewing; 10-

12, Miscophus eburneus sp. n. — 10, female, head, anterior view; 11, male, head anterior view; 12, sternite VII, ventral view;

13, Miscophus pseudochrysis sp. n., female forewing; 14, Miscophus sallitus Andrade, female, forewing.

Ziguinchor, 13.v.1983, mangrove, J. W. Everts

(ZMAN). — Paratypes 1 d 19, same data as in holo-

type (ZMAN).

Description

Female 4 mm. Black; tegula dark ferruginous. First

and basal half of second segment of labial palpi dark

brown, apical half of second segment and the next

two segments light brown. First two and basal half of

the third segment of the maxillary palpi dark brown,

apical half of third segment and the next three seg-

ments light brown. Mandibles basal third black, api-

cal two third ferruginous. Antennae black, flagello-

meres IIl-X ventrally with dense short bluish white

pubescence.

All segments of the palpi with some hairs at the ap-

ical end of the segments. Clypeus wider than high (17

124

: 7), bulging in the middle with narrow and high

midcarina. Clypeus and ventral half of frons with

dense silvery pubescence covering underlying struc-

ture. Dorsal part of frons and vertex finely rugose.

Interocular distance across base of clypeus nearly

twice as wide as across vertex. POL : OOL = 3.5, SOL

: OOL = 2.5, VOL : OOL = 2.0. Along interocular

rim of the compound eyes a line of white scales.

Pedicel length 0.6 of width. Flagellomeres I and II 1.6

times as long as wide. Flagellomeres III-VI 1.4 times

as long as wide. Flagellomeres VII-X 1.2 times as long

as wide. Flagellomeres X somewhat dorso-ventrally

flattened.

Dorsal side of pronotum separated from front side

by a transversal carina showing a triangle in the mid-

dle. Mesonotum transversally finely strigose, punctu-

lated in between. Mesonotum separated from scutel- |

Fig. 5. Distribution of the Miscophus bicolor species group in

the Ethiopian and west Eurasian regions (Lomholdt 1985)

and the species described in this paper. — % M. rufigaster

sp. n., @ M. senegalensis sp. n., À M. wieringi sp. n., I

M. eburneus sp. n., % M. pseudochrysis sp. n., WM. sal

litus Andrade, + M. funebris Honoré.

lum by a crenulated furrow. Scutellum and metano-

tum finely reticulated. Mesopleura strongly rugose

with a glabre, shining hypoepimeral area. Episternal

sulcus and hypersternaulus strongly developed.

Metapleura and sides of propodeum with longitudi-

nal strigose. Dorsal side of propodeum strongly retic-

ulate rugose, propodeal enclosure separated from ver-

tical sides by a carina. Ventral side of thorax with

scattered white hairs.

Tergites and sternites I-V black with apical ferrugi-

SIMON THOMAS: Sphecidae from West Africa

nous rim, tergite VI black without pygidium. Tergite

I nearly twice as wide as long on the dorsal surface.

Apex of sternites with some white hairs.

Wings hyaline with light brown veins and stigma.

All tibiae black with apical and basal brown rings.

All femurs black with apical brown rings. All tarsi

brown.

Male 4 mm. Colour and most of the structure as in

holotype. Apex of scutellum five longitudinal crenulate.

Etymology. — I am pleased to name this species af-

ter my wife Miek, who has helpfully assisted me in

the field and in preparing the manuscript.

Comparative notes. — Nitela (s.s.) miekae sp. n. is

the first species of this genus collected in West Africa,

resembling very much N. (s.s.) mochii Arnold (1940),

which was collected at Wadi Hoff, 60 km south of

Cairo, Egypt. The differences in characteristics of

both species are given in table 2.

Miscophus Jurine, 1807

(fig. 5)

Miscophus is represented by about 150 species in all

regions except Australia. Most species are found in

the Old World, especially the Mediterranean area.

Up till now no Miscophus species were found in West

Africa south of Spanish Sahara (20° N) (Lomholdt

1985). All species described below belong to the di

color species group. The bicolor species group holds

small species, rarely more than 6 mm long; genae

mostly without erect whitish pilosity; apex of fifth

sternite in the females without a complex of very shal-

low large rough puncturation (Andrade 1960).

Fig. 5 shows the known distribution of the bicolor

species group (after Lomholdt 1985) as well as the

collecting places of the new species in West Africa.

The Miscophus species are distinctive sanddwellers

(Honoré 1944). They are found along the desert, but

also around ‘wadis’. Mostly found in spring

(March-June) on the hottest part of the day on the

sand or around the stems of desert plants. Nest in the

soil. Spider catchers.

Table 2. Differences in characteristics of the species Nitela (s.s.) mochii Arnold and N. (s.s.) miekae sp. n.

Nitela (s.s.) mochii Arnold

Clypeus and lower part of face

Nitela (s.s.) miekae sp. n.

pubescence scarce and silvery

pubescence dense and silvery covering

Frons closely punctulated

Mesonotum punctulated

Proportinal length of pedicel and

flagello-meres I and II CORSIVO)

Tergite I

nearly three times as wide as long

underlying structure

finely rugose

transversal finely strigose

DATI

twice as wide as long

125

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Miscophus rufigaster sp. n.

(fig. 6)

Type material. — Holotype female, Côte d’Ivoire,

30-35 km N.Korghogo, mal. tr. 38, 10.11.1980, J.

W. Everts (ZMAN). — Paratypes 1 9, same data as hol-

otype (ZMAN); 1 ®, same place and collector, mal. tr.

25, 17.iv.1980 (LUW).

Additional specimens. — 19, Côte d'Ivoire, Katio-

la-Coton, 13.xi.1980, mal. tr., J. W. Everts (RMNH);

19, same place and collector, malaise, 20.iii.1981

(ZMAN).

Description

Female 7 mm. Black; mandibles ferruginous with

red brown apex after a black ring. Apex of subbasal

notch of mandible red brown. Scape, pedicel, and

gaster light ferruginous. Palpi dark greyish.

All segments of palpi with outstanding hairs. Inner

margins of compound eyes mostly parallel, at vertex

somewhat converging and near clypeus diverging (fig.

6). Median lobe of clypeus bulged, convex with

smooth and shining rim, rest of clypeus with long sil-

very hairs and punctulated. Frons with scarce silvery

pubescence. Apex of scape ending in sloping top; the

0.6 distal internal part of the scape with a shining

translucent area (fig. 6). Pedicel somewhat wider than

long. Flagellomeres I-III three times as long as wide at

apex, flagellomeres IV-X twice as long as wide at base.

Vertex finely and closely punctulated, bulging

between the ocelli. POL : OOL = 1.5, SOL : OOL =

LOS MOL KOOL SS

Pronotum, mesonotum, scutellum, and metano-

tum finely and closely punctulated. Metanotum with

lateral oblique silver pubescens, directed outwards.

Dorsal part of propodeum somewhat longer than

wide (15 : 20) wich minute, oblique strigose and clear

longitudinal midcarina. Posterior face of propodeum

rather abruptly cut‚ forming an angle of about 100

degrees with its dorsum, horizontal striated.

Sixth tergite sharply pointed apically, without pyg-

idium. Sternites II-VI each with two apical outstand-

ing hairs.

Forewing hyaline, somewhat clouded beyond the

nervation. First recurrent vein received by the first

submarginal cell, close to submarginal II, second re-

current vein received by submarginal cell II near the

apical end. Hindwing hyaline, media diverging be-

yond cu-a. Jugal lobe round and small with a very

deep incision.

Legs ferruginous; external side of the femora some-

what darker; coxae black. Tarsi apically brown.

Metatarsus I with two lateral spines. The apical spines

of metatarsus and first tarsus twice as long as the next

tarsus. The apical spine of second tarsus one third

longer than the third tarsus.

Male unknown.

Etymology. — From Latin rufo, meaning red, and

gaster, referring to all segments behind the propo-

deum.

Variation. — One female, Côte d’Ivoire, Katiola-

Coton, malaise, 20.11.1981, leg. J. W. Everts, differs

somewhat from the type in having the last three tergi-

tes and sternites dark brown and the forewing tip

white.

Comparative notes. — Miscophus rufigaster sp. n.

and M. funebris Honoré (1944) are much alike. The

differences in colour and characteristics of both spe-

cies are given in table 3.

Miscophus senegalensis sp. n.

(fig. 7)

Type material. — Holotype female, Senegal,

Ziguinchor, 13.v.1983, malaise, mangrove, J. W.

Everts (ZMAN).

Description

Female 4.5 mm. Black; palpi ferruginous; mandi-

bles dark ferruginous, black ring before amber red ap-

Table 3. Differences in colour and characteristics of the females of the species Miscophus rufigaster sp. n. and M. funebris

Honoré.

M. rufigaster sp. n. M. funebris Honoré

Legs excluding coxae ferruginous black

Last three gastral segments light ferruginous dark brown

Clypeus mostly shining without pubescence silvery pubescence

Pedicel somewhat wider than long somewhat longer than wide

Flagellomeres I-III 3 times as long as wide 2.1 times as long as wide

SOL : OOL 1.9 2.0

VOL : OOL 13 1.0

Dorsal part of propodeum obliquely strigose closely punctulated

Forewing hyaline, clouded beyond nervation basal part somewhat clouded; hyaline over

126

discoidal and submarginal cell, clouded

beyond nervation with white wing tip

ex, black base; apex of median lobe of clypeus brown;

pedicel and first three flagellomeres light ferruginous.

Scape ventrally ferruginous and dorsally dark brown.

Palpi wich scarce short pubescence. Clypeus and

lower part of frons with scarce white pubescence di-

rected apically. Medium lobe of clypeus smooth and

slightly bulging with a few large points. Apical rim of

medium lobe of clypeus smooth, no punctation, sep-

arated from basal part by a crenulated furrow (fig. 7).

Head with close and very fine punctulation. Inner

margins of compound eyes curved, near clypeus and

at vertex converging (17 : 20 : 13). Pedicel 0.9 of the

length of flagellomere I and as long as flagellomere II.

Flagellomeres II-VIII twice as long as wide.

Flagellomeres IX and X somewhat shorter.

Flagellomere X conical. POL : OOL = 3.0, SOL :

OOL = 2.0, VOL : OOL = 0.4.

Pronotal collar rather developed, transition to the

anterior declinity relatively sharp and well defined.

Mesonotum, scutellum, and metanotum very closely

and finely punctulated. Propodeum equally long and

wide at the base, very finely and obliquely strigose

and with longitudinal midcarina. Propodeum with

short silvery forwardly depressed pubescence. Side of

propodeum obliquely strigose. Mesopleuron and

metapleuron shining and finely punctulated. Post-

erior face of propodeum rather abruptly cut, forming

an angle of about 100 degrees with its dorsum.

Apical rim of tergites dark brown. Tergites and

sternites very finely punctulated. Tergite VI rounded

apically without pygidium.

Wings hyaline. Veins along front edge of forewing

dark brown. Other veins light ferruginous. 1m-cu en-

tering submarginal cell I at a distance from submargi-

nal cell II equal to the length of M2. Cul and 2m-cu

not reduced. Jugal lobe round and small equally long

and wide with very deep incision.

Inner side of tibia I, whole tibia II, and tarsi brown.

Other parts of the legs black. Hind tibia with two ap-

ical spines, one spine as long as 0.7 of metatarse III.

Male unknown.

Etymology. — The species is named after the coun-

try in which the type was collected (Senegal).

Comparative notes. — M. senegalensis belongs to the

aenigma species subgroup within the bicolor species

group. The species resembles M. nevesi Andrade,

1952 from Portugal. The differences are found in the

size of the body and the structure of the head. The

clypeus of M. senegalensis is more pointed; the dorsal

view of the head shows a much wider size behind the

eyes; POL : OOL = 3.0 and SOL: OOL = 2.0 in M.

senegalensis and POL : OOL = 1.8 and SOL : OOL =

1.4 in M. nevesi . The inner corners of the compound

eyes at the vertex are nearly right-angled in M. seneg-

SIMON THOMAS: Sphecidae from West Africa

alensis and rounded in M. nevesi . The ocelli are

placed more posteriorly, VOL : OOL = 0.4 in M. se-

negalensis and 1.0 in M. nevesi. Posterior face of pro-

podeum has a different cut, forming an angle with its

dorsum of about 130° in M. nevesi and 100° in M. se-

negalensis .

Miscophus wieringi sp. n.

(figs. 8, 9)

Type material. — Holotype female, Senegal, 25-35

km sud de Richard Toll, piege malaise D3, 31.

viii.1989, leg. H. v. d. Valk (LUW). — Paratype 19,

same place and collector as holotype, piège malaise

C3, 13.ix.1989 (ZMAN).

Description

Female 7 mm. Head black; palpi creamy white,

mandibles ferruginous, apex red, apex of subbasal

notch red. Scape, pedicel, and clypeus light ferrugi-

nous; flagellomeres I and II ventrally ferruginous dor-

sally brown, other flagellomeres ventrally brown and

dorsally dark brown. Pronotum, pronotal lobe, and

tegula ferruginous. Gaster light ferruginous.

External side of palpi with erect hairs. Clypeus

arched with rounded middle part. Clypeus narrow

(19 : 5) with broad translucent ferruginous rim with

some long hairs (fig. 8). Apex of scape ending in slop-

ing top (compare with fig. 6). Distance between com-

pound eyes at vertex half the distance just above cly-

peus POLAOOE=52 SO OOL = 4.0, VOL :

OOL = 2.5. Frons and clypeus with short and scarce

goldish pubescence covering the fine punctulation.

Flagellomere I two and a half times as long as wide.

Flagellomeres II and III twice as long as wide; flagel-

lomeres IV-X somewhat shorter. Flagellomere X con-

ical.

Pronotum rather long, collar half as long as wide,

lateral rounded. Collar finely and longitudinally stri-

gose. Mesonotum, scutellum, and metanotum with

short goldish pubescence and very closely punctulat-

ed. Mesopleuron strigose. Propodeum without pu-

bescence, shining, equally long and wide at the base,

lateral front part obliquely striated, middle apical part

very finely almost transversally striated, with rather

high midcarina. Angle between dorsum of propode-

um and hindslope about 100 degrees. Upper part of

mesopleuron with some goldish pubescence. Lower

part of the mesopleuron and mesosternum with sil-

very pubescence.

Gaster shining, with few small points. Tergites II-

IV translucent. Apex of tergite VI sharp, without pyg-

idium.

Forewing (fig. 9) hyaline with two clouded bands,

one just in front of submarginal cell I and the other

just over the marginal cell and submarginal cell II.

127

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Zone between the two clouded bands milky white

with white veins. Other veins light brown. Forewing

with petiole of submarginal cell II as long as height of

this cell. Cul and 2m-cu reduced. Jugal lobe round

and very small with deep incision.

All legs including coxa light ferruginous with

brown spines. Coxa I ventrally translucent and equal-

ly long and wide at the base.

Male unknown.

Etymology. — I am very pleased to name this spe-

cies after my friend H. Wiering, for his great help in

preparing this manuscript.

Comparative notes. — M. wieringi belongs to the

bicolor species group and resembles M. heliophilus

Pulawski, 1968 closely. Mesonotum, scutellum, and

metanotum are punctulated in M. wieringi and

strongly shagreen in M. heliophilus. The mesopleuron

is closely punctulated in M. wieringi and ridged lon-

gitudinally in M. heliophilus. There is a difference in

the place of the ocelli: POL : OOL = 3.2 and SOL:

OOL = 4.0 in M.wieringi and POL: OOL = 2.4 and

SOL : OOL = 3.0 in M.heliophilus.

Miscophus eburneus sp. n.

(figs. 10-12)

Type material. — Holotype female: Côte d'Ivoire,

30-35 km N.Korghogo, mal.tr. 39, 28.iv.1980, J. W.

Everts (ZMAN). — Paratypes 19, same data as holo-

type; 1d, Côte d'Ivoire, 30-35 km N.Korghogo,

mal.tr. 11, 3.1.1980, J. W. Everts (LUW); 29, same

data as holotype; 16, Côte d'Ivoire, 30-35 km

N.Korghogo, mal.tr. 40, 28.1v.1980, J. W. Everts

(ZMAN); 12, Côte d'Ivoire, 30-35 km N.Korghogo,

malitr 355) 17979) je We Everts; n lidi (Cote

dIvoire, 30-35 km N.Korghogo, mal.tr. 40,

28.iv.1980, J. W. Everts (RMNH).

Additional specimens. — 12, Côte d'Ivoire, Ka-

tiola-Savanne, malaise, 20.11.1981, leg. J. W. Everts

c.s., 12, Senegal, 25-35 km sud de Richard Toll,

piège malaise D3, 27.ix.1989, leg. H. v. d. Valk, 19,

Niger, Niamey, Centre Aghrymet, piége malaise-mil,

26.viii-1.1x.1988, Y. Jongema, 1 Q, Nigeria, savanne-

Côte d’Ivoire, 30-35 km N.Korghogo, mal.tr. 34,

28.1v.1980, J. W. Everts (LUW); 12, Côte d'Ivoire,

30-35 km N.Korghogo, mal.tr. 2, 28.11.1980, J. W.

Everts, 19, Côte d'Ivoire, Katiola, malaise,

20.iv.1980, leg. J. W. Everts, 1d, Ivory Coast, Kou-

dougou 10 km SE Bouaflé, malaise-trap, 16.11.1981,

J. W. Everts, 12, Senegal, 25-35 km sud de Richard

Toll, piège malaise G4, 21.viii.1989 H. v. d. Valk,

19, Senegal, 25-35 km sud de Richard Toll, piège

malaise G4, 30.ix.1989, H. v. d. Valk, 19, Senegal,

128

Vélingara, malaise, 15.v.1989 , J. W. Everts, 19,

Senegal, Ziguinchor, 13.v.1983, J. W. Everts (ZMAN).

Description

Female 5.5 mm. Black; palpi light ferruginous;

mandibles ferruginous, apex of mandible and apex of

subbasal notch red brown. Scape, pedicel, pronotal

lobe, tegula, and apical half of clypeus ferruginous.

Legs including coxae ferruginous; spines of all legs

dark brown. Sternites I and II, tergites I and II, and

lateral side of tergite III ferruginous.

Palpi externally hairy. Clypeus rather low (19 : 5)

with shining translucent apical rim (fig. 10). Clypeus

and frons just above antennal sockets with scarce sil-

very pubescence. Head very finely punctulated.

Frontal line shining. Inner margins of compound

eyes from clypeus up to halfway frons parallel, upper

half converging to apex (fig. 10). POL : OOL = 2.0,

SOL: OOL = 4.0, VOL: OOL = 2.5. Pedicel some-

what longer than wide, flagellomeres I and II four

times as long as wide, flagellomeres III and IV twice

as long as wide, flagellomeres V-X one and a half

times as long as wide.

Pronotum, mesonotum, scutellum, and metano-

tum finely punctulated. Propodeum with transversal

strigose and midcarina. Mesopleuron strigose. Propo-

deum equally long and wide at base (16 : 15). Side of

propodeum obliquely strigose. Mesopleuron finely

and horizontally strigose.

Sternites and tergites shining, very finely punctu-

lated. Tergite VI apically punctulated, without pygid-

ium. Sternite III-VI apically some long hairs.

Forewing somewhat clouded with hyaline band

over submarginal cell I and discoidal cell II. Veins

light brown and in hyaline band nearly hyaline. First

recurrent vein received by submarginal cell I at 0.3

from submarginal cell II. Cul and 2m-cu strongly re-

duced. Hindwing hyaline. In hindwing only medial

and submedial cell present. Jugal lobe small with a

deep jugal excision.

Coxa I and apical half of coxa II and III ferrugi-

nous, basal part of coxa II and III black. Femurs and

tibiae I-III for the greater part ferruginous, femurs I

and II with an apical, external black spot, tibia III ex-

ternally black; all tarsi dark brown. All coxae triangle

formed, 1.3 as long as wide at the base.

Male 5 mm. Colour largely as in holotype. Clypeus

trilobed, with narrow apical rim ferruginous (fig. 11).

Scarce silvery pubescence on basal part of clypeus and

on frons just above antennal sockets. Tarsi of all legs

dark brown. Sternite II with two lateral small round

dark brown spots. Tergite VII rounded apically.

Sternites III -VII apically two or three outstanding

long hairs. Apex of sternite VII straight with a line of

short hairs (fig. 12).

Etymology. — From Latin eburneus, meaning

made from ivory, so named in reference to the coun-

try in which the material was collected (Ivory Coast).

Variations. — The colour of the pronotum varies in

different areas. Some females from Ivory Coast have a

ferruginous spot at the lateral side of the pronotum.

The females of northern Senegal have a complete fer-

ruginous pronotum; all legs including the coxae are

completely ferruginous. The females of southern

Senegal, Niger, and Nigeria have a black pronotum as

in the holotype. In one male from Ivory Coast the

dark spots on the second sternite are missing.

Comparative notes. — The new species M. eburneus

sp. n. is very near to M. mineticus Honoré. The ante-

rior edge of the clypeus of M. eburneus without dis-

continuites is even more straight than the clypeus of

M. mineticus. The legs of M. eburneus are mostly fer-

ruginous and black in M. mineticus. M. eburneus has

no cupreous tinge. Propodeum of M. mineticus is

obliquely ridged with hairs directed forwards, while

M. eburneus has a transversally ridged propodeum

without hairs.

Miscophus pseudochrysis sp. n.

(fig. 13)

Type material. — Holotype female, Senegal, 25-35

km sud de Richard Toll, piege malaise C3,

13.ix.1989, H. v. d. Valk (LUW). — Paratype 14,

27.ix.1989, all other data as in holotype (ZMAN).

Description

Female 5 mm. Shining green; mandibles red

brown with dark green base; palpi brownish; scape

dark brown; pedicel and flagellum black. Tergites II

and III green with cupreous tint. Clypeus with dark

brown apical rim.

Ventral side of the segments of the palpi with two

rows of curved hairs. Clypeus almost straight at apex,

rounded towards compound eye, flat between antennal

sockets. Clypeus low (20 : 5). Lower half of frons very

roughly rugose. Upper half of frons with few large

points on punctulated area; vertex very finely and

closely punctulated. POL : OOL = 3.5, SOL : OOL =

3.5, VOL : OOL = 2.0. Pedicel a little longer than

wide at apex (5 : 4). Flagellomeres I - IV three times as

long as wide at apex. Flagellomeres V-VIII twice as

long as wide at apex. Flagellomeres IX and X a little

longer than wide. Flagellomere X flattened externally.

Pronotum, mesonotum, and scutellum very finely

punctulated, the points arranged in waves, only vis-

ible when strongly magnified. Sides of pronotum hor-

izontally strigose, pronotal lobe shining with some re-

clining white hairs. Dorsal part of propodeum, with

strong midcarina, finely strigose obliquely in front

and almost transversally apically. Propodeum at base

SIMON THOMAS: Sphecidae from West Africa

wider than long (16 : 12). Sides of propodeum

strongly and horizontally strigose. Mesopleuron retic-

ulated. Episternum of mesonotum with short hori-

zontal lines starting at episternal sulcus, rest of epis-

ternum punctated.

Tergites very finely punctulated. Sixth tergite apex

sharp and brown, no pygidium. Sternite V with two

outstanding hairs at apex; sternite VI with some out-

standing hairs at apical half.

Wings hyaline with light ferruginous veins. Veins

Cul and 2m-cu in forewing strongly reduced (fig.

13). Height of second submarginal cell 0.75 of base of

the cell.

All tarsi dark brown. Coxa I almost equilaterally

triangular and bulging. Metatarse I with two apical

spines and one lateral spine in the middle.

Male 4.5 mm. Colour and most characteristics as

in holotype. Base of mandible including basal notch

black, middle part brown, apex black. Scape with

brown ventral spot. Frons with two horizontal carinas

just above antennal sockets. Sternites II-VI with a few

setae along apex.

Etymology. — The species takes its name from its

great resemblance to M. chrysis Kohl, 1894.

Comparative notes. — M. pseudochrysis resembles

very much M. chrysis Kohl, 1894, of which only one

female was collected; a comparison can only be given

for the females. Reticulated part of the frons up to the

midocellus in M. chrysis and only half way in M. pseu-

dochrysis. Episternum of the mesothorax in M. pseu-

dochrysis is horizontally strigose and in M. chrysis re-

ticulated. Sternum of the mesothorax in M.

pseudochrysis is finely punctulated and horizontally

strigose in M. chrysis . The colour of M. chrysis is

bright green-blue and of M. pseudochrysis bright green

and the tergites II and III have a cupreous tint.

Miscophus sallitus Andrade, 1960

(fig. 14)

De Beaumont (1956) mentions a new species of

Miscophus from the Cape Verde Island of Sal. The

single male was later described by Andrade (1960) as

M. sallitus. Three females, collected in 1988 on the is-

land of Sal, were mentioned as belonging to the same

species (Simon Thomas & Wiering 1993). A descrip-

tion of a female is given below.

Description

Female 5 mm. Black; apical half of mandibles am-

ber ferruginous, apical internal side of scape light

brown; pedicel with light brown apical ring. Vertex,

pronotum, mesonotum, metanotum, and tergites

with rather marked cupreous tint.

129

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Clypeus arched with shining rim. Lower part of

face and clypeus with silver pubescence. Upper frons

very finely and closely punctulated. POL : OOL =

1:7, SOL 3 OO = 23, VOL 3 OOC = 20)

Flagellomeres IX and X are missing in all specimens,

VII and VIII on external side with shallow furrow.

Flagellomeres I-IV three and a half times as long as

wide at apex. Flagellomeres V-VII three times as long

as wide at apex. Flagellomere VIII twice as long as

wide at apex.

Pronotal collar, mesonotum, scutellum, and meta-

notum finely punctulated. Dorsal central part of pro-

podeum obliquely striated; side of dorsal part of pro-

podeum striation interrupted and sides of

propodeum obliquely striated.

Tergites extremely finely punctulated. Sternites

with a few long black hairs at apex.

Forewing hyaline, slightly clouded beyond the ner-

vation zone. Veins ending rather far from apex (fig.

14).

Material. - 32, Cabo Verde: Sal: Salinas Pedra da

Lume, 15.x1.1988, T. & M. Simon Thomas (ZMAN).

Miscophus funebris Honoré, 1944

Material. - 19, Niger, Niamey, Centre Agrymet,

piège malaise, 8.vi.1989, leg. J. Hollands (LUW), 16,

same data as female (ZMAN).

This species is new for Niger. Up till now

Miscophus funebris Honoré was only known from

Egypt. In structure the specimens from Egypt and

Niger are very much alike. The Niger specimens dif-

fer in extension of the ferruginous colour on legs and

first two gastral segments from the Egyptian speci-

men. All gastral segments of the Niger specimens are

light ferruginous, whereas the two first gastral seg-

ments of the Egyptian specimen are dark ferruginous.

The gastral segments from III to the apex of the

Egyptian specimen is also ferruginous. The legs of the

Niger specimens are ferruginous with an external

dark stripe on the tibiae, whereas the Egyptian speci-

men has dark brown legs.

130

Only one specimen of Egypt is known (Honore

1944) and one female and one male from Niger. No

specimens have been found in between Egypt and

Niger. From these few specimens it is impossible to

tell the colour variations are a constant characteristic

of the specimens from Egypt and Niger or a variation

in both populations.

Comparative notes. — See under M. rufigaster sp. n.

REFERENCES

Andrade, N. F., 1952. Sphecidae of Portugal. Genus

Miscophus Jurine. - Memorias e Estudos do Museo zool-

ogico da Universidade de Coimbra 211: 1-42.

Andrade, N. F., 1960. Palaearctic Miscophis: bicolor group

and isolated species (Hymenoptera, Sphecidae). —

Memorias e Estudos do Museo zoologico da

Universidade de Coimbra 262: 1-136.

Arnold, [G]., 1940. New species of African Hymenoptera

No. 4. — Annals of the Transvaal Museum 20: 101-143.

Beaumont, J. de, 1956. Sphecidae et Pompilidae (Hym.). —

Commentationes Biologicae 15 (21): 1-7.

Eady, R. D., 1968. Some illusstrations of microsculpture in

the Hymenoptera. — Proceedings of the royal

Entomological Society of London (A) 43: 66-72.

Everts, J. W., 1990. Environmental effects of chemical lo-

cust and grasshopper control. — F.A.O., Rome: 1-277.

Honoré, A.-M., 1944. Materiaux pour une monographie

des Miscophus d’Egypte. — Bulletin de la Societ€ Fouad

ler d’Entomologie 28: 45-79.

Kohl, F. F., 1894. Zur Hymenopterenfauna Afrikas. —

Annalen des naturhistorischen Museums, Wien (series 9)

3: 279-351.

Lomholdt, ©. C., 1985. A reclassification of the larrine

tribes with a revision of the Miscophini of Southern

Africa and Madagascar (Hymenoptera: Sphecidae). —

Entomologica scandinavica, supplement 24: 1-183.

Pulawski, W., 1968. Miscophus heliophilus sp. n. (Hym.,

Sphecidae). — Polskie Pismo entomologiczne 38: 285-

DISTA

Simon Thomas, R. T. & H. Wiering, 1993. Notes on the

Cape Verde Islands Fauna of Sphecidae and Apidae

(Hymenoptera). — Courier Forschungs-Institut Sencken-

berg 159: 403-409.

Received: 10 January 1994

Revised version accepted: 12 February 1995

J. van TOL' & Y. NORMA-RASHID*

! National Museum of Natural History, Leiden &° Department of. Zoology, University of Malaya,

Kuala Lumpur

THE GENUS EUPHAEA RAMBUR IN BORNEO

(ODONATA: EUPHAEIDAE)

Descriptions and records of Malesian Odonata, 3 2

The males are the most brilliant of oriental insects, and they are

exceeded by none in elegance of form.

Laidlaw 1924: 298

Tol, J. van & N. Norma-Rashid, 1995. The genus Euphaea Rambur in Borneo (Odonata:

Euphaeidae). Descriptions and records of Malesian Odonata, 3. — Tijdschrift voor

Entomologie 138: 131-141, figs. 1-40. [ISSN 0040-7496]. Published 15 June 1995.

The genus Euphaea Rambur is revised for the species occurring in Borneo. £. laidlawi Kimmins

of the tricolor species group is synonymized with E. subcostalis Selys. Euphaea ameeka sp. n., a

new species of the impar species group close to E. impar Selys, is described from Brunei. A key

to species of Euphaea occurring in Borneo is provided, and the distribution of all species is il-

lustrated.

Correspondence: J. van Tol, National Museum of Natural History, P.O. Box 9517, 2300 RA

Leiden, The Netherlands.

Key words. — Borneo, Euphaea, new species, revision.

) Part 2: Tijdschrift voor Entomologie 133 (1990): 273-279.

The dragonfly fauna of Brunei was virtually unex-

plored when a few years ago Drs. A. G. Orr and D. J.

Thompson started their studies on ecology and beha-

viour of these insects. A first collection identified by

the senior author revealed more than 80 species

(Thompson & van Tol in press), including several

species new to science. One of these species is de-

scribed in the present paper.

According to present knowledge there are eight

species of Euphaeidae in Borneo, viz. six species of

Euphaea and two of Dysphaea. Euphaea inaequipar

Selys, reported from Borneo by Selys (1859: sep. 7-8)

and Ris (1930), has proved to be a synonym of

Euphaea impar Selys from Sumatra (Lieftinck 1940).

The Euphaea species of Borneo are rather poorly

known, although adequate material is available in col-

lections worldwide, including the Leiden Museum.

One of the reasons is that the species of the complex

tricolor - subcostalis - laidlawi - subnodalis are very dif-

ficult to distinguish. Comparative overviews were

first provided by Laidlaw (1920, 1924), followed by

Ris (1930) who reviewed the species when he de-

scribed E. cora from the Philippines. In his descrip-

tion of E. laidlawi, Kimmins (1936) included sketch-

es of the characteristic seminal vesicles of the first

three species mentioned, and Lieftinck (1940) pro-

vided the same for subnodalis. Euphaea basalis seems

to be a more distantly related species.

The aim of the present paper is to provide an over-

view of all species indigenous to Borneo, and to de-

scribe their distribution. Since most species have been

sufficiently described before, we present a full de-

scription for the newly discovered species only, and

include short diagnoses for the other species.

Terminology of body parts follows Watson &

O’Farrell (1992).

MATERIAL

We have examined the specimens kept in the

Leiden Museum (RMNH), consisting of series of all

species known from Borneo and collected in most

parts of that island. From eastern Kalimantan there

are large series taken by M. E. Walsh-Held, while L.

Coomans de Ruiter collected extensively in the wes-

tern part of the island. Both collectors were active du-

ring the 1930’s. A smaller collection was made in the

central part of present-day Kalimantan, mainly taken

by Nieuwenhuis in the 1890’s. Collections from the

non-Indonesian part of Borneo are from Mt.

Merapok (Dent province, former British North

Borneo, collected at the end of the 19th century; this

131

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 1-12. Euphaea species, males. — 1, E. ameeka sp. n., thorax in lateral view. Brunei (JvT 8363); 2, E. cora, same. Samar Is.,

Philippines; 3, E. impar, same. Kalimantan (JvT 8362); 4, E. ameeka sp. n., inner view of superior anal appendage (JvT 5016);

5, E. cora, same; 6-12, last abdominal segment and anal appendages, left lateral view, 6, E. ameeka (JvT 5016); 7, E. cora; 8, E.

subcostalis (JvT 8737); 9, E. basalis (JvT 8733); 10, E. impar (JvT 8968); 11, E. subnodalıs, 12, E. tricolor (JvT 8802).

132

locality can be found on map 67 of Anonymus 1905,

see also Anonymus 1925/1926) (this locality is presu-

mably identical to Gn. Lumaku; Merapok is now a

village in the extreme NW corner of Sarawak, just at

the border with Sabah), from Sabah (Kinabalu area,

east and southern Sabah, among others collected by

the senior author), from Sarawak (leg. M. A. Lief-

tinck), and finally recent collections from Brunei,

brought together by Drs. A. G. Orr and D. J.

Thompson. Apart from these series, we were also able

to study single specimens of series kept in other insti-

tutions, obtained in exchange for the Leiden Museum

by M. A. Lieftinck.

SYSTEMATIC PART

Key to the males of Euphaea of Borneo

Keys to the Euphaea species of Borneo have been

published by Laidlaw (1920, 1924) and Ris (1930).

The present key includes some features not used in

those keys, and the new species described in this pa-

per. The males of all Bornean species of Euphaea have

a hyaline fore wing. The genus Dysphaea is characte-

rised by the nodus situated at the middle of the wing

(basal to nodus in Euphaea), and a smoothly rounded

dorsum of abdominal segment 10 (with dorsal pro-

jection in Euphaea). The superior anal appendages are

stronger curved inward in Dysphaea than in Euphaea.

The females are more difficult to distinguish, and are

not further discussed here.

1. Brown opaque area of hind wing at least from no-

dus to tip of wing (in some species area basal to

nodus also opaque to some extent); basal part of

opaque area with distinct metallic blue shine;

thorax brownish black (in life with greenish

tinge), sometimes with some slight yellowish lon-

gitudinal markings along the sutures; seminal

vesicle usually, but not in all species, with sharp

lateral projections, and posterior side broadly

LOUIS (OS 1ER) Ara 2

— Brown opaque area of hind wing confined to ap-

ical one-third, starting approximately halfway

between nodus and pterostigma, without con-

spicuous metallic shine, or hind wing fully hya-

line; thorax matt black with extensive blue lateral

markings; vesicle rounded laterally and posterior-

ly with a more or less conspicuous acute tip (figs.

DEE) en >

2. Hind wing fully opaque, brown, vesicle rounded

(figs. 25-26). Large species, hind wing 27-28

mim MviesKinabalutonly gn enen E. basalis

— Base of hind wing hyaline at least to Ax 10 ..... 3

3. ‘Auricles’ of tergite 2 pale yellowish on dorsal sur-

face, or at least much paler than rest of tergite 2;

opaque area of hind wing starting usually at or

VAN TOL & NORMA-RASHID: Euphaea in Borneo

just basal to nodus, subcostal space hyaline at

least in basal part; seminal vesicle rounded (figs.

27-28). Size variable, with hind wing 25-28 mm.

West, central and northern Borneo, but not in

theNinabalt region nt E. tricolor

— ‘Auricles’ on dorsal surface brownish black, con-

colorous with rest of tergite 2; opaque area of

hind wing distal to Ax 10-15 rather than distal to

nodus or just basal to it; lateral projections of ves-

ICICTACUILE RE een, 4

4. Larger species with hind wing more than 26 mm;

radial space in hind wing hyaline to nodus; vesi-

cle with lateral projections long and sharp (figs.

23-24); ‘auricles long and slender. Northwestern

Borneon me een E. subnodalis

— Smaller, but very variable species, hind wing 22-

24 mm; radial space of hind wing opaque from

same level as rest of opaque distal part; vesicle

very variable from broadly rounded and flat

(laidlawt) (figs. 13-14) to broadly transverse

with long acute lateral sides (‘subcostalis) (figs.

21-22); ‘auricles’ more or less triangular and usu-

ally shorter than vesicle in lateral view … … … … …

alia et: le AE Lo cene nd, CE E. subcostalis

5. Distal one-third to two-fifths of hind wing

opaque, brown without metallic blue shine; la-

brum, mandible and gena with distinct blue and

creamish blue markings … … E. impar

— Hind wing hyaline; labrum, mandible and gena

without, or with very faint, pale markings … … …

EENES Mate TUNE Seen E. ameeka sp. n.

The Euphaea tricolor group

The Euphaea tricolor group, as defined by Ris

(1930) consists of tricolor, subcostalis, subnodalıs,

amphicyana and basalis. Later, Kimmins (1936) ad-

ded Euphaea laidlawi to this group of species. All spe-

cies of this group, except amphicyana from

Mindanao, are endemic to Borneo. The status and

distribution of the Bornean species of Euphaea has

not been discussed since Lieftinck (1940: 343-344,

fig. 2) commented upon £. subnodalis.

The Euphaea tricolor group superficially represents

a homogeneous group of species, of which Laidlaw

(1924) presumes a close relationship to the masoni

group of mainland southeast Asia. On closer exam-

ination, two species are obviously distinct, viz. basalis

and tricolor (see key, and diagnoses below), but the

other three nominal species are closely related indeed.

Subnodalis, however, seems to be definable on its

large size, and the shape of the opaque area in the

hind wing. The seminal vesicle of specimens with this

combination of characters is also without variation,

but this shape, although of smaller size, also occurs in

subcostalis. We have been unable to find any charac-

133

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

ters that can unambiguously delimit a subdivision

among the rest of the specimens of the tricolor group.

Both the shape of the opaque area in the hind wing,

and the shape of the seminal vesicle, appeared to be

most variable. The shape of the vesicle, although fre-

quently considered of taxonomic importance, is also

very variable in both species of the Æ. impar group

discussed below. Consequently, subcostalis and laid-

lawi are here considered to represent only extremes of

a very variable species. Since the variation seems have

a geographical component, we cannot exclude that

microspecies within this complex can be distin-

guished when more material becomes available with

intermediate forms only occurring in narrow zones.

Actually, laidlawi specimens may represent such hy-

brids between ‘typical’ subcostalis and subnodalis. We

suggest a further study of the geographical variation

of this group in Borneo, based on an extensive sam-

pling program throughout Borneo, but particularly in

the northwestern part of the island.

Euphaea basalis (Laidlaw)

(figs. 9, 25-26, 36)

Pseudophaea basalis Laidlaw, 1915: 32 (type locality Kina

Balu).

Pseudophaea basalis. — Laidlaw 1920: 326-327 (Kinabalu);

Laidlaw 1924: 298 (key); Kimmins 1969: 306 (holotype

d).

Euphaea basalis. — Ris 1930: 88-89 (characters); Laidlaw

1931: 241; Laidlaw 1934: 550; Schmidt 1934: plate 17

(1) (pair of wings); Lieftinck 1954: 16 (distribution);

Asahina et al. 1983: fig. 39 (pair of wings).

Diagnosis. — Hind wing dark without basal hyaline

area, blue metallic in basal ca. 16 mm; costal, subcos-

tal and cubital space, and quadrangle black. Semical

vesicle round and rather flat, in lateral view hardly

reaching beyond ‘auricle’ (figs. 25-26); length 1.2

mm. Superior anal appendage (fig. 9) relatively slen-

der with inner ventral scoop-like tubercle visible in

distal three-fifths in lateral view.

Hind wing 27-29 mm.

Female unknown.

Distribution. — Only known from Mt. Kinabalu

(1000-1500 m) (fig. 36). Apparently occurring at low

densities only.

Material examined (RMNH). — 4 à , all from Sabah,

Kinabalu Mt. (Marai Parai, Tahuban river, Silau

Silau river, Liwagu river) (all in RMNH).

VAN TOL & NORMA-RASHID: Euphaea in Borneo

Euphaea subcostalis Selys

(figs. 10, 13-22, 37)

Euphaea subcostalis Selys, 1873: 483 (sep. 19) (original de-

scription, type locality Labuan).

Pseudophaea subcostalis. — Laidlaw 1915: 32-33 (record Kina

Balu); Laidlaw 1920: 326 (records Tampassuh, Sarabas,

Mt. Murud, Ulu Akar).

Euphaea subcostalis. — Laidlaw 1931: 241 (Borneo); Schmidt

1934: plate 17 (3) (pair of wings); Lieftinck 1954: 18

(references, distribution); Asahina 1983, fig. 39 (pair of

wings).

Euphaea laidlawi Kimmins, 1936: 77-78, figs. (original de-

scription, type locality Sarawak, Palawan). — Lieftinck

1954: 17 (references, distribution); Kimmins 1969: 307

(holotype d BMNH [= lectotype]). Syn. n.

Diagnosis. — Hind wing rather short and broad,

with opaque area distal to ca. Ax 12-15, with metallic

blue area extending to Px 6-8, distal margin oblique

to Costas costal space hyaline up to level of Ax 12-15,

but subcostal space brown. Seminal vesicle very vari-

able, see below. Anal appendages with superiors (fig.

10) rather robust, with scoop-like ventral inner tuber-

cle virtually extending from base to tip in lateral view.

Variation. — Kimmins (1936: 77-78) described £.

laidlawi based on two males collected by Everett at

Lawas in 1896. A specimen with the same locality

and collector is kept in the Leiden Museum, but it is

not certain that this represents the para(lecto)type.

The specimen fits the description exactly, including

the low profile of the vesicle in lateral view (figs. 13-

14). Such specimens have not been found on any oth-

er locality in Borneo. On the other hand, all kinds of

intermediates in the shape of the vesicle between this

form and the ‘genuine’ subcostalis can be found (figs.

15-22). The coloration of the wing and the shape of

tergite 10, is, as far as we can see, also very variable

and of no value for distinguishing taxa.

Distribution. — Widespread in lowland Borneo,

but rather scarce in NW part of Sabah, where it is

found only at lower altitudes of Mt. Kinabalu and

mountain ranges extending southward. At higher alti-

tudes it is replaced by £. subnodalis. Forms previously

considered as £. laidlawi are mainly from the coastal

region of NW Borneo (southern Sabah, Sarawak and

Brunei). E. subcostalis is especially common in streams

through lowland rain forest, where it has not been re-

corded above 600 m. The males can be found perch-

ing in high densities in sunny patches along rivulets

running through canopy gaps. Kimmins (1936) men-

Figs. 13-34. Seminal vesicles of male Euphaea species in ventral and lateral view. — 13-22, E. subcostalis. 13-14, Sarawak,

Lawas, paratype of E. laidlawi (JvT 9043); 15-16, Mt. Marapok (JvT 8775); 17-18, Sabah, Kinabalu area, Kamadaian river

(JvT 9153); 19-20, Sarawak, Merirai valley (JvT 8777); 21-22, Sabah, Danum valley, (JvT 8747); 23-24, E. subnodalis,

Sabah, Long Pasia, Maga river (JvT 8750); 25-26, E. basalis, Sabah, Kinabalu Park, Liwagu river (JvT 8735); 27-28, E. tri-

color, Borneo, Kalimantan, Singkawang (JvT 9052); 29-30, E. ameeka sp. n., Brunei (JvT 8375); 31, idem, ventral view on-

ly (vT 8376); 32, E. impar, Kalimantan, Bautau Bessi (JvT 8977); 33, same species and locality (JvT 8978); 34, E. cora,

Samar Is.

155

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

tions habitat segregation with £. tricolor. This obser-

vation is confirmed by Dr. A. G. Orr (pers. comm.)

‘At Kuala Belalong Field Studies Centre E. tricolor is

restricted to the main stream of the Sg. Belalong, and

is abundant. ZE. subcostalis is restricted to small side

tributaries and is also very abundant. The two may

occasionally interact at the mouths of such tribut-

aries’.

Material examined (RMNH). — 101 specimens from

Sabah (Gn Marapok, Kamadaian river, Quioin Hill,

Kalabakan, Danum Valley), Sarawak (Akar river,

Kapit, Matang, Lawas), Brunei (Belalong) and

Kalimantan (Midden Oost Borneo Expeditie, Batau

Bessi, Kariorang, Nunukan, Bengen river). The type

locality (Labuan) is not indicated on the map. This is-

land off the northwest coast of Borneo was an impor-

tant trading centre during the 19th century, and use-

less as an indication of the actual sampling station.

Euphaea subnodalis (Laidlaw)

(figs. 11, 23-24, 38)

Pseudophaea subnodalis Laidlaw, 1915: 31-32 (type locality

Mt. Kinabalu).

Pseudophaea subnodalis. — Laidlaw 1930: 326 (Kina Balu);

Kimmins 1969: 307 (holotype d BMNH).

Euphaea subnodalis. — Ris 1930: 88-89 (key, distribution);

Schmidt 1934: plate 17 (4) (pair of wings); Lieftinck

1940: 343-344, fig. 2 (references, material, compared

with subcostalis and laidlawi, anal appendages); Lieftinck

1954: 18 (references, distribution); Asahina et al 1983:

fig. 39 (pair of wings).

Diagnosis. — Very similar to subcostalis, but larger

and with hind wings relatively narrower. Hind wing

with opaque dark coloration distal to Ax 10-15, but

costal and radial space hyaline between base of wing

and nodus; metallic blue (or blue-green) basal area ex-

tending to Px 10, running only somewhat oblique to

Costa. Length of hind wing 26-28 mm. Seminal ves-

icle 1.1 mm, and profile usually considerably higher

than in swbcostalis, posteriorly broadly rounded and

lateral projections more or less acute and expanded.

‘Auricle’ considerably longer and narrower than in

subcostalis, reaching beyond the vesicle in lateral view

(figs. 23-24). Last abdominal segment remarkably el-

evated. Anal superior appendages with ventral tuber-

cle of more complex nature than in subcostalis, with a

transversal ridge running near base (see also tricolor)

(fig. 11).

Distribution and biology. — NW Borneo (Sabah)

above 1000 m (fig. 38). This species seems to inhabit

larger streams at higher altitudes than £. subcostalis s.].

It was apparently formerly common around 1000 m

on Mt. Kinabalu, but the population density seems to

be much lower today, possibly due to the heavy pres-

sure of urban settlements on river systems below 1500

136

m. In the southernmost part of Sabah, close to the

borders of Sarawak and Kalimantan, this species was

common at ca. 1000 m altitude in 1987, when this

area was still covered with virtually undisturbed sub-

montane rain forest. It occurs sympatrically with E.

tricolor in that area, with tricolor at much lower den-

sities.

Material examined (all RMNH). — 33 specimens

from Sabah (Kinabalu, Gn. Marapok, and various

rivers around Long Pasia).

Euphaea tricolor Selys

(figs. 12, 27-28, 39)

Euphaea tricolor Selys, 1859: sep. 8 (original description,

type locality Saratoga [= lapsus pro Sarawak]).

Pseudophaea tricolor. — Laidlaw 1920: 326 (records Retuh,

Saribas, Baram).

Euphaea tricolor. — Laidlaw 1931: 241 (Borneo); Schmidt

1934: plate 17 (2) (pair of wings); Lieftinck 1954: 18;

Asahina et al. 1983: fig. 39 (pair of wings).

Diagnosis. — Hind wing with opaque area distal to

nodus; blue metallic area from nodus to Ax 12 with

distal margin somewhat oblique. ‘Auricles’ pale

coloured on dorsal surface. Seminal vesicle more or

less diamond-shaped with rounded lateral sides (figs.

27-28). Anal appendages with superiors relatively

short with tip distinctly squarish; in lateral view

superior appendage is broader terminally than basal-

ly.

Distribution and biology. — Widespread in western

and central Borneo, but possibly lacking in the

Kinabalu region and certainly absent from Southeast

Borneo (fig. 39).

Material examined (RMNH). — 27 specimens from

Sabah (Gn. Marapok, Padas river), Sarawak (Dingey

river, Tebang), Brunei (Belalong), Kalimantan

(Mahakkam, Bloe-oe, Penaring, Bengkajang).

The Euphaea impar group

The group of Euphaea impar consists of three spe-

cies, viz. E. impar, E. cora and E. ameeka sp. n. Ris

(1930) considered E. inaequipar as a distinct species,

but Lieftinck (1940) synonymized both taxa. £. cora

has been recorded from the Philippine islands

(Mindanao, Samar) and is not further discussed here

(but see figs. 2, 7 and 34). It seems to be a rather rare

species (M. Hämäläinen, personal comm.). We were

able to study a male and female from Samar from the

collection of Dr. Hämäläinen. In the Leiden Museum

there are two female specimens from the Philippine

Islands (Basilan I., leg. Baker (ex UMMZ, no 2448-

1920), and Tobedo, 300 m, 15 May 1951, leg. H.

Townes) [locality not found].

Den D DR CECCO E

Eee

fun che

Euphaea ameeka sp. n.

(figs. 1, 6, 29-30, 40)

Type material. — Holotype d, ‘BRUNEI Darussa-

lam. Surroundings Ingai base camp along Ingai river.

Stream in freshwater swamp forest running into S.

Ingai, 20 min. walk S of base camp. 4°09°21”N

114°4256E. 12 Sep 1992. D. Thompson’ (in

RMNH) [no 12/9/11] [JvT 4888]. — Paratypes 146

1® (all Brunei Darussalam): Ulu Belai, between

Ingai and Topi rivers, c. 4°08°N 114°43’E, 3 Oct

1992, 15 (D. Thompson) [JvT 5016] in RMNH; Bkt

Teraja, 9 Sep 1993, 2d (A. G. Orr) [JvT 8363/8364]

in RMNH; Belait district, Sg. Mendaram, Sep. 1993,

2d (A.G. Orr) in RMNH [JvT 8374/8375]; Menteri

pools, 28 Oct 1993, 1d (A.G. Orr) in RMNH [JvT

8376]; Belait district, near Labi, Sg. Mendaram, 3

Feb. 1995, 88 19 (A. G. Orr) in RMNH [JvT

9792/9800].

Description

Male holotype. — Head. Labium in ventral view

with base pale coloured, the central part suffused with

brown; lateral lobe and movable hook brownish black

except for the pale base of lobe, median lobe glossy

brownish black with central part pruinose, median

cleft reaching to two-fifth the height of median lobe;

maxilla as far as visible in ventral view pale; labrum,

VAN ToL & NORMA-RASHID: Euphaea in Borneo

Fig. 35. Euphaea ameeka sp. n.

Secondary genitalia in ventral view.

(a) with seminal vesicle round (JvT

8375), and more elongate (JvT

8376).

1mm

mandibles and clypeus glossy brownish black; re-

maining part of head matt black without any pale

markings.

Thorax. Prothorax matt black with a paired elon-

gate transversal marking on middle lobe. Synthorax

(fig. 1) matt black with extensive bright blue (pale

blue in preserved material) coloration covering most

parts; dorsal carina with anterior triangle and ante-

alar region black, mesepisternum blue except for nar-

row line against mesopleural (=humeral) suture and

pre-alar ridge and posteriormost part of mesopleural

suture; mesokatepisternum (= mesinfraepisternum)

brownish black with a small blue spot against mese-

pisternum, and a pale creamish spot near lower side of

mesepimeron; metepisternum above interpleural su-

ture blue, anteriorly leaving only one-fifth black, met-

episternum below interpleural suture black leaving

anterior part up to and inclusive the stigma black, a

small black spot against interpleural suture circa half-

way the blue area; metepimeron black with an elon-

gate blue marking on upper half from anterior one-

fourth to hind margin, the marking widening at

posterior end and axtending along hind margin of

metepimeron.

Legs black. Wings with veins black, an indistinct

yellowish brown lustre; fore and hind wing transpar-

ent without any opaque black marking; fore wings

with 18-19 Ax, 24-26 Px; hind wing with 17 Ax, 20-

197

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Euphaea basalis Euphaea subcostalis

® Euphaea impar

MW Euphaea ameeka

138

21 Px; discoidal cell in all wings with one cross-vein;

2-3 Cux; origin of R3 2'/,-3 cells distal to subnodus;

pterostigma dark brown, covering 7-8 cells.

Abdomen. Fully matt black except for a paired sub-

quadrangular blue spot lateral on tergite 1, covering

circa 3/5 of segment length, and a very inconspicuous

longitudinal line on both sides of segment 2; segment

10 with distal half triangularly raised into a dorsal

projection. Seminal vesicle variable, from broadly

rounded (fig. 35a) to more elongate, posteriorly

acute. Appendages with superior (fig. 6) in distal half

with smoothly rounded ventral tubercle.

Measurements. Hind wing 25 mm, abdomen incl.

appendages 32 mm.

Female (paratype JvT 9800). — Pale coloration

darker than in male, dirty yellow (in life dark olive

green).

Head. Labium including lateral lobes yellowish

white, with only tips of middle lobe and movable

hooks glossy black; labrum glossy brownish black

with a paired subtriangular yellowish white spot, each

covering the lateral third, but leaving narrow black

anterior and lateral borders; mandibles glossy yellow-

ish white with a brownish black border dorsally and

anteriorly; clypeus brownish black; remaining part of

head matt black, except yellowish white gena, more

or less extending above clypeus, and an elongate pale

marking between lateral ocellus and antenna.

Thorax. Prothorax matt black, but middle lobe

with triangular dirty yellow markings laterally and

lateral tubercle dirty greyish yellow. Synthorax most-

ly blueish or dirty yellow; mesepisternum blueish, but

leaving black borders against dorsal carina, hind mar-

gin and dorsal one-fourth over mesopleural suture; al-

so a darker central line over dorsal two-thirds of mes-

episternum; rest of synthorax mostly pale coloured,

but dark patches over dorsal part of interpleural and

metapleural sutures, a subquadrangular marking on

mesokatepisternum, and metepimeron brown in cen-

tral part. Wings hyaline, with 19-21 Ax and 20-22 Ax

in fore wing, and 15-16 Ax and 19-20 Px in hind

wing.

Abdomen. Brown, darker on dorsal side, but seg-

ment 1 on lateral side with pale triangular marking

pointed anteriad, widening posteriorly and emargi-

nated on dorsal side; segment 2 with yellow lateral

line, segment 3-7 latero-anteriorly with triangular

pale marking, relatively large on segment 3 and small-

er on each following segment; these segments fol-

lowed by a longitudinal line on segments 3-5, wider

on 3 than on on following segments; lateral lines on

VAN TOL & NORMA-RASHID: Euphaea in Borneo

all segments posteriorly ending against transversal su-

ture.

Measurements: hind wing 24 mm, abdomen 25

mm.

Remarks

Variation. — Males. There is hardly any variation in

the coloration of the paratypes. Wing venation is as

follows: fore wing with 18-21 Ax and 23-26 Px; hind

wing with 15-18 Ax and 19-24 Px.

The shape of the seminal vesicle is variable as in E.

impar. in some specimens it has a distinctly elongate

shape, while in others in is more or less round (fig.

35). Hind wing 24-25 mm, abdomen incl. appendag-

es 29-31 mm.

Differential diagnosis. — Apparently a close relative

of Euphaea impar. Both species are structurally simi-

lar, but differ in the shape of the wing. The coloration

of male head and thorax differs in most characters

rather subtlely. However, the shape of the blue mark-

ings on the mesepisternum (cf figs. 1 and 3), and the

absence of any opaque dark markings in the hind

wing of Euphaea ameeka are distinctive. The blue and

creamish markings on the head of £. impar are hard-

ly visible in ameeka. The female is very similar to the

E. impar, including the markings on the head, but the

central black line over labrum is very narrow in £. im-

par. Dorsal half of mesepisternum brownish black in

E. impar, and with extensive dark olive green mark-

ings in £. ameeka. Pale lateral lines wider and extend-

ing to segment 7 (rather than to 5) in £. impar. E.

ameeka is also a close relative of another species of the

Euphaea impar group as defined by Ris (1930), viz. E.

cora Ris. Since this species also has fully hyaline

wings, the general appearance is very similar. The

structure of the male superior anal appendages of £.

cora (fig. 5), however, differs distinctly from both £.

impar and E. ameeka (fig. 4).

Although we have not found any structural genital-

ic characters, E. impar and E. ameeka are considered

distinct species, since they occur sympatrically in

Brunei. In that region these species show habitat seg-

ragation in the upper courses of streams in Brunei fo-

rests (A. G. Orr, pers. comm.). Despite their sympat-

rical occurrence, no intermediates have been noticed.

Specific isolation of calopterygoids is, as far as known,

especially enforced during the pre-mating phase of re-

productive behaviour. Isolation during copulation,

e.g. by differences in the shape of the secondary geni-

talia, is less common. It is generally only observable

between distantly related species. This phenomenon

Figs. 36-40. Distribution patterns of Euphaea species occurring on Borneo. All species except E. imparare endemic to Borneo.

Based on collection of RMNH Leiden and published records. — 36, E. basalis 37, E. subcostalis, the square is the type locality

of E. laidlawi; 38, E. subnodalis 39, E. tricolor, 40, E. impar (circle) and E. ameeka sp. n. (square).

139

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

is also known in the Calopterygidae and Chlorocy-

phidae. For instance, morphological differences

between the two European species of Calopteryx, C.

virgo (L.) and C. splendens (Harris), are nearly absent,

while hybrids have never been documented.

À comparative study of the reproductive behaviour

of E. impar and E. ameeka would provide an excellent

opportunity to contribute to our understanding of

the speciation processes of the euphaeids.

Etymology. — Named after daughter Ameeka of the

first collector of this species, Dr. D. J. Thompson. A

noun in apposition.

Distribution. — Brunei (fig. 40).

Euphaea impar Selys

(figs. 3, 8, 32-33, 40)

Selected references

Euphaea impar Selys, 1859: sep. 7 [type locality Mt. Ophir,

Malacca].

Euphaea impar. — Laidlaw 1902: 87 (record Aring river,

Peninsular Malysia); Ris 1930: 85-86 (key); Lieftinck

1940: 341-343 (synonymy, references, material);

Lieftinck 1954: 16-17 (references, synonymy, habitat,

distribution Malaya, Anambas, Sumatra, Bangka,

Borneo); Asahina et al. 1983, fig. 39 (pair of wings);

Asahina 1993: 5 (record Thailand); Thompson & van

Tol 1995 (records Brunei).

Euphaea inaequipar Selys, 1859: sep. 7 [type locality

Saratoga [= lapsus pro Sarawak]).

Euphaea inaequipar. — Ris 1930: 86 (key); Lieftinck 1940:

342 (status discussed).

Diagnosis. — Head with significantly more blue

coloration than in E. ameeka. Labrum dark brown

with a large central blue marking, leaving only narrow

dark margins on all sides, blue marking medially

transversed by a dark stripe, which is widest in the

middle; mandibles pale blue with anterior and basal

side dark, clypeus glossy brownish black; remaining

part of head matt black, except for blue coloured ge-

nae below antennae.

Thorax. Prothorax mainly matt black, middle lobe

without blue markings. Synthorax mainly blue

coloured, although blue markings are less extensive

than in £. ameeka. Mesepisternum only blue in ante-

rior lower third (fig. 3), remaining part matt black.

Wings with fore wing hyaline, base of hind wing hya-

line, opaque brown marking in hind wing from Px 12

to tip of wing.

Abdomen. Brownish black to black. Shape of sem-

inal vesicle as variable as in E. ameeka (fig. 32). Anal

appendages with distal half of superior smoothly

rounded ventral tubercle (fig. 10).

Measurements. Hind wing c. 25 mm, abdomen

incl. appendages c. 31 mm.

Geographical variation. — Specimens from Sumatra

140

are similar in structure and markings to those from

Borneo.

Distribution. — Thailand, Peninsular Malaysia,

Sumatra, Borneo.

Material examined (all RMNH). — 214 specimens

from Sabah (Gn Marapok), Brunei (Ingai river),

Kalimantan (Katoergau Mts, Raja Mts, Bengkajang,

Bagak river, Penaring, Poteng Mts, Batau Bessi,

Kariorang, Maloewi, Ampah, Santubang,

Gunungsari, Bengen river), Sumatra (Tanggamoes

Mts, Deli Laut Tador), Bangka Island and West

Malaysia (Penang Island, Kledang Mts, Templer

Park, Mupor river).

ACKNOWLEDGEMENTS

We would like to thank D. J. Thompson and A. G.

Orr most cordially for the opportunity to study and

describe Euphaea ameeka. DaveThompson and Bert

Orr also put their field notes of the ecology and beha-

viour of Euphaea species of Borneo at our disposal,

and extensively commented on a draft of this paper.

We also wish to thank Dr. M. Hämäläinen (Espoo,

Finland) for the loan of a pair of Euphaea cora, and

Dr. D.A.L. Davies for comments. Figure 35 was pre-

pared by E. J. Bosch (National Museum of Natural

History, Leiden).

REFERENCES

Anonymus, 1905. Stielers Hand-Atlas. Ed. 9. — Justus

Perthes, Gotha: map 1-100.

Anonymus, 1925/1926. Namenverzeichnis zu Stielers

Hand-Atlas. Ed. 10. [Hundertjahr-Ausgabe]. — Justus

Perthes, Gotha: 1-315.

Asahina, S., 1993. Further contributions to the odonate fau-

na of Thailand based on the recent collection made by

Bro. A. Pinratana and Mr. Yunosuke Kimura. — Tombo

36: 2-11, figs. 1-25.

Asahina, S., Y. Wada & T. Yamasaki, 1983. A revisiting to

North Borneo. Part 3. — Gekkan Mushi no 149: 23-29,

figs. 31-42. [In Japanese].

Coomans de Ruiter, L., 1936. Biologische waarnemingen

over beekjuffers van Borneo (Een groep van mooi ge-

kleurde libellen). — Tropische Natuur 25 (5): 69-78, figs.

1-5, plate 1.

Hämäläinen, M., 1985. Euphaea pahyapi spec. nov. from

peninsular Thailand (Zygoptera: Euphaeidae). —

Odonatologica 14: 341-344.

Kimmins, D. E., 1936. The Odonata of the Oxford

University Sarawak Expedition. — Journal Federated

Malay States Museums 18: 65-108, figs. 1-17.

Laidlaw, F. F., 1915. Contributions to a study of the drag-

onfly fauna of Borneo. — Part III. A collection made on

Mount Kina Balu by Mr. J. C. Moulton in September

and October 1913. — Proceedings Zoological Society of

London: 25-39, figs. 1-5.

Laidlaw, F. F., 1920. Contributions to a study of the drag-

onfly fauna of Borneo. Part IV. A list of species known to

occur in the island. — Proceedings of the Zoological

Society of London 1920: 311-342, figs. 1-4.

Laidlaw, F. F., 1924. A survey of the dragonfly fauna of the

Malay Peninsula, with notes on that of neighbouring

countries. Part II. Journal of the Malaysian Branch of the

Royal Asiatic Society 2 (3): 296-308, plates 3-5.

Laidlaw, F. F., 1931. A revised list of the dragonflies

(Odonata) of Borneo. — Journal of the Federated Malay

Museums 16 (3/4): 234-250.

Laidlaw, F. F., 1934. A note of the dragonfly fauna

(Odonata) of Mount Kinabalu and of some other moun-

tain areas of Malaysia, with a description of some new or

little known species. — Journal of the Federated Malay

States Museums 17(3): 549-561.

Lieftinck, M. A., 1940. Descriptions and records of South-

east Asiatic Odonata (II). — Treubia 17 (4): 337-390, figs.

1-18.

Lieftinck, M. A., 1954. Handlist of Malaysian Odonata. A

catalogue of the dragonflies of the Malay peninsula,

Sumatra, Java and Borneo, including the adjacent small

islands. — Treubia 22 (Suppl): i-xiii + 1-202 + 1 map.

Ris, F., 1930. Vier neue Calopterygiden (Odonata) von den

Philippinen und Palawan. — Mitteilungen der Münchner

Entomologischen Gesellschaft 20: 71-92, plate 4-7.

VAN Tor & NORMA-RASHID: Euphaea in Borneo

Schmidt, E., 1934. Odonata der Deutschen Limnolo-

gischen Sunda-Expedition. I. Imagines. Mit Beschrei-

bungen zweier neuer /ctinus aus Celebes und Neu-

Guinea. — Archiv für Hydrobiologie (Suppl) 13:

316-397, figs. 1-93 + plates 14-17.

Selys Longchamps, E. de, 1859. Additions au synopsis des

Caloptérygines. — Bulletin de l’Académie royale de

Belgique (série 2) 7 (7): sep. 1-16.

Selys Longchamps, E. de, 1873. Troisièmes additions au

synopsis des Caloptérygines. — Bulletin de l’Académie

royale de Belgique (série 2) 35 (5): sep. 1-66.

Thompson, D.J. & J. van Tol, 1995. Dragonflies from four

forest types in Brunei. — Brunei Museum Journal (in

press).

Watson, D. A. L. & A. F. O’Farrell, 1992. Odonata

(Dragonflies and damselflies), p. 294-310, figs. 17.1-

17.23. In: CSIRO, The insects of Australia. A textbook

for students and research workers. Melbourne University

Press.

Received: 23 December 1994

Accepted: 22 March 1995

141

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

BOOK REVIEW

Wiebes, J. T., 1994. The Indo-Australian Agaoninae

(pollinators of figs). — Verhandelingen, Koninklijke

Nederlandse Akademie van Wetenschappen,

Afdeling Natuurkunde (Series 2) 92: 1-208. [ISBN

0-444-85779-6]. Price NLG 85.00.

As a counterpart to the treatment of the African

Agaoninae, as published by Wiebes in 1992 (same se-

ries, volume 89: 1-298), this publication summarizes

the present knowledge of the systematics of the Indo-

Australian Agaoninae. In a short introductory chapter

the challenges for the future are enumerated, partly

consisting of previously described taxa incertae sedis,

and partly of still incompletely understood phyloge-

netic relationships. Wiebes also provides a short his-

tory of the study of the Indo-Australian fig wasps,

starting only in 1883. A special word is devoted to the

work of Girault whose taxa ‘took a long time before a

few of these were recognized, and most had better

been forgotten’. Not a hymenopterist myself, and

looking through the bibliography, I noticed that

these species were described in privately published

brochures with titles as ‘Some gem-like or marvellous

inhabitants of the woodlands heretofore unknown

and by most never seen nor dreamt of and ‘Some

beauties inhabitant not of commercial boudoirs but

of nature’s bossom, notably new insects’. I sincerely

hope that publications with such melodious titles will

142

be referred to for many years to come, since I consider

it very unlikely that articles with such titles will pass

an editor of any ‘serious’ journal now.

The present book, however, qualifies for any high

standard of the 1990’s. The keys are clear and exten-

sive, but not too verbose. The descriptions all follow

the same format, and provide careful enumerations of

relevant characters, as well as data on host fig and dis-

tribution. There is a surprisingly small amount of il-

lustrations.

The quality of printing of the illustrations is also

inferior to that of the text. I presume that only xerox

copies of the original art work have been used for re-

production in the present publication.

All students of Chalcidoidea will very much appre-

ciate the efforts of Wiebes and the publishers. The in-

formation, otherwise only available in many short pa-

pers, can now easily be retrieved from this book. I also

expect that other systematists with interest in co-evo-

lution will start a re-evaluation of the phylogeny of

this group in future (and, of course, of the figs as

well). Previously published authoritive papers by

Wiebes on this topic (e.g. in Ann. Rev. Ecol. Syst. 10:

1-12), and the summaries of basic data as in the pre-

sent volume, provide a sound basis for every student

of this difficult field of research.

[J. van Tol].

A. WELLS' & T. ANDERSEN’

! Australian Biological Resources Study, Canberra, Australia &° Museum of Zoology, Bergen,

Norway

TANZANIAN MICRO-CADDISELIES

(RRICGHOMMERA"EMDROPIIEIDAE)

Wells, A. & T. Andersen, 1995. Tanzanian micro-caddisflies (Trichoptera: Hydroptilidae). —

Tijdschrift voor Entomologie 138: 143-167, figs. 1-63. [ISSN 0040-7496]. Published 15 June

1995

The genus Tangatrichia is erected for a new species from the West Usambara Mountains in

north-eastern Tanzania, 7. gracilenta. In addition, 24 other new species of micro-caddisflies are

described from north-eastern and central Tanzania: Catoxyethira apicospinosa, C. bombolensis,

C. ciliata, C. crenulata, C. crinita, C. elongata, C. incompta, C. lanceolata, C. ruvuensis,

Scelotrichia glandulosa, Stactobia kaputensis, Dhatrichia cinyra, D. divergenta, Hydroptila bum-

bulensis, H. mazumbaiensis, H. morogorensis, H. tannerorum, H. usambarensis, Ugandatrichia

dentata, U. tanzaniensis, Orthotrichia bisetula, O. hydroptiloides, O. nigrovillosa, O. scutellata.

Two other established species, Catoxyethira ocellata Statzner, 1977 and Orthotrichia barnardi

Scott, 1963, are recorded from Tanzania for the first time and a new record is given for

Hydroptila cruciata Ulmer, 1912, one of the two species known previously to occur in

Tanzania.

The East African Hydroptilidae fauna now totals 36 species; the Tanzanian fauna comprises 29

species in eight genera including three genera in the tribe Stactobiini: Stactobia and Scelotrichia

each with one species and Catoxyethira with ten species. In Hydroptilini, a new monotypic ge-

nus Tangatrichia as well as two species in Dhatrichia, two species in Ugandatrichia, and six spe-

cies in Hydroptila are recorded. For the Orthotrichiini six species are listed in Orthotrichia.

Notes are given on distributions and apparent affinities of new taxa and a key to males of

Tanzanian genera and species is provided.

Correspondence: T. Andersen, Museum of Zoology, University of Bergen, Musépl. 3, N-5007

Bergen, Norway.

Key words. — Trichoptera, Hydroptilidae, new genus, new species, Tanzania.

Information on caddisflies of the Afrotropical

Region was summarised by Scott (1986) who listed

eleven hydroptilid genera (one endemic), and 49 spe-

cies. Gibon (1985, 1987a, 1987b, 1991) has de-

scribed 21 species from the Ghana, Guinea, Mali, the

Ivory Coast and Togo and Mey (1992) recently de-

scribed a new species from Kenya. For East Africa

alone, six micro-caddisfly genera and nine species

were listed by Johanson (1992), not including the

species described by Mey (1992). In a preliminary ac-

count of part of the Tanzanian fauna studied here,

from the Kaputu stream only, Andersen & Johanson

(1993) recorded eight genera and 12 species. In this

present study, which covers a broader area of north-

eastern and central Tanzania, 28 species are listed in

eight genera, 25 species and one genus being newly

described. New distribution records are given for

three established species, one of which is a widespread

African/Asian species, another a Central African spe-

cies and the third South African.

Of the two hydroptilid subfamilies only Hydropti-

linae has been recorded from the Afrotropical Region;

the more primitive Ptilocolepinae is restricted to the

Holarctic Region. Three tribes are represented: Stac-

tobiini Botosaneanu, 1956, Orthotrichiini Nielsen,

1948 and Hydroptilini Stephens, 1836. Two stacto-

biine genera, Catoxyethira Ulmer, 1912 and Stactobia

McLachlan, 1880 were listed for the East African fau-

na by Johanson (1992), but none for Tanzania. New

Tanzanian species are here described in each of these

genera, and in a third stactobiine genus, Scelotrichia

Ulmer, 1951. One species is referred to each of Scelo-

trichia and Stactobia and nine to Catoxyethira, and a

new record is given for Catoxyethira ocellata Statzner,

1977. The relatively high diversity of Catoxyethira in

tropical Africa, the features of the genus and its pos-

sible relationships with other members of the

Stactobiini are discussed briefly.

Five genera in the Hydroptilini are recorded here

for East Africa. Two of these, Ugandatrichia Mosely,

1939 and Hydroptila Dalman, 1819, were included

by Johanson (1992) in his list for East Africa.

Johanson also listed Argyrobothrus Barnard, 1934

which was synonymised with Oxyethira Eaton, 1873

143

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

by Ross (1948); since Johanson gave no support for

reinstating the name, his action is not accepted here.

The record is based on an unidentified species from

Uganda in a paper by Kimmins (1958). As Gibon

(1987b) notes, Oxyethira is poorly represented in

Africa in comparison, for example, with North

America or Eurasia. De Moor (1993) lists only three

species for South Africa and no Oxyethira males were

found in collections on which the present study is

based. Perhaps when more is known of the biology of

some of the other African genera, it will be found that

other species are filling the filamentous green algal-

feeding niche that Oxyethira species generally occupy

elsewhere.

Two new Ugandatrichia species are described here,

representing the first Tanzanian records of the genus;

other East African records are from Uganda and

Kenya. In Hydroptila, only H. cruciata Ulmer, 1912,

described from Tanzania (Ulmer 1912), was previ-

ously known from East Africa. Five new Tanzanian

species are described here, and a new record is given

for H. cruciata. The fourth Hydroptilini genus re-

corded for Tanzania, Dhatrichia Mosely, 1948, is al-

so recorded for the first time for East Africa. Two new

species are described, bringing the total for this

Afrotropical genus to five. The fifth genus, Tangatric-

hia gen. n., is newly described and monotypic; its af-

finities within the tribe are discussed.

Of the two genera in the tribe Orthotrichiini, only

Orthotrichia Eaton, 1873 is known to occur in East

Africa. The other genus, /thytrichia Eaton, 1873 is

recorded from South Africa, from immatures only

(Scott 1986). Four new species in Orthotrichia are de-

scribed here, raising the total for Tanzania to six spe-

cies, and a new record is given for Orthotrichia bar-

nardi Scott, 1963, known previously from South

Africa only.

MATERIAL AND METHODS

All material studied here was collected in connec-

tion with an expedition by the Museum of Zoology,

University of Bergen, to central and north-eastern

Tanzania during October to December 1990. One

Malaise trap at the Kaputu Stream in the West Usam-

bara Mountains was run by the staff at the Mazumbai

Forest Reserve Station for shorter periods during

January to August 1991. The main collecting sites

along the Kaputu Stream are described in detail by

Andersen & Johanson (1993). They also outline the

main methods used to collect and preserve the speci-

mens. Additional material was taken by sweep-net-

ting and light-trapping at other localities in the West

and East Usambara Mountains in the Tanga region

and at two lowland sites in the Morogoro region.

These localities are listed below; where maps were

144

available, Universal Transverse Mercator Grid (UTM)

reference and accurate altitude are given.

Amani East Forest Reserve, Zigi River (UTM:

37MDE609365), 470 m a.s.l., East Usambara Mts,

Tanga region, 2.xii.1990, sweep net. Fast flowing riv-

er, 10 m wide, with large stones; in lowland rain for-

est area.

Bombole (UTM: 37MDE596388), 830 m a.s.l., East

Usambara Mts, Tanga region, 1.xii.1990, sweep net.

Small rapid river with large stones; in forested area.

Dule, Bumbuli River (UTM: 37MDE403633), 1220

m asl, West Usambara Mts, Tanga region,

26.xi.1990, sweep net. Rapid river, 5 m wide, with

large stones; in a cultivated area.

Gogoi, (UTM: 37MDE378509) 1100 m a.s.l., West

Usambara Mts, Tanga region, 30.xi.1990, sweep net.

Two small streams with waterfalls; in area with rain

forest and cultivated land.

Gologolo, Lushoto (UTM: 37MDE182770), 1860 m

a.s.l., West Usambara Mts, Tanga region, 25.xi.1990,

sweep net. Rapid stream, 1.5 m wide, with stony and

sandy bottom and waterfalls; in rain forest area.

Kimboza, Ruvu River, about 150 m a.s.l., Moro-

goro region, 20.x.1990, sweep net. Large slow flow-

ing river, 20-30 m wide, with sandy base, in parts

with faster flow over stones; banks forested.

Mlesa (UTM: 37MDE6003745), 800 m a.s.l., East

Usambara Mts, Tanga region, 1.xii.1990, sweep net.

Rapid river with large stones and waterfalls; in forest-

ed area.

Shakoi River (UTM: 37MDE404730), 1420 m a.s.l.,

West Usambara Mts, Tanga region, 24.xi.1990,

sweep net. Rapid stream with large stones and heavi-

ly vegetated banks; in cultivated area.

Campus, Sokoine University of Agriculture, about

550 m a.s.l., Morogoro, Morogoro region, 26.x.-

11.xi.1990, light trap. Garden at the foothills of the

Uluguru Mts.

Specimens were prepared for study following the

methods of Wells (1990a). Attempts to associate fe-

males proved difficult and thus only males of new

species are described. All types are lodged in the col-

lection of the Museum of Zoology, University of

Bergen, Norway (ZMBN).

SYSTEMATIC PART

Check list of East African Hydroptilidae

Tribe Stactobiini

Catoxyethira Ulmer, 1912

syn. Sperotrichia Marlier, 1978

1. apicospinosa sp. n., Tanzania.

2. bombolensis sp. n., Tanzania.

ciliata sp. n., Tanzania.

crenulata sp. n., Tanzania.

crinita Sp. n., Tanzania.

elongata sp. n., Tanzania.

incompta sp. n., Tanzania.

lanceolata sp. n., Tanzania.

. ocellata Statzner, 1977, Zaire; Tanzania.

10. pinheyi Kimmins, 1958, Zimbabwe; Uganda.

11. ruvuensissp. n., Tanzania.

Do ND A Do

Scelotrichia Ulmer, 1951

syn. Madioxyethira Schmid, 1960

12. glandulosa sp. n., Tanzania.

13. kenyella (Mey, 1992), Kenya.

Stactobia McLachlan, 1880

syn. Afritrichia Mosely, 1939

14. aurea (Mosely, 1939), Uganda.

15. kaputensis Sp. n., Tanzania.

Tribe Hydroptilini

Dhatrichia Mosely, 1948

16. cinyra sp. n., Tanzania.

7% divergenta sp. n., Tanzania.

Hydroptila Dalman, 1819

18. bumbulensis sp. n., Tanzania.

19. cruciata Ulmer, 1912, Tanzania.

syn. H. hirra Mosely, 1948

20. mazumbaiensis sp. n., Tanzania.

21. morogorensis sp. n., Tanzania.

22. tannerorum sp. n., Tanzania.

23. usambarensis sp. n., Tanzania.

Tangatrichia gen. n.

24. gracilenta sp. n., Tanzania.

Ugandatrichia Mosely, 1939

25. acuta Mosely, 1939, Kenya; Uganda.

26. dentata sp. n., Tanzania.

27. minor Mosely, 1939, Kenya.

28. nigra Mosely, 1939, Uganda.

29. tanzaniensis sp. n., Tanzania.

Tribe Orthotrichiini

Orthotrichia Eaton, 1873

30. aequatoriana Kimmins, 1957, Uganda.

31. barnardi Scott, 1963, South Africa; Tanzania.

32. bisetula sp. n., Tanzania.

33. hydroptiloides sp. n., Tanzania.

34. nigrovillosa sp. n., Tanzania.

35. scutellata sp. n., Tanzania.

36. straeleni Jaquemart, 1956, Zaire;

Uganda.

Tanzania,

Fast Africa, as defined here, comprises the coun-

WELLS & ANDERSEN: Tanzanian micro-caddisflies

tries Tanzania, Kenya, Uganda, Burundi and Rwan-

da. From Burundi and Rwanda, however, no hydro-

ptilids are recorded. In the list only synonyms applied

to the Fast African species are included.

Key to males of hydroptilid genera and species in

Tanzania

1. Mesoscutellum with transverse suture ............. 2

— Mesoscutellum without suture ........................ 4

Db alispurto mula 0 DAN N ner

12%

13:

IR DARLE Scelotrichia glandulosa sp. n.

Jibialispur formulati 3, 4ior 142,40 ee. 3

diibialfspurstormulanl 0A En

DELAIN EN Stactobia kaputensis sp. n.

Tibial spur formula 1, 3, 4 (Catoxyethira Ulmer) 8

@cellifabsen kN as 5

lihreerocelliipreseneer. ne. RER 6

Forewing with jugal lobe, metascutellum penta-

gonal to triangular (Hydroptila Dalman) … … 17

Forewing without jugal lobe, metascutellum rec-

tangular (Orthotrichia Eaton) … … 22

Wings slender, attenuate apically, venation re-

ducedilsecitiesy4,92, 5A) en a Rene 7)

Wings broad, forewing rounded apically, venation

complete (fig. 26) (Ugandatrichia Mosely)....... 27

Antennal flagellar segments with scattered cloth-

ing hair; metascutellum triangular, truncate ante-

riorly (fig. 50) … … Tangatrichia gracilenta sp. n.

Antennal flagellar segments with clothing hair in

basal whorl; metascutellum rounded anteriorly

(DhatrichiaiNosely) Mesen en en. 28

.Abdominal sternite VIII with one pair of stout,

apico-lateral or mesal spines (figs. 12-25) … … 9

ze Abdominal sternite VIII without stout spines

(figs. 8-10)

Abdominal sternite VIII with apico-lateral spines

(ARS DIS RE 10

-Abdominal sternite VIII with mesal spines (figs.

EDs) vie: a see EA UT ent ERP ARS Eee 8 erie. 12

„in ARs Abdominal sternite VIII with cluster of lon

stout setae mid-ventrally (fig. 22) … … … …………

ERWIN SANE Catoxyethira crinita sp. n.

Abdominal sternite VIII without cluster of long,

stoutisetaelmidvenitrally pee. 11

. „Inferior appendages positioned mid-ventrally in a

deep excision in abdominal sternite VIII (see Statz-

ner, 1977, fig. 24) .. Catoxyethira ocellata Statzner

Abdominal sternite VIII without deep excision mid-

ventrali 25)" Catoxyethira ciliata sp. n.

re Abdominal sternite VIII with a shorter, thi

spine between the paired spines (fig. 21)

NE Catoxyethira crenulata sp. n.

..Abdominal sternite VIII with one pair of spines

ONE JA Aen GAREN 13

...Paired spines on abdominal sternite VIII about

equal in length to inferior appendage (fig. 13) ...

145

14.

15:

16.

18.

19?

20.

DIE

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

DER OR DE nc Catoxyethira ruvuensis sp. n.

Paired spines on abdominal sternite VIII almost

2 X length of inferior appendage, or longer ...14

Length of paired spines on abdominal sternite

VIII 2.5 to 3 X length of inferior appendage (fig.

een Catoxyethira elongata sp. n.

.... Length of paired spines on abdominal sternit

VIII no more than 2 X length of inferior append-

age (tigs3 14, loro. 15

Paired spines on abdominal sternite VIII straight

in lateral view (fig. 16), curved inwards in ventral

or (Enter MM) Catoxyethira lanceolata sp. n.

.Paired spines on abdominal sternite VIII curved

upwards in lateral view (fig. 14), more or less

stratehesinventraliview (HE LS) PE PU

Scbodndvocenndooonineco906 Catoxyethira apicospinosa sp. n.

Subgenital plate and inferior appendages elon-

gate, subequal in length; inferior appendages cy-

lindricaly(tiest Syl) peewee nn...

EE Catoxyethira bombolensis sp. n.

Subgenital plate about 2 X length of inferior ap-

pendages; inferior appendages broader basally

thankdistallya ss) ne...

en MES A Catoxyethira incompta sp. n.

With pair of sclerotised strap-like structures

above inferior appendages (fig. 48) ................ 18

Without pair of sclerotised strap-like structures

above inferior appendages (figs. 39, 42, 43, 46)

Inferior appendages in ventral view slender,

curved, narrowed slightly towards apex without

black spine apically (see Mosely, 1948, fig. 48)

EN ENGE Hydroptila cruciata Ulmer

Inferior appendages in ventral view stout basally,

apically bifid, with strong, black spine ventrally,

pale, slender spine dorsally (fig. 48) … … … … … …

Po ATER ER PERI Hydroptila bumbulensis sp. n.

Inferior appendage short, sub-glubose in ventral

viewsirregularinsshapen(fio 40) ea

eee RR Hydroptila tannerorum sp. n.

Inferior appendage elongate, cylindrical or some-

what sinuous, with length at least 3X width …

ASI E E e le PTE EEE EE 20

Aedeagus greatly dilated distally, with single

Smallispinesubapicall (ie)

sn det Hydroptila usambarensis sp. n.

Aedeagus slender or weakly dilated distally, with

onetomitworspinestapically ee. 21

Inferior appendages in ventral view dilated in ba-

sal half, tapered and out-turned apically; aedea-

gus with small spine apically (figs. 41, 42) ..........

N EN Hydroptila morogorensis sp. n.

Inferior appendages in ventral view sub-cylindri-

cal; aedeagus divided distally to form a pair of

spimesjinkseries (tips AS AA) nee

ruines oa Hydroptila mazumbaiensis sp. n.

146

DD. Abdominal segment IX laterally with pai

membranous, digitate processes with 1 or 2 apical

setaei (ee PS 5257) Seccccecoocace200d00000900000000006 23

— Abdominal segment IX without paired processes

laterallya(e e81183.5 803) ccovccsacoasocas00000000000000 24

23. Inferior appendages symmetrical (fig. 7) ............

weet Erden Orthotrichia barnardi Scott

— Inferior appendages asymmetrical (fig. 55) .........

tenia on Ann Orthotrichia bisetula sp. n.

24%ibialispurtfo mula 2 Aero Re

LITI pa ae. Orthotrichia hydroptiloides sp. n.

— libial/spurstormula 08 FRA 25

25. Inferior appendages fused, in ventral view rectan-

raul ee (Gel 499) cocesccce Orthotrichia scutellata sp. n.

— Inferior appendages discrete or partially fused, in

fonmyoftwounequalllobes) nen nen 26

26. Inferior appendages rounded, asymmetrical, the

left apically with small sclerotised knob; an elon-

gate process extending distally into a spine at

right apico-lateral angle of segment IX (fig. 61)

en era An Orthotrichia nigrovillosa sp. n.

— Inferior appendages tapered distally, the right one

twisted; a simple, apically rounded process at

right apico-lateral angle of segment IX (see

Jacquema SiR EEA

ee are in Orthotrichia straeleni Jacquemart

27. Inferior appendages set into deep excision in ab-

dominal sternite IX, irregular in shape, with small

inner spur subapically (fig. 28) … …… … nn

RR Ugandatrichia tanzaniensis sp. n.

— Abdominal sternite IX with shallow, ventral exci-

sion; inferior appendages stout, with inner spur at

base (figs. 27, 30) .... Ugandatrichia dentata sp. n.

28. Length of inferior appendages in ventral view

about Zwide SS) en

ee Fehde Dhatrichia divergenta sp. n.

— Length of inferior appendages in ventral view 3 to

4X width (fig. 37)......... Dhatrichia cinyra sp. n.

Stactobia McLachlan

Stactobia McLachlan, 1880: 515. Type species Hydroptila fusci-

cornis Schneider, 1845 by subsequent designation (Mosely

1933). For full generic synonymy see Marshall (1979).

The genus Stactobia was reviewed by Marshall

(1979). It is well represented in the Palaearctic and |

Afrotropical regions and some parts of the Oriental |

region. The larvae are scrapers, feeding on organic |

material in madicolous and hydropetric habitats. One |

species, placed by Marshall (1979) in the vaillanti |

species group, was known previously from East |

Africa, from Uganda. The new species resembles the |

West African S. vaillanti Schmid, 1959 described

from Guinea by Schmid (1959), more closely than |

the East African S. aurea (Mosely, 1939).

WELLS & ANDERSEN: Tanzanian micro-caddisflies

Figs. 1-3. Stactobia kaputensis sp. n., male genitalia. 1, dorsal view; 2, ventral view; 3, lateral view. — Figs. 4-7. Scelotrichia glan-

dulosa sp. n., male. 4, wings; 5, genitalia, ventral view; 6, genitalia, lateral view; 7, genitalia, dorsal view. — Abbreviations. —

ae: aedeagus; c.gl.: costal gland; inf.app.: inferior appendage; mes.pr.: mesal process; sp.: spine; subg.pl.: subgenital plate;

VIII, IX, X: abdominal segments VIII, IX, X.

Stactobia kaputensis sp. n. trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

(figs. 1-3) 194). — Paratypes: 364 (3 slides) data as for holotype

except loc. 2-11, 30.x.-26.xi.1990; 26 data as for

Type material. — Holotype d, TANZANIA, Tanga holotype except loc. 7, 1.-8.viii.1991; 2d data as for

region, West Usambara Mts, Mazumbai, Kaputu holotype except loc. 5, 6. xi. 1990, sweep net; 2d

Stream, loc. 9, 1450 m a.s.l., 19.-23.xi.1990, Malaise data as for holotype except Gogoi, 30.xi.1990, sweep

147

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

net; 36 data as for holotype except East Usambara

Mts, Amani Fast Forest Reserve, Zigi River,

2.xii.1990, sweep net; 9d data as for holotype except

East Usambara Mts, Bombole, 1.xii.1990, sweep net;

14 data as for holotype except East Usambara Mts,

Mlesa, 1.xii.1990, sweep net.

Description

Small, and (in alcohol) uniformly brown; antennae

with 18 segments; ocelli 3; tibial spurs 1, 2, 4.

Anterior wing length 1.4-2.1 mm. Genitalia as in figs.

1-3. Abdominal sternite VII with a long straight me-

sal process, reaching posteriorly almost to the apical

margin of segment VIII. Segment IX well developed.

Segment X extending distally in a pair of stout, elon-

gate, sclerotised spines. Inferior appendages broad-

based, forming irregular lobes distally. A distinct lobe

above the inferior appendages and apparent in lateral

view probably represents the subgenital plate.

Aedeagus elaborate and swollen apically, with several

sets of spines.

Etymology. — Named for the Kaputu Stream in the

West Usambara Mountains, Tanzania.

Biology and distribution. — Taken at several sites in

both East and West Usambara Mountains, both

along smaller streams and larger rivers at altitudes

between 470 m and 1650 m a.s.l. In the Malaise traps

along the Kaputu Stream this species was trapped fre-

quently both at sites with slow flow over gravel,

stones and mud, and at sites with fast flow over bed-

rock and stones (see Andersen & Johanson 1993).

Swarming specimens were netted by day mostly in ar-

eas of fast flow over bedrock and stones or near water-

falls.

Remarks. — Stactobia kaputensis sp. n. closely re-

sembles S. vaillanti from which it is distinguished by

the shape of the inferior appendages in lateral view

and the bigger spines on the aedeagus.

Scelotrichia Ulmer

Scelotrichia Ulmer, 1951: 73. Type species: Scelotrichia sa-

ranganica Ulmer, by original designation and monotypy.

See Wells (1990a) for full generic synonymy.

Worldwide, the genus Scelotrichia is represented by

some 29 species, the majority described from SE Asia.

Only two African species were known previously, S.

marshalli (Statzner, 1977) from Zaire and S. kenyella

(Mey, 1992) from Kenya. Both of these species and a

new species described here are relatively conservative

in genitalic form compared to some of the Bornean

and New Guinean species. All lack areas of scent

scales (androconia) on the forewings, but the new

species has a small sac-like costal gland at the base of

148

a tuft of hair in the forewing. At least in the

Tanzanian species, the body vestiture is hairy, not

scaly as in many of the species of the Malay

Archipelago. Conceivably, the African species repre-

sent a relict fauna.

Like Stactobia, some Scelotrichia species are madic-

olous. Others are found living on aquatic mosses on

rocks in fast flowing waters (see Schmid 1960, Wells

1990a, 1990b). No details are known of the biology

of these African species.

Scelotrichia glandulosa sp. n.

(figs. 4-7)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 5, 1650 m a.s.l., 2.-6.x1.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

195). — Paratypes: 334 (5 slides) data as for holotype

except loc. 2-11, 30.x.-26.xi.1990; 14d (4 slides)

data as for holotype except 31.x. & 6.xi.1990, sweep

net; 5d data as for holotype except loc. 8, 5.xi.1990,

sweep net; 254 data as for holotype except Gologolo,

Lushoto, 25.xi.1990, sweep net.

Description

Males relatively large, in alcohol uniformly dark

coloured; antennae 19-segmented, segments elonga-

te; ocelli 3; tibial spurs 0, 2, 4, with a small rounded

knob in the place of the spur on foreleg. Forewing

(fig. 4) with a small sac-like gland adjacent to a tuft of

hair in the proximal costal region; length 2.8-3.5

mm. Genitalia as in figs. 5-7. Segment VIII reduced

ventrally, rounded dorsally. Segment IX (dorsal plate)

appears to be reduced, possibly represented by the

paired sclerotised lobes shown in fig. 7. Inferior ap-

pendages stout, tapered apically and with small serra-

tions on inner margin subapically, fused basally and

extended anteriorly to a pronounced round mesal

structure at the margin of sternites VII and VIII.

Subgenital plate bifid and rugose apically, lobes sepa-

rated by a deep, rounded concavity. Aedeagus simple,

straight, apically bilobed in ventral view.

Etymology. — From the Latin glandulosus, glandu-

lous, referring to the gland in the forewing.

Biology and distribution. — Known only from the

West Usambara Mountains, where it was taken along

streams at altitudes between 1400 m and 1860 m

asl. In the Malaise traps along the Kaputu Stream

this species was trapped in highest numbers at a site

with fast flow over gravel and stones above a waterfall

(see Andersen & Johanson 1993). Swarming speci-

mens were netted by day mostly near waterfalls.

Remarks. — This species appears to resemble quite

closely the other African congeners, but differs in the

form of the ventral plate and tergite X and in having

fine serrations on inner distal margin of inferior ap-

pendages. Neither Statzner (1977) nor Mey (1992)

mention any glands on the wings or make any men-

tion of anything unusual about the wings of their spe-

cies. Females of Scelotrichia glandulosa sp. n. were

collected with the male and have the usual conserva-

tive form.

Catoxyethira Ulmer

Catoxyethira Ulmer, 1912: 82. Type species: Catoxyethira

fasciata Ulmer, by monotypy.

Sperotrichia Marlier, 1978: 294. Type species: Sperotrichia

mali Marlier, by original designation and monotypy.

Synonymised by Marshall (1979).

Catoxyethira is proving to be particularly diverse in

tropical Africa (see Gibon 1985, 1987a, 1991). This

work adds nine new species to the previous 17 (one

with two subspecies) known from Africa. The only

other members of the genus are a dubious Taiwanese

species known only from immatures and originally

described in Hydroptila (see discussion in Marshall

1979), and a Vietnamese species (Oläh 1989) which

may well be referable to Chrysotrichia Schmid, 1958.

One species has been described from each of South

Africa (Kimmins 1958), Zimbabwe (Morse 1974),

and Mali (Marlier 1978), three from Zaire (Ulmer

1912, Statzner 1977), six from the Ivory Coast

(Gibon 1985) and five from Guinea (Gibon 1987a,

1991);

Catoxyethira shares many features of general body

form with the Holarctic genus Stactobiella Martynov,

1924, the neotropical Bredinia Flint, 1968 and the

SE Asian/New Guinean/northern Australian genus

Chrysotrichia, particularly the very short metascutel-

lum on the adult thorax. Only the spur number dif-

fers, Catoxyethira having a tibial spur formula of 1, 3,

4. The African species fall into two main groups on

the basis of genitalic form: those with and those with-

out a pair of dark spines on abdominal sternite VIII.

Species lacking the spines are scarcely distinguishable

from some of the SE Asian Chrysotrichia. Although

from available information it appears that the larval

forms of Catoxyethira and Chrysotrichia are also simi-

lar, these similarities are probably plesiomorphous.

Members of the two genera probably also occupy

comparable niches, but again the hydropetric niche

may well be plesiomorphic in the tribe.

One African species, Catoxyethira pinheyi Kim-

mins, 1958, is quite widespread, being recorded from

Victoria Falls, Mali and the Ivory Coast. It closely re-

sembles (and may be the senior synonym of)

Catoxyethira ocellata Statzner, 1977 from Zaire,

which is here recorded from Tanzania. Most other

WELLS & ANDERSEN: Tanzanian micro-caddisflies

species appear to be quite restricted in their distribu-

tions.

Catoxyethira incompta sp. n.

(fig. 8)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 4, 1680 m a.s.l., 20.-26.x1.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

196).

Description

Antennae damaged. Anterior wing length 1.7 mm.

Genitalia as in fig. 8. Abdominal segment VIII long,

produced distally into lateral lobes, spines absent.

Segment IX short, retracted within segment VII.

Segment X membranous, broadly rounded and

slightly concave apico-mesally. Subgenital plate bro-

ad-based, tapered distally to narrow apex; apex bifid,

lobes acute. Inferior appendages short, slender and in

close proximity distally, with paired long setae dorsal-

ly. Aedeagus simple, straight.

Etymology. — From the Latin incomptus, un-

adorned, referring to the relatively simple appearance

of the male genitalia.

Biology and distribution. — Known only from the

type locality, where the specimen was collected from

a site of moderate flow over fine sand, gravel and

stones (see Andersen & Johanson 1993).

Remarks. — Catoxyethira incompta sp. n. most

closely resembles C. improcera Statzner, 1977 and C.

mali (Marlier, 1978). It differs in lacking the lateral

processes above the inferior appendages seen in C.

improcera and the paired dorsal structures of C. mali

which are referred to by Marlier (1978) as anal ap-

pendages.

Catoxyethira bombolensis sp. n.

(figs. 9, 10)

Type material. — Holotype d, TANZANIA, Tanga

region, East Usambara Mts, Bombole, 830 m a.s.l.,

1.xii.1990, sweep net, ZMB’s Tanzania Expedition

(slide, ZMBN No. 197). — Paratypes: 7d (4 slides) data

as for holotype; 16 data as for holotype except Amani

East Forest Reserve, Zigi River, 2.x11.1990, sweep net.

Description

Antennae 18-segmented. Anterior wing length 1.3-

1.5 mm. Genitalia as in figs. 9, 10. Abdominal ster-

nite VIII elongate, without paired spines, with a U-

shaped apico-mesal cleft. Tergite X about half length

of inferior appendages, membranous. Inferior appen-

dages elongate, cylindrical. Subgenital plate forming a

149

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

inf.app.

= inf.app.

Fig. 8. Catoxyethira incompta sp. n., male genitalia, ventral view. — Figs. 9, 10. Catoxyethira bombolensis sp. n., male genitalia.

9, lateral view; 10, ventral view. — Figs. 11-13. Catoxyethira ruvuensis sp. n., male genitalia. 11, dorsal view; 12, ventral view;

13, lateral view. — Figs. 14, 15. Catoxyethira apicospinosa sp. n., male genitalia. 14, lateral view; 15, ventral view.

Abbreviations. — As for figs. 1-7.

pair of straight narrow processes in ventral view; in la- in the East Usambara Mountains, Tanzania.

teral view slightly swollen apically, and curved vent- Biology and distribution. — Known only from East |

rally. Usambara Mountains, from beside rivers with fast

flow over large rocks at altitudes between 470 m and

Etymology. — Named after the village of Bombole 830 ma.s.l.

150

Remarks. — This species groups with Catoxyethira

improcera and C. mali in lacking paired, dark, socket-

ed spines. However, the form of sternite VIII with the

small apico-mesal excision is distinctive.

Catoxyethira ocellata Statzner

Catoxyethira ocellata Statzner, 1977: 396.

Biology and distribution. — Catoxyethira ocellata is

known from Zaire and Tanzania. The Ruvu River at

the site at which the five specimens were collected is

large and slow flowing with a sandy and stony sub-

strate.

Remarks. — This species closely resembles Catoxye-

thira pinheyi Kimmins, 1958 from Zimbabwe, Ivory

Coast and Uganda. It shares with C. pinheyi features

such as inferior appendages set into a deep mid-ven-

tral excision in the apical margin of abdominal ster-

nite VII and dorsal spines on sclerite VIII at apico-

lateral angles. Catoxyethira ocellata has three pairs of

fine setae on each side of the inferior appendages

whereas, according to Kimmins (1958) and verified

in the type by Wells, there is a single slender, acute

spine on each side of the inferior appendages in C.

pinheyi the shape of the inferior appendages also dif-

fers slightly. These differences are probably minor

and the two may be synonyms.

Material examined. — 54 (4 slides) TANZANIA, Mo-

rogoro region, Kimboza, Ruvu River, 20.x.1990,

sweep net, ZMB’s Tanzania Expedition.

Catoxyethira ruvuensis sp. n.

(figs. 11-13)

Type material. — Holotype d, TANZANIA, Moro-

goro Region, Kimboza, Ruvu River, 150 m a.s.l.,

20.x.1990, sweep net, ZMB’s Tanzania Expedition

(slide, ZMBN No. 198). — Paratypes: 4d (2 slides) data

as for holotype.

Description

Antennae 18-segmented. Anterior wing length 1.1-

1.3 mm. Genitalia as in figs. 11-13. Abdominal seg-

ment VIII stout, laterally extended into broadly roun-

ded lobes about as long as inferior appendages, spines

ventrally at base of lobes; posterolaterally a pair of

elongate moderately thick setae. Tergite X broadest in

basal half, rounded apically, covered with spinules.

Inferior appendages subrectangular in ventral view, a

pair of thick setae mesolaterally. Subgenital plate ap-

pears to be reduced. Aedeagus simple, straight.

Etymology. — Named after the Ruvu River, origi-

nating in the Uluguru Mountains, Tanzania.

Biology and distribution. — Known only from the

WELLS & ANDERSEN: Tanzanian micro-caddisflies

type locality at Ruvu River, a large slow-flowing river

with sandy and stony substrate.

Remarks. — This species groups with Catoxyethira

pinheyi and C. ocellata in the form of tergite X and

general arrangement of the genitalia. However, the

inferior appendages are only shallowly set into the

posterior margin of segment VIII, and the paired

spines are positioned ventrally beside the inferior ap-

pendages, not apico-laterally.

Catoxyethira apicospinosa sp. n.

(figs. 14, 15)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Dule, Bumbuli River,

1220 m a.s.l., 26.xi.1990, sweep net, ZMB’s Tanzania

Expedition (slide, ZMBN No. 199).

Description

Antennae damaged. Anterior wing length 2 mm.

Genitalia as in figs. 14, 15. Abdominal segment VIII

stout, short mid-dorsally and ventrally, paired spines

of moderate length, positioned ventro-laterally, cur-

ving upwards in lateral view, slightly curved in ventral

view, and with a number of spinules in apical region.

Tergite X membranous, rounded. Inferior appenda-

ges tapered slightly in distal half. Subgenital plate in

form of a small sclerotised median crescentic structu-

re. Aedeagus straight, simple.

Etymology. — From the Latin, apex, top, and spine,

thorn, referring to the few stout spinules at the apex

of the ventral spines.

Biology and distribution. — Known only from the

type locality in West Usambara Mountains, from a

river with fast flow and a base of large rocks.

Remarks. — This species closely resembles

Catoxyethira lanceolata sp. n., but abdominal segment

VIII is far wider, the ventral spines curve upwards in

lateral view and have few and stouter spinules near

the apex; the inferior appendages are narrower.

Catoxyethira lanceolata sp. n.

(figs. 16, 17)

Type material. — Holotype d, TANZANIA, Tanga

region, East Usambara Mts, Mlesa, 800 m a.s.l.,

1.xii.1990, sweep net, ZMB’s Tanzania Expedition

(slide, ZMBN No. 200). — Paratype: 16 data as for

holotype.

Description

Antennae damaged. Anterior wing length 1.6 mm.

Genitalia as in figs. 16, 17. Abdominal sternite VIII

very short, paired spines stout, curved, covered in fine

spinules and arising slightly off centre midway along

151

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

152

the length of the sternite, extending distally beyond

the inferior appendages, inturned apically, in lateral

view slender, straight. Tergite X membranous, broad

in ventral view, triangular in lateral view. Inferior ap-

pendages stoutly ovate in ventral view, in lateral view

about three times as long as wide. Subgenital plate in

ventral view broad with an apico-mesial downwardly

directed sclerotised process, in lateral view appearing

as a small down-turned lobe. Aedeagus simple,

straight.

Etymology. — From the Latin /ancea, light spear,

referring to the lance-like ventral spines on abdomi-

nal sternite VIII.

Biology and distribution. — Known only from the

type locality in the East Usambara Mountains, from a

small, fast-flowing river with waterfalls and a base of

large rocks.

Remarks. — This species and C. apicospinosa sp. n.

from the same locality show close resemblance, yet

each is quite distinctive. Particularly, in this species

the paired spines are longer and in ventral view are

curved, in lateral view straight and lance-like.

Catoxyethira elongata sp. n.

(figs. 18, 19)

Type material. — Holotype d, TANZANIA, Moro-

goro region, Kimboza, Ruvu River, 150 m a.s.l.,

20.x.1990, sweep net, ZMB’s Tanzania Expedition

(slide, ZMBN No. 201). — Paratypes: 2d (slides) data

as for holotype.

Description

Antennae 18-segmented, flagellar segments elon-

gate. Anterior wing length 1.5-1.9 mm. Genitalia as

in figs. 18, 19. Abdominal sternite VIII sub-quadra-

te in ventral view; paired spines symmetrical, slen-

der, somewhat sinuous, covered in minute spinules,

arising in close proximity slightly behind antero-

ventral margin and extending beyond tergite X.

Tergite X short, membranous, rounded apically.

Inferior appendages elongate. Subgenital plate

slightly longer and narrower than tergite X.

Aedeagus straight, simple.

Etymology. — From the Latin elongatus, prolonged,

referring to the general shape of the male genitalia.

Biology and distribution. — Known only from the

type locality at Ruvu River, a large slow-flowing river

with sandy and stony substrate.

WELLS & ANDERSEN: Tanzanian micro-caddisflies

Remarks. — In general form, male genitalia of this

species resemble those of Catoxyethira lanceolata sp.

n. and C. apicospinosa sp. n., but differ mainly in

that the ventral spines are narrower and more elon-

gate and arise closer to the anterior margin of ster-

nite VIII.

Catoxyethira crenulata sp. n.

(figs. 20, 21)

Type material. — Holotype d, TANZANIA,

Morogoro region, Morogoro, Sokoine University of

Agriculture, 550 m a.s.l., 26.x.-11.xi.1990, light trap,

ZMB’s Tanzania Expedition (slide, ZMBN No. 202). —

Paratypes: 114 (3 slides) data as for holotype.

Description

Antennae 17-segmented. Anterior wing length 1.5-

1.8 mm. Genitalia as in figs. 20, 21. Sternite VIII

with paired spines slender, sclerotised, barbed in basal

half, extending posteriorly to just slightly below the

tips of the inferior appendages, basally between the

spines another short spine. Tergite X broad, mem-

branous. Inferior appendages stout, short, with a sub-

apical notch on the inner margin. Subgenital plate

produced and downturned apically. Aedeagus slen-

der, simple.

Etymology. — From the diminutive form of the

Latin crena, notch, and the suffix -atus, provided

with, referring to the small, subapical notch on the

inner margin of the inferior appendage.

Biology and distribution. — Known only from the

type locality on the campus of Sokoine University of

Agriculture in Morogoro. The university is situated

in the foothills of the Uluguru Mountains and sever-

al small, slow-flowing streams and ditches are found

on the area.

Remarks. — This species closely resembles

Catoxyethira veruta Morse, 1974 from Kariba,

Zimbabwe (as Southern Rhodesia) from which it is

distinguished in ventral view by the more closely

placed ventro-lateral spines, the small subapical

notch on the inner margin of the inferior appendag-

es, and the shape of the subgenital plate, and in lat-

eral view by the irregular shape of the inferior ap-

pendages and of tergite X (dorsal plate). Morse

(1974) likened C. veruta to C. pinheyi. However, re-

semblances between C. pinheyi and C. crenulata sp.

n. are slighter than those of the new species and C.

veruta.

Figs. 16, 17. Catoxyethira lanceolata sp. n., male genitalia. 16, lateral view; 17, ventral view. — Figs. 18, 19. Catoxyethira elon-

gata sp. n., male genitalia. 18, lateral view; 19, ventral view. — Figs. 20, 21. Catoxyethira crenulata sp. n., male genitalia. 20,

lateral view; 21, ventral view. — Figs. 22, 23. Catoxyethira crinita sp. n., male genitalia. 22, ventral view; 23, lateral view. —

Figs. 24, 25. Catoxyethira ciliata sp. n., male genitalia. 24, lateral view; 25, ventral view.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

3 se È

Figs. 26, 28, 29. Ugandatrichia tanzaniensis sp. n., male. 26, wings (scale bar = 1.0 mm); 28, genitalia, ventral view; 29, gen-

italia, lateral view. — Figs. 27, 30, 31. Ugandatrichia dentata sp. n., male genitalia. 27, ventral view; 30, dorsal view; 31, later-

al view. — Abbreviations. — inf.app.: inferior appendage; subg.pl.: subgenital plate.

Catoxyethira crinita sp. n.

(figs. 22, 23)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 5, 1650 m a.s.l., 6.-12.x1.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

203). — Paratypes: 4d (slides) data as for holotype ex-

cept loc. 2-10, 30.x.-3.xii.1990; 1d (slides) data as

for holotype except East Usambara Mts, Bombole,

1.x11.1990, sweep net.

154

Description

Antennae 18-segmented. Anterior wing length 1.5-

2.0 mm. Genitalia as in figs. 22, 23. Sternite VIII

elongate, length about twice width, spines short, at

apico-lateral angles; a cluster of long, stout, dark setae

mesally on sternite. Tergite X broad, a little longer

than inferior appendages, in lateral view stout, apical-

ly truncate. Inferior appendages simple lobes, about

twice as long as wide, rounded apically. Subgenital

plate a narrow, apically rounded lobe. Aedeagus |

simple, slender, straight.

Etymology. — From the Latin crinis hair, and the

suffix -atus, provided with, referring to the meso-ven-

tral tuft of long hair on sternite VIII.

Biology and distribution. — Taken both in the West

and East Usambara Mountains, beside slow to fast flow-

ing streams and rivers flowing over stones, sand, gravel

and mud, at altitudes between 830 m and 1650 m a.s.l.

Remarks. — In the position and form of the lateral

spines on segment IX, C. crinita sp. n. is similar to C.

disymetrica Gibon, 1991 from Guinea. It is distin-

guished, however, from all other African species by

having a cluster of elongate dark coloured setae medi-

ally on sternite VIII.

Catoxyethira ciliata sp. n.

(figs. 24, 25)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 3, 1720 m a.s.l., 20.-29.x1.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

204). — Paratypes: 5034 (8 slides) data as for holo-

type except loc. 2-10, 30.x.-13.xii.1990; 29d data as

for holotype except loc. 7, 4.i.-8.vii.1991; 36 data as

for holotype except Mazumbai Forest Reserve, small

stream at 1480 m a.s.l., 24.xi.-4.xii.1990; 1 Â data as

for holotype except Gogoi, 30.xi.1990, sweep net.

Description

Anterior wing length 1.8-2.1 mm. Genitalia as in

figs. 24, 25. Abdominal sternite VIII with paired spi-

nes unequal, stout, elongate and curved and positio-

ned apico-laterally. Tergite X expanded in basal part,

narrowed sub-apically, slightly expanded distally, ap-

ex almost truncate, covered in fine spinules. Inferior

appendages short, about half length of tergite X,

rounded distally. Subgenital plate membranous.

Etymology. — From the Latin cilium, eyelash, and

the suffix -atus, provided with, referring to the fine

spinules on the tergite X.

Biology and distribution. — Known only from the

West Usambara Mountains, from streams and rivers

at altitudes between 1100 m and 1650 m a.s.l. In the

Malaise traps at Kaputu Stream this species was

trapped in highest numbers at sites with moderate to

fast flow over gravel, stones and bedrock near water-

falls (see Andersen & Johanson 1993).

Remarks. — This species is clearly identified by the

stout, unequally curved paired sclerotised spines

which arise mid-laterally.

Ugandatrichia Mosely

Ugandatrichia Mosely, 1939: 36. Type species:

Ugandatrichia minor Mosely, by original designation.

WELLS & ANDERSEN: Tanzanian micro-caddisflies

Ugandatrichia occurs in Africa, South and South-

east Asia and New Guinea. Scott (1967) described the

hydropetric larva of a species from Zimbabwe and

since similar larvae have been collected from the type

locality of the New Guinean species, life in fast flow-

ing waters is probably the usual niche for the genus.

The two new species described here appear to oc-

cur sympatrically in the Kaputu Stream. Only males

are described here as although a number of females

were collected, they could not be assigned with cer-

tainty to species. They closely resemble the holotype

female of U. acuta Mosely, 1939 in having patches of

black androconia on sternite VIII, but differ in gener-

al genitalic shape.

Ugandatrichia tanzaniensis sp. n.

(figs. 26, 28, 29)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 7, 1535 m a.s.l., 2.-6.xi.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

205). — Paratypes: 164 (3 slides) data as for holotype

except loc. 3-11, 30.x.-29.xi.1990; 16 data as for

holotype except loc. 5, 6.xi.1990, sweep net; 1d data

as for holotype except loc. 8, 5.xi.1990, sweep net;

26 data as for holotype except Gologolo, Lushoto,

25.ix.1990, sweep net; 2d data as for holotype except

East Usambara Mts, Bombole, 1.xii.1990, sweep net.

Description

Large, and in alcohol uniformly dark coloured; an-

tennae all damaged; wings (fig. 26) broad, venation

complete; anterior wing length 3.4-3.7 mm. Male ge-

nitalia as in figs. 28, 29. Segment IX short mid dor-

sally and ventrally, produced laterally to form a lobe

that is stout and apically somewhat truncate in lateral

view, apically acute in ventral view. Segment X bro-

adly rounded in lateral view, in dorsal view broadly

bi-concave apically. Inferior appendages divergent, ir-

regularly divided in distal half, with inner sclerotised

spur on ventral lobe, small digitiform lobe mid-dor-

sally and apically sclerotised process more distally.

Subgenital plate in lateral view forming a pair of scle-

rotised lobes. Aedeagus simple, with titillator.

Etymology. — Named for the country Tanzania.

Biology and distribution. — Known both from the

West and East Usambara Mountains, beside rapidly

flowing streams and smaller rivers at altitudes

between 830 m and 1860 m a.s.l. In the Malaise traps

along the Kaputu Stream this species was trapped

most frequently at sites with fast flow over bedrock

and stones, but also at sites with slow flow over grav-

el, stones and mud (see Andersen & Johanson 1993).

Swarming specimens were netted by day mostly near

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

waterfalls or in areas of fast flow over bedrock and

stones.

Remarks. — Both this species and the following, U.

dentata sp. n. group with the other East African

Ugandatrichia species in having tergite IX deeply ex-

cavated dorsally. Ugandatrichia tanzaniensis sp. n.

differs from U. dentata in having sternite IX very

short medially. In this respect it resembles the south-

ern African U. rhodesiensis Scott, 1976, from which it

is distinguished clearly by narrower, longer lateral

lobes on segment IX which appear subquadrate in lat-

eral view.

Ugandatrichia dentata sp. n.

(figs. 27, 30, 31)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 8, 1510 m a.s.l., 5.xi.1990, sweep net,

ZMB’s Tanzania Expedition (slide, ZMBN No. 206). —

Paratypes: 44d (3 slides) data as for holotype except

5.xi. & 6.xi. 1990; 5 data as for holotype except loc.

8, 29.x. & 5.xi. 1990; 156 (1 slides) data as for holo-

type except loc. 5-7, 29.x.-13.xii.1990, Malaise trap;

26 data as for holotype except loc. 7, 4.ii.-8.-

viii.1991, Malaise trap; 84d (5 slides) data as for

holotype except Dule, Bumbuli River, 26.xi.1990;

704 data as for holotype except East Usambara Mts,

Mlesa, 1.x11.1990

Description

Large; body uniformly very dark coloured; anten-

nae dark in proximal third, pale in median section

and dark in distal third; wings broad, anterior wing

length 2.4-4.6 mm. Male genitalia as in figs. 27, 30,

31. Segment IX with deep cleft mid-dorsally and a

pair of small ovoid lobes laterally. Segment X difficult

to discern. Inferior appendages stout, in lateral view

with inner margins almost parallel, closely aligned,

outer margins rounded, with large black spur dorsally

at base on inner margin. Subgenital plate short, roun-

ded apically. Aedeagus simple, titillator present.

Etymology. — From the Latin dentatus, toothed, re-

ferring to the large, black spur on the inner margin of

the inferior appendage.

Biology and distribution. — Taken both in the

West and East Usambara Mountains, beside rapidly

flowing rivers at altitudes between 800 m and 1650 m

asl. In the Malaise traps along the Kaputu Stream

this species was trapped most frequently at a site with

fast flow over gravel and stones above a waterfall (see

Andersen & Johanson 1993). Swarming specimens

were abundant by day mostly near waterfalls or beside

stretches of fast flow over bedrock and stones.

Remarks. — This species groups with Ugandatrichia

156

tanzaniensis sp. n., U. minor Mosely, 1939, U. nigra

Mosely, 1939 and on the basis of similarities berween

females of U. acuta and unassociated females of U. tan-

zaniensis and/or U. dentata sp., n. probably also with

that species. The form of the inferior appendages of U.

dentata is almost identical with that of U. sourya

(Schmid, 1960) from Pakistan. Ugandatrichia dentata

is readily distinguished from U. tanzaniensis by the

short lateral lobes and stouter inferior appendages.

Dhatrichia Mosely

Dhatrichia Mosely, 1948: 78. Type species: Dhatrichia ina-

sa Mosely, by original designation and monotypy.

The genus Dhatrichia was first recorded from the

Yemen and subsequently a second species was de-

scribed from Zaire (Statzner 1977) and a third from

the Ivory Coast (Gibon 1987b). Two new species are

described here and are the first to be recorded from

East Africa. Male genitalia of each are distinctive al-

though conforming in general features with congen-

ers. Ihe two new species were collected together

beside slower reaches of the Kaputu Stream where the

substrata are sand, gravel, some mud and stones.

Dhatrichia divergenta sp. n.

(figs. 33-35)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 7, 1535 m a.s.l., 4.-12.ii.1991, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

208). — Paratypes: 11 4 (9 slides) data as for holotype;

738 (3 slides) data as for holotype except loc. 2-10,

29.x.-29.xi.1990; 16 data as for holotype except

Mazumbai Forest Reserve, small stream at 1480 m

a.s.l., 26.x1.-4.x11.1990.

Description

Small, pale, antennae short, 19-segmented, seg-

ments quadrate; wings as for Dhatrichia cinyra sp. n.,

anterior wing length 1.4-1.8 mm. Genitalia as in figs.

33-35. Abdominal segment IX short mid-dorsally

and ventrally, forming pronounced lobes laterally in

ventral and dorsal views. Abdominal segment X a

short membranous band only. Inferior appendages

short and stout, bilobed and strongly upturned in la-

teral view, the ventral lobe with a slender apical pro-

cess. Subgenital plate membranous, also with a slen-

der projection; the origins of two other lobes seen in

lateral view are unclear.

Etymology. — From the Latin divergens, wide-

spreading, referring to the diverging lobes of the infe-

rior appendages in ventral view.

WELLS & ANDERSEN: Tanzanian micro-caddisflies

Figs. 32, 36-38. Dhatrichia cinyra sp. n., male. 32, wings; 36, genitalia, dorsal view; 37, genitalia, ventral view; 38, genitalia,

lateral view. — Figs. 33-35. Dhatrichia divergenta sp. n., male genitalia. 33, lateral view; 34, dorsal view; 35, ventral view. —

Abbreviations. — ae: aedeagus; inf.app.: inferior appendage; subg.pl.: subgenital plate; VIII, IX, X: abdominal segments VIII,

IX, X.

Biology and distribution. — Known only from

streams in the Mazumbai area in the West Usambara

Mountains, at altitudes between 1420 m and 1650 m

a.s.l. In the Malaise traps along the Kaputu Stream

this species was most abundant at a site with moder-

ate flow over mud, fine sand and larger stones (see

Andersen & Johanson 1993).

Remarks. — As with the next species, Dhatrichia

divergenta sp. n. shows similarities in general form

to other congeners. It can be recognised by the

stouter, more robust appearance of the inferior ap-

pendages.

Dhatrichia cinyra sp. n.

(figs. 32, 36-38)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Stream, loc. 4, 1680 m a.s.l., 1.-6.x1.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

207). — Paratypes: 2514 (8 slides) data as for holo-

type except loc. 2-11, 27.x.-3.xii.1990; 1d data as for

holotype except loc. 7, 4.-12.11.1990; 16 data as for

holotype except Gogoi, 30.xi.1990, sweep net.

Description

Small, uniformly pale fuscous in colour; antennae

short, 19-segmented, segments stout; wings (fig. 32)

slender, tapered to acuminate apices; forewing with

jugal lobe, length 1.7-1.9 mm. Genitalia as in figs.

36-38. Abdominal segment IX well retracted into

VII, with lateral lobes prominent. Segment X for-

ming a short, stout plate, almost truncate apically.

Inferior appendages slender, comprising three lobes,

the ventral-most pair in ventral view in the form of a

harp, being slender and bowed, with slightly out-

turned apices. In lateral view, the subgenital plate ap-

pears to be divided into two lobes, a rounded ventral

lobe and a narrower, upturned dorsal lobe. Aedeagus

straight, tightly constricted medially and tapered dis-

tally to a flared apex, titillator present.

Etymology. — From the Latin cinyra, a kind of

harp, referring to the shape of the ventral-most pair of

lobes of the inferior appendages in ventral view.

Biology and distribution. — Collected along

streams in the West Usambara Mountains, at alti-

tudes between 1100 m and 1650 m a.s.l. In the

Malaise traps along the Kaputu Stream this species

was most abundant at sites with fast flow over gravel,

stones and bedrock (see Andersen & Johanson 1993).

Remarks. — In general form, males of this species

resemble those of Dhatrichia inasa Mosely, 1948, the

type species from Yemen and D. bipunctata Statzner,

1977 from Zaire, but D. cinyra is sp. n. is clearly dis-

tinguished by the shape of the inferior appendages.

Hydroptila Dalman

Hydroptila Dalman, 1819: 125. Type species: Hydroptila ti-

neoides Dalman, by monotypy. See Marshall (1979) for

full generic synonymy.

Several of the Hydroptila species groups recognised

by Marshall (1979) are represented in Africa, particu-

larly in northern Africa. These include the sparsa, ca-

pensis, occulta and pulchricornis groups. Prior to this

work, however, only the widespread Hydroptila crucia-

ta Ulmer, 1912 (with A. hirra Mosely, 1948 in synon-

ymy), in Marshall’s occulta-group, was recorded from

East Africa (see Johanson 1992). A new record is given

here for this species, and five new species are described.

Two of the new species H. usambarensis sp. n. and HH.

morogorensis sp. n., clearly align with sparsa-group spe-

158

cies and H. bumbulensis sp. n. shares features of the oc-

culta-group. The affinities of Hydroptila mazumbaien-

sis sp. n. and H. tannerorum sp. n. are unclear.

Sites at which Hydroptila species were collected

varied from slow reaches of streams to fast riffles.

Hydroptila usambarensis sp. n.

(figs. 39, 40)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Dule, Bumbuli River,

1220 ma.s.l., 26.xi.1990, sweep net, ZMB’s Tanzania

Expedition (slide, ZMBN No. 209). — Paratypes: 56

(slides) data as for holotype; 14 (slides) data as for

holotype except East Usambara Mts, Mesa,

1.xii.1990; 38 data as for holotype except East

Usambara Mts, Bombole, 1.xii.1990.

Description

Post-occipital scent organs on head each compris-

ing a cluster of pale androconia beneath inner side of

post-occipital lobes; antennae 24-segmented; flagellar

segments rectangular, with scattered sensilla placo-

dea. Anterior wing length 1.3-1.8 mm. Genitalia as in

figs. 39, 40. Without lateral lobes of the usual form

on abdominal segment IX, but with apico-lateral ang-

les of abdominal segment IX rounded, setose. Tergite

X membranous, quadrate in dorsal view. Inferior ap-

pendages elongate, in ventral view parallel-sided, api-

cally produced slightly on inner side, in lateral view,

irregular in shape. Subgenital plate subtriangular, pai-

red setae basal. Aedeagus dilated in distal half, a small

spine subapically.

Etymology. — Named after the Usambara Moun-

tains, Tanzania.

Biology and distribution. — Taken from rivers both

in the East and West Usambara Mountains, at sites

with fast flow over large stones at altitudes between

800 m and 1220 m a.s.l.

Remarks. — This is probably a sparsa-group species,

although the form of the lateral lobes differs from

others. Hydroptila usambarensis is distinguished by

the unusual shape of the aedeagus which is dilated

distally anterior to a small, subapical spine.

Hydroptila morogorensis sp. n.

(figs. 41, 42)

Type material. — Holotype d , TANZANIA, Morogoro region,

Morogoro, Sokoine University of Agriculture, 550 m

asl, 26.x-11.xi.1990, light trap, ZMB’s Tanzania

Expedition (slide, ZMBN No. 210).

Description

Post-occipital scent organs on head small; antennae

WELLS & ANDERSEN: Tanzanian micro-caddisflies

MD subg pl.

Figs. 39, 40. Hydroptila usambarensis sp. n., male genitalia. 39, ventral view; 40, lateral view. — Figs. 41, 42. Hydroptila mo-

rogorensis sp. n., male genitalia. 41, lateral view; 42, ventral view. — Figs. 43-45. Hydroptila mazumbaiensis sp. n., male geni-

talia. 43, ventral view; 44, dorsal view; 45, lateral view. — Figs. 46, 47. Hydroptila tannerorum sp. n., male genitalia. 46, ven-

tral view; 47, lateral view. — Abbreviations. — ae: aedeagus; inf.app.: inferior appendage; subg.pl.: subgenital plate.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

25-segmented; flagellar segments with sparse scatte-

red sensilla placodea. Anterior wing length 1.4 mm.

Genitalia as in figs. 41, 42. Lateral lobes on abdomi-

nal segment IX rounded, setose. Tergite X mem-

branous, in lateral view stout, rounded apically.

Inferior appendages elongate, irregular in shape,

swollen near base, tapered and turned outwards to-

wards apex in ventral view; in lateral view swollen

subapically, upturned distally. Subgenital plate

broad, membranous, apically acute in ventral view, a

triangular medial projection in lateral view. Aedeagus

slender with a small apical spine.

Etymology. — Named after the city of Morogoro,

Tanzania.

Biology and distribution. — Known only from the

type locality on the campus of Sokoine University of

Agriculture in Morogoro, in an area with several

small, slow-flowing streams and ditches.

Remarks. — This is a sparsa-group species showing

general resemblance to H. usambarensis sp. n., but

differing in detailed shape of genitalic structures, par-

ticularly in having the aedeagus simple and slender,

and inferior appendages in ventral view slightly ex-

panded proximally, slender and out-turned distally.

Hydroptila mazumbaiensis sp. n.

(figs. 43-45)

Type material. — Holotype d, TANZANIA, Tanga re-

gion, West Usambara Mts, Mazumbai, Kaputu Stream,

loc. 7, 1535 m a.s.l., 4.-12.11.1991, Malaise trap, ZMB’s

Tanzania Expedition (slide, ZMBN No. 211). —

Paratypes: 1d (slide) data as for holotype; 25 data as

for holotype except loc. 5-10, 30.x.-26.xii.1990.

Description

Post-occipital scent organs on head small, densely co-

vered with setae and in the form of a balloon-like struc-

ture with a central duct which appears to open ventral-

ly, without discernible androconia; antennae damaged;

flagellar segments with sparse sensilla placodea on apical

rim, elsewhere scattered sensilla auricillica. Meso-

scutellum diamond-shaped. Anterior wing length 2.2

mm. Genitalia as in figs. 43-45. Lateral lobes on abdo-

minal segment IX rounded apically, membranous.

Tergite X broad, rounded laterally and in lateral view

produced and upturned apico-laterally. Inferior appen-

dages slender, tapered, apically with a pale spur on inner

margin. Subgenital plate broad in ventral view, paired

setae on apical margin, in lateral view with a median

spur ventrally. Aedeagus with slender apical and subapi-

cal spines, swollen below each spine.

Etymology. — Named after the village of

Mazumbai, West Usambara Mountains, Tanzania.

160

Biology and distribution. — Known only from the

Kaputu Stream in the West Usambara Mountains,

where it was taken between 1420 m and 1650 m a.s.l.,

at sites with slow to fast flow over mud, sand, gravel

and larger stones (see Andersen & Johanson 1993).

Remarks. — The affinities of this species are un-

clear. The ventral spur on the subgenital plate sug-

gests that it may be an occulta-group species. The

form of the aedeagus is distinctive in having distally a

pair of flange-like structures in series.

Hydroptila tannerorum sp. n.

(figs. 46, 47)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 7, 1535 m a.s.l., 4.-12.11.1991, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

212). — Paratypes: 4d (slides) data as for holotype;

75 data as for holotype except loc. 4-8, 30.x.-

13.x11.1990.

Description

Post-occipital scent organs on head small, in the

form of a cluster of black androconia; antennae da-

maged. Anterior wing length 1.9-2.0 mm. Genitalia

as in figs. 46, 47. Abdominal sternite VII with a small

acute mesal process. Abdominal segment IX produ-

ced apico-laterally into broadly rounded lobes.

Tergite X membranous, broad-based, tapered distally

to truncate apex. Inferior appendages short, fused

mesally, forming rounded lobes laterally in ventral

view, in lateral view with a hooked process dorsally.

Subgenital plate wide in ventral view, with paired

setae at sides of a slender apico-mesal cleft. Aedeagus

slender, a short curved spine subapically.

Etymology. — Named after Mrs and Mr Tanner,

who initiated the establishment of the Mazumbai

Forest Reserve.

Biology and distribution. — Collected from the

Kaputu Stream, West Usambara Mountains only,

from stream sites with slow to moderate flow over

sand, gravel and stones and some mud at altitudes

between 1510 m to 1680 m a.s.l. (see Andersen &

Johanson 1993).

Remarks. — Males of this species have the genitalic

structures, particularly the inferior appendages, re-

duced and the antero-lateral angles of segment IX

strongly produced posteriorly. The affinities of this

species are unknown.

Hydroptila cruciata Ulmer

Hydroptila cruciata Ulmer, 1912: 83.

Hydroptila hirra Mosely, 1948: 81; Malicky, 1986: 234.

Biology and distribution. — In Africa this species is

found in Cape Verde, Benin, Guinea, the Ivory

Coast, Niger, Tanzania, Togo, Transvaal and it also

occurs in Yemen and Palestine (see Malicky 1986,

Gibon 1987b). Aspects of the life history were de-

scribed by Botosaneanu & Guidicelli (1981). The

present specimens were taken in a light trap in an ar-

ea with several small slow-flowing streams and ditch-

es.

Remarks. — Hydroptila cruciata is widely distribut-

ed and can be recognised amongst congeners by the

long strap-like spines that intersect mid-ventrally in

the male genitalia. Another prominent feature of the

male is the shape of the post-occipital scent organs

which are large, subrectangular and occupy about one

third of the dorsal head.

Material examined. —8 4 (4 slides), 99, TANZANIA,

Morogoro region, Morogoro, Sokoine University of

Agriculture, 18.x.-11.xi.1990, light trap, ZMB’s

Tanzania Expedition.

Hydroptila bumbulensis sp. n.

(figs. 48, 49)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Dule, Bumbuli River,

1220 m a.s.l., 26.xi.1990, sweep net, ZMB’s Tanzania

Expedition (slide, ZMBN No. 213).

Description

Post-occipital caps on head, angular and covering

small tufts of scent hairs; antennae 31-segmented; fla-

gellar segments sub-quadrate with dense scattered

sensilla placodea. Wings slender, elongate; anterior

wing length 2.1 mm. Genitalia as in figs. 48, 49.

Lateral lobes on abdominal segment IX short, mem-

branous. Tergite X broad, membranous. Inferior ap-

pendages in ventral view stout, about twice as long as

greatest width, apico-ventrally with a darkly scleroti-

sed process; in lateral view, inferior appendages great-

ly expanded dorsally in distal half. Below the subgeni-

tal plate, and dorsal to the inferior appendages is a

pair of sclerotised processes which lie across each ot-

her. Subgenital plate narrow medially, with the usual

paired apical seta situated on each side of a small me-

dian cleft. At the base of and between the inferior ap-

pendages is a short, slender, sclerotised median pro-

cess. Aedeagus simple.

Etymology. — Named after the River Bumbuli in

the West Usambara Mountains, Tanzania.

Biology and distribution. — Known only from the

type locality, the Bumbuli River at Dule in the West

Usambara Mountains. At the collecting site the river

was fast flowing over large stones.

Remarks. — This is an occulta-group species closely

WELLS & ANDERSEN: Tanzanian micro-caddisflies

resembling Hydroptila cruciata and the Malaysian

species, H. berkait Wells & Huisman, 1992, both in

the occulta-group. It differs from H. cruciata in the

form of the inferior appendages, and the parameres.

Tangatrichia gen. n.

Type species. — Tangatrichia gracilenta sp. n., male,

by present designation.

Diagnostic characters. — Showing strong resem-

blance to the Stactobiini taxa in the form of wings,

and male genitalia, especially the elongate internal ap-

odemes and form of the aedeagus, but groups with

the Hydroptilini genera on basis of presence of post-

occipital lobes on head and absence of suture on me-

soscutellum. In general form of male genitalia resem-

bling Hydroptila, but distinguished by presence of

ocelli, and slender, elongate apodemes on abdominal

segment IX; and by scattered arrangement of vestiture

and absence of sensilla placodea on antennal seg-

ments.

Etymology. — From Tanga region of Tanzania re-

ferring to the type locality, and the Greek trichos, hair.

Description

Adult. Wings (fig. 54) attenuate, with strongly re-

duced venation; forewing with jugal lobe. Head with

3 ocelli; tentoria, including posterior bridge, comple-

te; post-occipital lobes on head, but no associated

scent glands; antennae with segments elongate, clot-

hing hair scattered and sensory structures sensilla au-

ricillica only. Thorax (fig. 50) with mesoscutellum

sub-rectangular, without suture; metascutellum trian-

gular. Tibial spur formula 0, 2, 4.

Genitalia resembling some Stactobiini taxa in gen-

eral form, in having long internal apodemes on ab-

dominal segment IX, but with basic arrangement of

genitalic structures as in Hydroptila. The aedeagus is

slender and straight, with a short straight titillator.

Remarks. — This genus is erected for a single species

that groups with Hydroptila on the basis of presence

of jugal lobe on forewing, form of tentorium, anten-

nal segments, shape of thoracic scutellae, and of male

genitalic structures in general, but not presence of oc-

elli and shape of abdominal segment IX. In these lat-

ter respects, and in having only sensilla auricillica on

the antennal segments this genus shows similarity to

the Australian/Malaysian genus Jabitrichia Wells,

1990 (Wells 1990c, O'Connor & Ashe 1992), from

which it is distinguished by differences in male geni-

talic structures and arrangement of hair on antennal

segments.

The one species referred to this genus was collected

only in the West Usambara Mountains, from streams

161

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

Figs. 48, 49. Hydroptila bumbulensis sp. n., male genitalia. 48, ventral view; 49, lateral view. — Figs. 50-54. Tangatrichia grac-

ilenta gen. n., sp. n., male. 50, head and thorax, dorsal view; 51, genitalia, lateral view; 52, genitalia, ventral view; 53, aedea-

gus; 54, wings. — Abbreviations. — inf.app.: inferior appendage; mes.sc.: mesoscutellum; met.sc.: metascutellum; p-o.l.: post-

occipital lobes; subg.pl.: subgenital plate.

with slow to fast flow over stones, gravel, sand and

mud.

Tangatrichia gracilenta sp. n.

(figs. 50-54)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 5, 1650 m a.s.l., 2.-6.x1.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

214). Paratypes: 51d (8 slides), data as for holotype

except loc. 1-10, 30.x.-13.xii.1990; 1354 (some

162

damaged) data as for holotype except loc. 7, 4.i.-

8.vi.1991; 3d (2 slides) data as for holotype except

Mazumbai Forest Reserve, small stream at 1480 m

a.s.l., 26.xi.-4.x11.1990; 3d (2 slides) data as for holo-

type except loc. 8, 5.xi.1990, sweep net.

Description

In general features as for genus. Small, pale, wings

as in fig. 54, anterior wing length 1.5-1.6 mm.

Genitalia as in figs. 51-53. Abdominal segment IX

short mid-ventrally, antero-lateral apodemes long and

slender. Segment X membranous, bifid distally.

Inferior appendages simple, elongate rectangular in

ventral view, well separated medially. Subgenital plate

narrowed abruptly in distal half, paired setae apically.

Aedeagus long, slender, slightly twisted distally, with

a short titillator arising at one-third length, but not

twisted as is more usual in hydroptilids.

Female unknown.

Etymology. — From the Latin gracilis, slender, gra-

cile, referring to the general shape of the species.

Biology and distribution. — Collected in the

Mazumbai area in the West Usambara Mountains

only, from streams with slow to fast flow between

1420 m to 1770 m a.s.l. In the Malaise traps at the

Kaputu Stream this species was most abundant at a

site with slow flow over mud, sand, gravel and stones

(see Andersen & Johanson 1993).

Remarks. — See generic description.

Orthotrichia Eaton

Orthotrichia Eaton, 1873: 141. Type species:

Hydroptila angustella McLachlan, 1865 by original

designation. For full generic synonymy see Marshall

(1979).

Orthotrichia is a diverse genus in tropical Africa.

This work adds four new species to the previous 16

described from Africa. Most species are recorded

from Central Africa, while two are listed in the South

African fauna (de Moor 1993), two others are known

from East Africa (Johanson 1992), including O. strae-

leni Jacquemart, 1956 which is recorded from

Tanzania, as well as Uganda and Zaire, and one from

Mali in West Africa (Marlier 1978). Marshall (1979)

included the African species in the angustella- and cos-

talis- species groups.

The South African species, Orthotrichia barnardi

Scott, 1963, is here recorded from Tanzania for the

first time. One of the four new species, O. bisetula sp.

n., differs only slightly from O. barnardi and the

Central African O. kalengiensis Statzner, 1977 and

another, O. nigrovillosa sp. n., groups with O. straele-

ni, O. spinicauda Kimmins, 1958, from Zimbabwe

and O. nova Marlier, 1978, from Mali. A third, O.

scutellata sp. n., is distinctive and not aligned closely

with any of the other species, although conforming

with congeners. The fourth new species is enigmatic

and is referred tentatively to Orthotrichia as O. hy-

droptiloides sp. n.

Orthotrichia bisetula sp. n.

(figs. 55, 56)

Type material. — Holotype d, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

WELLS & ANDERSEN: Tanzanian micro-caddisflies

Stream, loc. 10, 1420 m a.s.l., 20.-26.x1.1990,

Malaise trap, ZMB’s Tanzania Expedition (slide,

ZMBN No. 215). — Paratypes: 19d (3 slides) data as

for holotype except loc. 10-11, 14.xi.-3.xii.1990; 24

(slides) data as for holotype except Shakoi River,

24.xi.1990, sweep net.

Description

Antennae 29-segmented; flagellar segments elonga-

te, with dense sensilla placodea and at least one sensory

pit per segment. Anterior wing length 2.4-2.8 mm.

Genitalia as in figs. 55, 56. Abdominal segment IX

with a pair of unequal, membranous, lateral digitate

processes, each with a pair of setae apically. Tergite X

broadly rounded apically, not sclerotised, covered with

tiny spinules. Aedeagus of typical elongate slender

form. Inferior appendages asymmetrical, left shorter

than right, its inner apical angle produced; dorsal pro-

cess V-shaped, anterior apodeme slender, elongate. A

pair of unequal sclerotised spines dorso-laterally.

Etymology. — From the Latin bis, two and setula,

diminutive for bristle, to describe the paired setae on

the digitiform process in the male genitalia.

Biology and distribution. — Collected from streams

and rivers with slow to rapid flow in the West

Usambara Mountains, between 1400 m and 1420 m

a.s.l. In the Malaise traps at the Kaputu Stream this

species was only taken at the lowermost sites where

the stream is moderate to slow flowing over mud,

sand, gravel and stones (see Andersen & Johanson

1993):

Remarks. — This species, Orthotrichia barnardi and

O. kalengiensis, all have lateral digitate processes on

segment IX. They are distinguished by differences in

the shape of the inferior appendages and lateral

spines.

Orthotrichia barnardi Scott

(fig. 57)

Orthotrichia barnardi Scott, 1963: 470.

Biology and distribution. — Known from South

Africa and northern Tanzania. The present specimens

were taken at slow to fast flowing streams and rivers,

at altitudes between 1220 m and 1400 m a.s.l.

Remarks. — This species closely resembles

Orthotrichia bisetula sp. n. and O. kalengiensis. It is

distinguished by the nearly symmetrical inferior ap-

pendages, see fig. 57.

Material examined. — 14, TANZANIA, Tanga re-

gion, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 11, 13.-14.xi.1990, Malaise trap, ZMB’s

Tanzania Expedition; 14 (slide) as previous except

Dule, Bumbuli River, 26.x1.1990, sweep net.

163

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

164

Orthotrichia scutellata sp. n.

(figs. 58, 59)

Type material. — Holotype 4, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 5, 1650 m a.s.l., 6.-12.xi.1990, Malaise

trap, ZMB’s Tanzania Expedition (slide, ZMBN No.

216).

Description

Antennae damaged. Anterior wing length 2.4 mm.

Genitalia as in figs. 58, 59. Abdominal segment IX

without membranous lateral digitate processes, but

with a pair of unequal sclerotised spines. Tergite X re-

duced to a stout spine. Aedeagus of typical elongate

slender form. Inferior appendages fused, rectangular,

with a seta at each apical angle and with a pair of thin

sclerotised lobes dorsally; dorsal process also fused,

quadrate, setae at apical angles; anterior apodeme

slender, elongate. A pair of unequal, sclerotised spines

laterally, the left slender and curved, the right broader

and hooked distally. Internally a pair of short, slender

parameres.

Etymology. — From the Latin scutella, little flat

dish, plate, and the suffix -atus, provided with, refer-

ring to the fused dorsal process of the inferior ap-

pendages, forming a single plate.

Biology and distribution. — Known only from the

type locality, Kaputu Stream in the West Usambara

Mountains, where it was taken at a site where the

stream is fast flowing over gravel and stones (see

Andersen & Johanson 1993).

Remarks. — This is a distinctive species, character-

ised by the fused inferior appendages and their dorsal

process in the form of a simple plate.

Orthotrichia nigrovillosa sp. n.

(figs. 60, 61)

Type material. — Holotype 4, TANZANIA, Tanga

region, West Usambara Mts, Mazumbai, Kaputu

Stream, loc. 10, 1420 m asl, 29.x1.-3.xii. 1990,

Malaise trap, ZMB’s Tanzania Expedition (slide,

ZMBN No. 217). — Paratypes: 4d (1 slide) data as for

holotype except loc. 4-11, 12.-29.xi.1990.

Description

Antennae damaged; flagellar segments elongate,

with dense scattered sensilla placodea and one sensory

WELLS & ANDERSEN: Tanzanian micro-caddisflies

pit per segment. Anterior wing length 2.3-2.4 mm.

Genitalia as in figs. 60, 61. Abdominal sternite VIII

with a mesal process bearing blunt, dark setae.

Abdominal segment IX without lateral digitate pro-

cesses. Tergite X sclerotised, distally in the form of

stout hooks and spines. Aedeagus of typical elongate

slender form. Inferior appendages superficially sym-

metrical save for a small, square sclerotised process

apically on the left inferior appendage; dorsal process

U-shaped, well displaced to right side; anterior apo-

deme slender, elongate. A pair of unequal, sclerotised

processes laterally, the left slender, with long setae dis-

tally, the right with curved setate process basally and

sclerotised spine distally.

Etymology. — From the Latin niger, black, dark,

and villus, tuft of hair, referring to the dark, blunt se-

tae on the mesal process of abdominal segment VII.

Biology and distribution. — Known only from the

Kaputu Stream in the West Usambara Mountains, at

sites between 1400 m and 1680 m a.s.l. with slow to

moderate flow over mud, sand, gravel and stones (see

Andersen & Johanson 1993).

Remarks. — Orthotrichia nigrovillosa sp. n., O. spir-

alina Statzner, 1977, O. straeleni, O. spinicauda and

O. nova all have a more complex development of lat-

eral processes on segment IX than the above men-

tioned group of species. In O. nigrovillosa this takes

the form of an elongate lobe with setae distally on the

outer side on the left and on the right a similar struc-

ture basally, extended dorsally in a sclerotised spine.

The shape of the inferior appendages differs between

species.

Orthotrichia hydroptiloides sp. n.

(figs. 62, 63)

Type material. — Holotype d, TANZANIA, Moro-

goro region, Morogoro, Sokoine University of Agri-

culture, 550 m a.s.l., 11.xi.1990, light trap, ZMB’s

Tanzania Expedition (slide, ZMBN No. 218).

Description

Ocelli absent. Tibial spurs 0, 2, 4. Meso- and me-

tascutellum subrectangular, metascutellum about half

length of mesoscutellum. Antennae damaged, remai-

ning flagellar segments with scattered clothing hair,

basal whorl of hair and sensilla placodea lacking.

Anterior wing length 1.4 mm. Genitalia as in figs. 62,

63. Abdominal segment IX broad anteriorly, without

Figs. 55, 56. Orthotrichia bisetula sp. n., male genitalia. 55, ventral view; 56, lateral view. — Fig. 57. Orthotrichia barnardi

Scott, 1963, male genitalia, ventral view. — Figs. 58, 59. Orthotrichia scutellata sp. n., male genitalia. 58, lateral view; 59, ven-

tral view. — Figs. 60, 61. Orthotrichia nigrovillosa sp. n., male genitalia. 60, lateral view; 61, ventral view. — Figs. 62, 63.

Orthotrichia hydroptiloides sp. n., male genitalia. 62, ventral view; 63, lateral view. Abbreviations. — d.po.inf.: dorsal process

of inferior appendage.

165

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 138, 1995

spines or digitate processes. Tergite X membranous,

broad. Aedeagus without spiral titillator, swollen and

twisted distally, with divided apex. Inferior appenda-

ges symmetrical, stout, fused basally, in lateral view

expanded distally and broadly rounded apically; dor-

sal process fused, more or less T-shaped, with a setae

laterally on the right side of base; anterior apodeme

slender, elongate and curved.

Etymology. — Named after the genus Hydroptila,

using the suffix -oides, resembling, referring to the

general shape of the male genitalia.

Biology and distribution. — Known only from the

type locality on the campus of Sokoine University of

Agriculture in Morogoro in an area with several

small, slow-flowing streams and ditches.

Remarks. — This species is placed in Orthotrichia

on the basis of general head, thoracic and wing fea-

tures and the form of the male genitalic structures.

The tibial spur formula is aberrant, however, and in

the male genitalia the aedeagus is quite unlike the typ-

ical and generally invariant form, and the anterior,

internal shape of abdominal segment IX is unusual.

The spur formula conforms with that of Hydroptila

and in general appearance the male genitalia resemble

those of H. cortensis Mosely, 1937. However, the

wing lacks a jugal lobe, the thoracic scutellae conform

with those of Orthotrichia and upon close inspection

the genitalia certainly show features typical of

Orthotrichia, with the characteristic form of the infe-

rior appendages, their dorsal process and basal apo-

deme.

Close examination of species of Tricholeiochiton

Kloet & Hincks, 1944 reveals a basal apodeme simi-

lar to, but less distinctive than that of males of most

species in Orthotrichia. The aedeagus of O. hydropti-

loides sp. n. is dissimilar to that of other Orthotrichia,

but shows some resemblance to the form typical of

Tricholeiochiton. However, the arrangement of hair

on the antennal segments appears to be of the form

found in Orthotrichia.

ACKNOWLEDGEMENTS

The expedition to Tanzania was funded by The

Norwegian Council for Science and the Humanities

(NAVE). Thanks are due to the staff at the Department

for Forest Biology, Sokoine University of Agriculture

(SUA), Morogoro, Tanzania and to the Norwegian

Agency for Development (NORAD), Dar es Salam,

Tanzania, for their cooperation and support. We also

want to thank the Australian Biological Resources

Study (ABRS) and CSIRO Division of Entomology,

Canberra, Australia for making available facilities for

the laboratory and word-processing aspects of this

study. Robert Scougall, Drafting Office, Australian

166

Nature Conservation Agency very kindly prepared

the bromides from the line drawings.

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Received: 3 August 1994

Revised version accepted: 9 January 1995

167

Tijdschrift voor Entomologie

Volume 138, no. I

Articles

| A. J. de Boer

The taxonomy and biogeography of the cicada genus Papuapsaltria gen. n.

(Homoptera, Tibicinidae).

45 P. E. Bragg

A review of the subfamily Korinninae (Phasmida: Pseudophasmatidae), with the

description of a new species.

51 N. Moller Andersen & P. P. Chen

A taxonomic revision of the ptilomerine genus Rhyacobates Esaki (Hemiptera:

Gerridae), with five new species from China and adjacent countries.

69 N. Nieser

Nine new species of Pseudovelia and a new Xiphovelia (Heteroptera: Veliidae)

from Sulawesi (Indonesia) and Mindanao (Philippines). Notes on Malesian

aquatic and semiaquatic bugs (Heteroptera), V.

89 J. D. Oswald

Revision of the southeast Asian silky lacewing genus Balmes (Neuroptera:

Psychopsidae).

103 L. M. Roth

Revision of the cockroach genus Homopteroidea Shelford (Blattaria,

Polyphagidae)

117 L. M. Roth

Description of a new species of Ctenoneura Hanitsch from Sabah (Blattaria,

Polyphagidae)

121 R. T. Simon Thomas

New and rare Sphecidae (Hymenoptera) from West Africa

131 J. van Tol & Y. Norma-Rashid

The genus Euphaea Rambur in Borneo (Odonata: Euphaeidae). Descriptions

and records of Malesian Odonata, 3.

143 A. Wells & T. Andersen

Tanzanian micro-caddisflies (Trichoptera: Hydroptilidae)

Announcements and book reviews

142 J. T. Wiebes, The Indo-Australian Agaoninae (pollinators of figs). [J. van Tol]

Published 15 June 1995 ISSN 0040-7496

VOIUME 150, NO. 4, 1775 pr ISSN 0040-7496

Tijdschrift

voor

Entomologie

A journal of systematic and evolutionary

entomology since 1858

Published by the Nederlandse Entomologische Vereniging

Tijdschrift voor Entomologie

À journal of systematic and evolutionary entomology since 1858