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volume 156, no. 1, 1775 ISSN 0040-7496

Tijdschrift
voor >
Entomologie

A journal of systematic and evolutionary
entomology since 1858

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Published by the Nederlandse Entomologische Vereniging: _ ze

Tijdschrift voor Entomologie

A journal of systematic and evolutionary entomology since 1858

Scope

The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long
tradition in the publication of original papers on insect taxonomy and systematics.
The editors particularly invite papers on the insect fauna of the Palaearctic and
Indo-Australian regions, especially those including evolutionary aspects e.g.
phylogeny and biogeography, or ethology and ecology as far as meaningful for
insect taxonomy. Authors wishing to submit papers on disciplines related to
taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied
entomology, are requested to contact the editorial board before submitting.
Usually, such papers will only be published when space allows.

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

Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)

Advisory board
M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff).

The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
Amsterdam.

Editorial address
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Correspondence regarding membership of the society, subscriptions and
possibilities for exchange of this journal should be addressed to:

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Special rate for members of the society. Please enquire.

Instructions to authors
Published with index of volume 135 (1992).

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

DISTRIBUTION AND ECOLOGY OF HYDRAENA

]. G. M. CUPPEN ee
RARY
Department of Nature Conservation, Wageningen Agricultural University

AUG 0 2 1993

KUGELANN IN THE NETHERLANDS (COLEOPTERWIVERS/ Ty

HYDRAENIDAE)

Cuppen, J. G. M., 1993. Distribution and ecology of Hydraena in The Netherlands. —
Tijdschrift voor Entomologie 136: 1-10, figs. 1-16, tables 1-5. [Issn 0040-7496]. Published
1 July 1993.

The distribution patterns of the species of the genus Hydraena in The Netherlands are figured.
Their habitats are briefly described and compared with references. Physical/chemical data, phe-
nologies and co-existence of species, based on observations by the author, are presented.
Hydraena species mainly living in stagnant waters have relatively wide distributional areas in
The Netherlands. Hydraena species from running waters are confined to the most eastern and
southern part of the country and some of them may have disappeared at the beginning of this
century. Chlorinity explains best the different distribution patterns of the stagnant water
Hydraena. All stagnant water species can co-exist and their phenology is more or less the same
with a high (early) spring maximum and most often a (lower) autumn maximum, indicating an
univoltine life cycle.

J. G. M. Cuppen, Department of Nature Conservation, Wageningen Agricultural University,
Ritzema Bosweg 32a, 6703 AZ Wageningen, The Netherlands.

Key words. — Hydraena; distribution maps; ecology; co-existence; The Netherlands.

Beetles of the genus Hydraena Kugelann are small
water beetles (1.5-2.5 mm) characterised by parallel
sides, club-shaped antennae and very long maxillary
palps. Sexes can be distinguished easily as males have
six visible abdominal sternites, while females have
seven. Many species show secondary sexual differen-
ces, which sometimes offer useful characters for the
identification of the species. Many species, however,
can only be reliably identified in the male sex by their
genitalia. In the past the study of genitalia has been
neglected in The Netherlands owing to the lack of il-
lustrated (Dutch) keys and by the former practice of
relying entirely on external characters. An illustrated
key to the Dutch species is now available (Cuppen
1992) to which one is referred for identification.

The present survey is the first covering all species of
Hydraena in The Netherlands and gives distribution
maps based on old records, which have been checked
in relation with taxonomic uncertainties, and recent-
ly collected material. A survey of phenology and ecol-
ogy is presented on the basis of literature and own
field-work on mainly the stagnant water Hydraena.

MATERIAL AND METHODS

The material of this study includes the collections

‘Mededeling E. I. S. Nederland, no. 67

of the National Museum of Natural History, Leiden
(RMNH), the Zoological Museum, Amsterdam (ZMA),
the Departments of Entomology and Nature
Conservation of the Wageningen Agricultural
University, and the private collections of H. Cuppen
(Apeldoorn), B. Drost (Wadenoyen), G. van Ee
(Haarlem), J. Huijbregts (Leidschendam), B. van
Maanen (Wageningen), H. Vallenduuk (Boxtel), C.
Visser (Wageningen), B. van Vondel (Hendrik-Ido-
Ambacht), O. Vorst (Utrecht) and the author. Some
data from the Provincial Water Authorities of Noord-
Holland and Utrecht, and from the Research
Institute for Nature Management are also included.
The maps are based on specimens, identified by the
author. For Hydraena riparia Kugelann, Hydraena as-
similis Rey and H. melas Dalla Torre in the H. ripa-
ria-complex (see Jäch 1988), the maps are based on
records of males only.

The distribution of the Hydraena species in The
Netherlands is plotted in the 10 km-squares of the
UTM-grid (figs. 1-12) in which small dots refer to rec-
ords before 1950 and large dots to records since 1950.

On most localities visited by the author a water
sample has been taken, which has been analysed the
same or the next day on pH, electrical conductivity,
chlorinity and total hardness. The results, viz. the
ranges, 90% ranges and median values are presented
in tables 1-4.

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-4. Distribution of Hydraena species in the Netherlands. — 1, Hydraena testacea Curtis; 2, Hydraena palustris Erichson;
3, Hydraena britteni Joy; 4, Hydraena riparia Kugelann (males only).

Data on phenology and ecology are mainly based
on the authors observations from The Netherlands
and are compared with data from literature. Owing to
the paucity of records for most species, statistical

treatment of ecological data was not possible. For
some species only old records are available and there
is little information concerning ecology as labelling
was insufficient in the past.

HABITAT AND DISTRIBUTION

All Dutch Hydraena species are widely distributed
in Europe (lenistea 1978, Hebauer 1980, Horion
1949, Berthélemy 1986, d’Orchymont 1925, Jäch
1988). Detailed distribution maps for Hydraena,
using any grid system, are given by Nilsson (1984) for
the northern part of Sweden (50 km-squares) and
Foster (1990) for the British Isles (10 km-squares).

Relatively small on an European scale are the distri-
bution areas of H. assimilis, H. melas, H. belgica
d’Orchymont and A. excisa Kiesenwetter. The Dutch
localities of H. excisa form an extreme western out-
post of this eastern European species.

Running waters including spring-fed streams are
confined to the southern, central and eastern part of
The Netherlands and therefore many rheophilic spe-
cies of Hydraena are restricted to these parts of the
country. Stagnant waters, however, occur all over the
country. Brackish waters are mainly found in the
south-western and northern part of the country (and
before 1950 also around the Zuiderzee). The sandy
soils in the southern, central and eastern part of the
country, and the coastal dune area have usually a
chlorinity of less than 100 mg/l, while the clay soils in
the western and northern part have a chlorinity of
more than 50 mg/l and gradually pass into the brack-
ish areas. The main soil types per 10 km UTM grid-
square are presented by Van Tol (1981).

NOTES PER SPECIES

H. testacea Curtis

(fig. 1).

This species is recorded from 89 squares since 1950
and additionally from 10 squares before 1950. It is
the commonest species of the genus Hydraena in The
Netherlands, collected mainly in the northern, east-
ern and southern part of the country. From the west-
ern part of the country there is only one recent
record, apart from some pre 1950 records, despite
large sampling programmes in recent years. The lack
of records from Flevoland is probably due to under-
sampling. South-Limburg apparently has no proper
habitats for this species.

Most records of H. testacea in the present investiga-
tion are from temporary, semi-permanent and per-
manent stagnant waters with a well developed margi-
nal vegetation dominated by Carex, Glyceria maxima
(Hartman) Holmberg, Phragmites australis (Cav.)
Steudel, Calamagrostis canescens (Weber) Roth or
Juncus effusus L. Certainly the marginal vegetation is
not a necessity as the species was frequently found in
waters without any marginal vegetation or with only
a submerged vegetation. Also, in slowly running wa-

Cuppen: Hydraena in The Netherlands

ters such as regulated streams, H. testacea occurs fre-
quently. In fast running waters the species is-extreme-
ly rare in The Netherlands. This species is generally
mentioned from stagnant waters, often with a well
developed marginal vegetation as well as slowly run-
ning wa.crs (d’Orch,.nont 1925; Horion 1949;
Hrbacek 1951; Balfour-Browne 1958). Derenne
(1952) mentions its occurrence in slowly running wa-
ters dominated by algae. The occurrence in fast run-
ning waters is only mentioned by Cuppen (1985) and
Horion (1949) in Fontinalis in clear mountain-
brooks. The habitat investigations by the author fair-
ly well agree with literature for this species.

H. palustris Erichson
(fig. 2).

This is an uncommon species recorded from 35
squares since 1950 and additionally from two squares
before 1950. The general distribution pattern of H.
palustris resembles that of H. testacea, but with much
larger gaps and some isolated populations in the
coastal dune area. The only area with a more or less
continuous cover of squares is formed by the fresh
water part of the river district.

With one exception (in a regulated stream), all
Dutch records of H. palustris are from temporary,
semi-permanent and permanent stagnant waters. The
species prefers temporary and semi-permanent, over-
grown ditches and pools with Carex, Phragmites aus-
tralis, Phalaris arundinacea L., Calamagrostis canescens
and Glyceria maxima, in which the soil is covered by
organic debris of the dominant plants. Most localities
are unshaded or only partly shaded by Salix. In per-
manent ponds, ditches and canals H. palustris has
been found only where a well developed marginal
vegetation of helophytes is present. Generally H. pa-
lustris is considered to be a species of stagnant waters
(d’Orchymont 1925; Derenne 1952), which are cov-
ered by aquatic macrophytes (Horion 1949; Hebauer
1980) or mosses (Balfour-Browne 1958) as forest-
pools, fens, ponds and ditches. The rare occurrence in
running water is only mentioned by Horion (1949)
and Hebauer (1980). Hebauer (1980) mentions a
preference for acid waters, which could not be con-
firmed in the present investigations.

H. britteni Joy
(fig. 3).

This species was recorded from The Netherlands
from 18 squares in 1982 by Cuppen & Cuppen. At
present this rather common species is known from 73
squares since 1950 and additionally. from three
squares before 1950. The low number of specimens
in museum collections in comparison with other
Hydraena species is very remarkable. The large in-
crease of the number of records is caused by specific

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

sampling in autumn, winter and early spring. H. brit
teni is confined to the southern and eastern part of the
country with one old record from the province of
Groningen (Haren; coll. ZMA). H. britteni fails in
South-Limburg and occurs locally in the river area be-
hind dikes of the rivers Rhine and Waal (Drost
1989). Records of H. nigrita by Everts (1898; 1922)
belong to this species.

H. britteni is considered to inhabit smaller stagnant
waters, such as vegetation-rich pools and ditches, of-
ten rich in mosses or with decaying leaves (Hrbacek
1951; Derenne 1952; Hebauer 1980; Nilsson 1984).
Dutch habitats of H. britteni are characterized by
Cuppen & Cuppen (1982) as temporary, weakly
acid, stagnant waters with peaty soils covered by
coarse organic debris. Such habitats are alder-brooks,
forest-drains and -pools, marshes and swampy ditch-
es dominated by Phragmites australis, Phalaris arundi-
nacea, Calamagrostis canescens, Glyceria maxima or
Carex. The acidophilic character of the habitat of H.
britteni mentioned by Hebauer (1980) is confirmed
above. The occurrence in a variety of running waters
(Hebauer 1980; Balfour-Browne 1958), especially
those with grassy edges in partial shade (Foster 1990)
and cold springs (Nilsson 1984) is only confirmed for

small sluggish streams receiving seepage-water. The
incidental records in The Netherlands in larger
streams most probably concern specimens that are
washed out from their normal habitats during high
water or flyers from habitats which have been dried
out during the summer. There are some records of H.
britteni, sifted from litter, in the summer period in
dried out habitats (Vorst, pers. comm.). Landin
(1976) mentions the presence of A. britteni and lar-
vae of Hydraena above the water line.

H. riparia Kugelann
(fig. 4).

After the splitting up of H. riparia and H. assimilis
by Jäch (1988), a comparison between literature and
the present data is difficult. H. riparia is known from
38 squares since 1950 and additionally from 13 be-
fore 1950. The map gives only records of males and
the species can be considered as rather common.
Most records are confined to the river area with some
recent records from the northern and western part of
the country, and some old records from the south.

Jäch (1988) mentions A. riparia from very fast
flowing streams and not in stagnant waters, while
Nilsson (1984) mentions exposed lakeshores among

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9 10 11 el 12

Figs. 5-12. Distribution of Hydraena species in The Netherlands. — 5, Hydraena assimilis Rey (males only); 6, Hydraena
melas Dalla Torre (males only); 7, Hydraena pygmaea Waterhouse; 8, Hydraena pulchella Germar; 9, Hydraena minutissima
Stephens; 10, Hydraena gracilis Germar; 11, Hydraena belgica d’Orchymont; 12, Hydraena excisa Kiesenwetter.

sand and gravel in Sweden. In England Balfour-
Browne (1958) records the species from both stag-
nant and running waters. The Dutch data contradict
the findings of Jach (1988) and Nilsson (1984): there
are no records of H. riparia in fast flowing waters nor
on exposed lake shores. The species is mainly con-
fined to stagnant waters and is occasionally found in
slowly running, small streams. In fact most records are
from temporary and semi-permanent habitats with a
well developed vegetation of Phragmites australis,
Glyceria maxima or Carex and soils covered by coarse
organic debris or, in the case of running waters, small
shady brooklets less than one meter wide. The occur-
rence in larger ditches and ponds has been found only
when there exists a well developed marginal vegetation
or, single specimens have been found, when the ap-
propriate habitats are within close vicinity.

H. assimilis Rey
(fig. 5).

This very rare species is recorded from 9 squares
since 1950 in the south-eastern part of the country
(provinces of Limburg and Gelderland), but very
probably some pre-1950 females from Limburg be-
long to this species. The map gives only records of
males.

The few Dutch records of this species originate
from small ditches and pools, receiving seepage-wa-
ter, with or without vegetation, and from some small,
relatively fast flowing streams. The only reference
(Jäch 1988) mentions the same habitats as for A. ri-
paria as the most probable.

H. melas Dalla Torre
(fig. 6).

This very rare species was recorded for the first
time from The Netherlands by Cuppen (1981) from
Beertsenhoven. There are additional records of single
males from Epen (coll. RMNH and coll. O. Vorst)
and the Middelsgraaf near Echt (coll. J. Cuppen).

The few Dutch records of H. melas are from
spring-fed, small streams and, in one case, a larger,
sun-exposed, slowly running stream. The references
for H. meias are contradictory: Hrbätek (1951) and
Hebauer (1980) mention preference for cold streams
in mountainous areas, as well as acid ditches with
much detritus in river valleys, while Jach (1988) men-
tions mainly stagnant waters and small springs, but
occasionally in larger streams and rivers.

[H. nigrita Germar]

The specimens from Warnsveld and Breda, men-
tioned by Everts (1898, 1922), are deposited in the
RMNH-collection and appear to belong to H. britteni.

For this reason H. nigrita has to be removed from the
Dutch list.

CuppEn: Hydraena in The Netherlands

H. pygmaea Waterhouse
(fig. 7).

This very rare species is recorded from three
squares since 1950 in South-Limburg (Bunde, Epen,
Houthem, Noorbeek en Cottessen) with the last
record from 1991.

The few recent Dutch records of H. pygmaea are
from small (width less than 1 m), relatively fast flow-
ing streams, which are heavily shaded in one case and
unshaded in two. Generally this species is confined to
running waters in hills and mountains (Horion 1949;
Hrbätek 1951; Derenne 1952; Balfour-Browne
1958). Apparently H. pygmaea prefers sites with
aquatic mosses and periphyton on wires and boulders
(Hebauer 1980, Foster 1990). Hebauer (1980) refers
to the species as cold-stenotherm and crenophilic,
confined to the upper courses of streams and rarely
found in the middle course. The Dutch data confirm
the restriction of H. pygmaea to the smallest streams.

H. pulchella Germar
(fig. 8).

Forty specimens of this very rare species were taken
in four squares in South-Limburg and the vicinity of
Winterswijk, all at the end of the 19th century or the
beginning of the 20th century. A large series from
1921 (Wylre) represents the last record. H. pulchella
is regarded to be extinct in The Netherlands.

The Dutch records of H. pulchella suggest that, in
the past, this species mainly occurred in the larger,
fast-flowing streams (rivers Geul, Gulp and Ratumse
Beek). Generally H. pulchella is confined to running
waters, especially mossy stones and grassy edges
(Hrbatek 1951; Derenne 1952; Balfour-Browne
1958; Hebauer 1980). Foster (1990) mentions ex-
posed muddy pools in a small stream with riffles and
pools, and amongst fine tree roots in a clayey gorge in
a river. In comparison with H. pygmaea, this species is
more restricted to middle courses and open, exposed
parts of a stream. Hebauer (1980) refers to it as a rhe-
obiontic, eurythermous species.

H. minutissima Stephens

(fig. 9).

This very rare species is only known from two
squares with 15 specimens altogether. The last record
is from 1920 (Gulpen) and the species is regarded to
be extinct in The Netherlands.

The few (old) records from The Netherlands prob-
ably all originate from the middle courses of the Geul
and the Gulp, which are relatively fast flowing
streams. Abroad this running water species has been
found in small and large streams between gravel and
aquatic mosses (Hrbacek 1951; Derenne 1952;
Balfour-Browne 1958; Hebauer 1980; Valladares

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

1989). The distribution of H. minutissima seems to
depend on strong current, rather than on tempera-

ture-regime (Hebauer 1980).

H. gracilis Germar
(fig. 10)

There are only three specimens from two squares
known from The Netherlands. A record from 1923
(coll. ZMA) is the last and the species is regarded to be
extinct in The Netherlands. Most records of this spe-
cies by Everts (1898, 1922) appear to belong to either
H. belgica or H. excisa.

Generally A. gracilis is considered to be the most
common running water Hydraena in western and
central Europe, which occurs from the upper course
till the lower course, as well in cold as in exposed
streams and as well in mountains as on plains (Pretner
1930, Hebauer 1980, Valladares 1989). Also, with
reference to the current velocity, the species shows a
wide amplitude as Derenne (1952) even mentions
stagnant waters. In these habitats the species can be
found in mosses, under stones and in gravel (Horion
1949, Balfour-Browne 1958, Nilsson 1984). Despite
this wide ecological amplitude H. gracilis has only
been found twice in probably the middle and lower
course of the Geul.

H. belgica d’Orchymont
(fig. 11)

The only Dutch record is from the middle course
of the Voer river (width of 3-5 m), probably from the
beginning of the 20th century (coll. RMNH). No fur-
ther details are known. H. belgica most probably no
longer occurs in The Netherlands. Everts (1922) re-
fers to these specimens as H. gracilis.

The habitat of H. belgica in The Netherlands cor-
responds with Derenne (1952), who simply refers to
streams as its habitat. Hrbäcek (1951) mentions small
(shady) streams with a moderate slope.

H. excisa Kiesenwetter

(fig. 12).
This very rare species is known from only two
Table 1. Range, 90% range and median of observed pH-

values on localities of Hydraena species in The Netherlands.
N is the number of observations.

range 90% range median N
Hydraena assimilis 6.3 - 9.0 6.8 6
Hydraena britteni 47-78 5.5-7.3 6.5 88
Hydraena melas 7.1 - 9.0 2
Hydraena palustris 3.5-9.2 5.3-7.7 6.8 45

Hydraena pygmaea 7.9 1
Hydraenariparia 3.3-83 6.2-7.8 7.0 174
Hydraena testacea 3.5-9.9 5.5 - 8.0 6.8 151

squares since 1950 in the vicinity of Winterswijk.
Tolkamp (1980) lists H. excisa for the first time for
The Netherlands, without mentioning it as an addi-
tion to the Dutch fauna. However, Everts (1898) al-
ready recorded H. gracilis var. erosa and var. excisa
from Winterswijk, but the material could not be
found in the museum collections. The first specimens
from Winterswijk, labelled as H. gracilis, in these col-
lections date from 1918 and all later references
(Everts 1922, Laijendecker & Nieser 1971) belong to
H. excisa.

The recent Dutch records are from the Ratumse
Beek (width about 3-5 m with shaded and unshaded
sections) and some smaller tributaries. The species
has been found here amongst gravel in relatively fast
flowing parts of the stream (Laijendecker & Nieser
1971, Tolkamp 1980). Elsewhere in Europe H. excisa
occurs in small, shady streams with a moderate slope
(Hrbacek 1951) or sun-exposed streams (Hebauer
1980). According to the latter author the species is
not cold-stenotherm and has no preference with re-
spect to calcium. Within the streams it can be found
in mosses or between and under stones (Knie 1974;
Hebauer 1980).

PHYSICAL AND CHEMICAL DATA

The ranges of the observed pH-values (table 1) are
large for most Hydraena species. However, the me-
dian values of pH for all frequently encountered
Hydraena species are between 6.5 and 7.0, with 90%
of the observed values within 1.3 pH-unit above or
below the median pH of that species. A. britteni has
the lowest median pH-value and H. riparia the hig-
hest. As the 90% ranges of pH for the different spe-
cies considerably overlap and the median values are
very close to each other, the pH can not be conside-
red a very important factor for the explanation of the
different geographic distribution patterns of these
Hydraena species in The Netherlands.

Ranges for the values of the electrical conductivity
(table 2) are very large for A. riparia and H. testacea
and large for A. britteni and H. palustris. The 90%
ranges for the most common species reduce the upper
limit for electrical conductivity considerably with the
smallest range for H. britteni and a larger, more or less
the same, range for the other species. The median val-
ue of electrical conductivity is distinctly higher for H.
riparia in comparison with the other species, which
partly can be explained by the different distribution
patterns of these species. However, also in this respect
there is a considerable overlap in ranges.

Ranges, 90% ranges and median values of total
hardness (table 3) are more or less the same for the
common Hydraena species and certainly cannot ex-
plain their different geographic distribution patterns.

Table 2. Range, 90% range and median of observed values
of electrical conductivity (US cm') on localities of Hydraena
species in The Netherlands. N is the number of observa-
tions.

range 90% range median N
Hydraena assimilis 330- 710 490 6
Hydraena britteni 107-1260 192- 730 450 88
Hydraena melas 420- 620 2
Hydraena palustris 124-1590 150 - 1030 440 45
Hydraena pygmaea 580 I
Hydraena riparia 55 - 2030 228-1140 560 174

Hydraena testacea 87-2030 174-1030 460 151

Table 3. Range, 90% range and median of observed values
for total hardness ("D) on localities of Hydraena species in
The Netherlands. N is the number of observations.

range 90% range median N
Hydraena assimilis 6-14 10 6
Hydraena britteni 1-23 3-16 8 46
Hydraena melas 9 1
Hydraena palustris 2-25 3-21 6 33
Hydraena riparia 1226. AD] 9 Vo,
Hydraena testacea We) Mall 6 88

Based on only 6 observations, H. assimilis has a rela-
tively small range with a high median value.

Of the four investigated parameters, chlorinity is
the most informative with respect to the distribution
patterns of Hydraena (table 4). All species are con-
fined to fresh water with only one sample with a chlo-
rinity of more than 200 mg/l for H. britteni, H. palus-
trisand H. testacea, while for H. riparia there are five.
Chlorinity is certainly highly responsible for the ab-
sence of Hydraena species in the western and north-
ern part of the country of which the coastal dune area
is excluded due to the presence of fresh waters.
Median and 90% ranges of the most common
Hydraena species are also in agreement with their
general distribution patterns: H. riparia occurs main-
ly between the large river systems, H. palustris is scat-
tered all over the country with the exception of brack-
ish parts, H. testacea occurs on sandy soils in the

Cupren: Hydraena in The Netherlands

Table 4. Range, 90% range and median of Cl--values (mg | ')
on localities of Hydraena species in The Netherlands. N is
the number of observations.

range 90% range median N
Hydraena assimilis 36.5 - 102.8 41.9 6
Hydraena britteni 6.8 - 270.0 14.5 - 92.7 36.5 88
Hydraena melas 19.8 - 41.9 2
Hydraena palustris 6.8 - 355.0 17.1 - 130.0 60.7 45
Hydraena pygmaea 19.6 1

Hydraena riparia 7.1 - 534.0 18.8 - 172.2 64.4 174
6.9 - 492.4 16.6 - 126.3 49.2 151

Hydraena testacea

eastern, southern and northern parts of the country
and between the large rivers and A. britteni is mainly
found on the sandy soils in the southern, central and
eastern part of the country.

Co-EXISTENCE

In table 5 the co-existence of five Hydraena species
in The Netherlands is shown. The number of co-exis-
tences for each species exceeds the total number of oc-
currences because more than two species often co-
exist. H. assimilis is found alone or in company with
H. testacea. H. britteni lives most often alone or in co-
existence with H. testacea. H. palustris lives most often
in company with H. riparia and/or H. testacea, and
only occasionally alone. H. riparia occurs most often
alone, very frequently with H. testacea and frequently
with H. palustris. H. testacea most often lives alone,
very frequently with H. riparia and frequently with
H. britteni and/or H. palustris.

Proportionally H. britteni is most often the only
Hydraena species at a locality, indicating that its hab-
itat requirements may be more exclusive than those of
the other Hydraena species. H. palustris rarely occurs
alone, and it shares its habitat quite often with A. ri-
paria and/or H. testacea. H. testacea, though frequent-
ly occurring alone, co-exists quite often with all other
Hydraena species, indicating that it can live in quite
different habitats. H. testacea can be considered as
the most euryoecious Hydraena species in The

Netherlands.

Table 5. Association of Hydraena species in The Netherlands. N is the number of observations.

N Alone with H. with H. with H. with A. with A.

assimilis brittent palustris riparia testacea
Hydraena assimilis 10 4 = 2 = 1 6
Hydraena britteni 104 78 2 = 3 5 21
Hydraena palustris 52 11 - 3 = 28 19
Hydraena riparia 196 126 ] 5 28 — 51
Hydraena testacea 164 82 6 21 19 51 =

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

PHENOLOGY

Phenological data at monthly intervals of adult
Hydraena species are given for H. palustris, H. britte-
ni, H. riparia and H. testacea (figs. 13-16). For the re-
maining species the number of records of adults is too
low to present meaningful diagrams. Keys for the
identification of the larvae of Hydraena are not availa-
ble and the larvae of only very few Hydraena species
have been described so far (Hrbácek, 1943-44). As
the number of collected larvae of this genus is very
low and the above mentioned species quite often co-
exist, it is impossible to include these data in the figu-
res.

H. palustris has been collected throughout the year
with a distinct maximum in April, which probably
coincides with the breeding period. Cuppen (1983)
reports a different phenology for H. palustris with a
spring maximum in May and a (lower) autumn max-

imum in October. The same phenology was found by
Landin (1976) in Lake Sagsjon near Stockholm,
though the autumn peak occurred in September.
Adults in copula, however, have never been observed.
Teneral adults have been observed between 7.viii and
18.xi, indicating a rather long emergence period, and
without a clear autumn maximum. This type of life
cycle agrees with life cycle type I of Nilsson (1986).
H. britteni can be collected throughout the year
with clear maxima in March/April and November
(the absence of records in January is certainly artifi-
cial). A similar phenology has been reported by
Cuppen (1983) in a study of a population of A. brit-
teni in a seepage-marsh near Apeldoorn (The
Netherlands) during one year. Landin (1976) found a
spring maximum in the beginning of May and a (low-
er) autumn maximum in September. The spring
maximum coincides with the breeding period as
many adults in copula have been observed between

340
|
120
50 100 +
40 u ao
30 so}
20 40
|
10 20 IL
0 0
J F M À M J IAS ON DO J
13

F M À MJ

JF M À M J

AES ORNE
14

Figs. 13-16. Number of
specimens Aydraena spe-
cies at monthly intervals. —
13, Hydraena palustris
Erichson; 14, Hydraena
britteni Joy; 15, Hydraena
riparia Kugelann; 16,
16 Hydraena testacea Curtis.

JAS OWN D

28.xii and 10.iv and the species can be considered as a
winter breeder. Some pairs in copula even have been
found in ice-covered pools. Teneral adults have been
found in a very few occasions between 30.vii and 13.x
before the November maximum. Cuppen (1983)
mentions teneral adults in October and November.
This low number of observations of teneral adults can
be explained by the drying out of most habitats of A.
britteni in the summer period. Adults probably leave
their pupal chambers after the rising of the water lev-
el due to autumn rains, when they are already fully
developed. The November maximum coincides with
the filling up with water of the habitats of H. britteni
during the period in which their gonads develop.
This type of life cycle with winterbreeding is not
mentioned by Nilsson (1986).

H. riparia has been collected throughout the year
with two peaks in abundance in April and
August/October. The spring maximum probably co-
incides with the breeding period but pairs in copula
have been found only once on 4.iii. Teneral adults
have been observed in great numbers between 23.vii
and 13.xi (mainly August and the beginning of
September), and therefore coinciding with the late
summer maximum. The life cycle agrees with the life
cycle type I of Nilsson (1986).

H. testacea has been collected quite evenly through-
out the year with a maximum in April. This April
maximum probably coincides with the breeding peri-
od though pairs in copula never have been found.
Teneral adults have been found in abundance
between 24.vii and 14.xi with a distinct maximum in
September. Also in this species its life cycle is in agree-
ment with type I of Nilsson (1986).

Based on the museum material, all other species
have been collected mainly between May and
September indicating more the activity of the collec-
tors than the activity of the beetles. This also applies
to the data for the above-mentioned species in the
museum collections. The low number of records of
Hydraena in museum collections is certainly partly
based on the low activity of collectors in autumn,
winter and spring.

DISCUSSION

The various distribution patterns of Hydraena in
The Netherlands are in the first place determined by
stream-velocity or a parameter associated with it.
Most species are rheophilic or rheobiontic and there-
fore confined to the southern and eastern part of The
Netherlands. The maps suggest that nearly all species
of this group are restricted to areas with fast running
waters and do not occur in areas with more slowly
running lowland streams. Paucity of recent records of
these species permits no further conclusions about

Cuppen: Hydraena in The Netherlands

their preferred habitats. As running water species of
Hydraena have no obvious morphological adaptations
to that type of habitat it is difficult to judge which pa-
rameters are responsible for their restricted occurren-
ce. A permanent high oxygen content of the water
could be a main factor, as most species disappeared
after pollution and subsequent decrease of oxygen.

For the stagnant water species chlorinity seems to
be an important parameter for the explanation of
their distribution patterns. All species are mainly re-
stricted to waters with a chlorinity less than 200 mg/l.
Other, less well known parameters, as soil type or the
occurrence of seepage, probably also play an impor-
tant role in their distributions. Quite often, however,
two or three species can co-exist in the same locality,
indicating that the different distribution patterns are
not gouverned by the measured parameters.

Until 1991 twelve species of the genus Hydraena
have been found in The Netherlands. Most of them,
if not all, have been collected before 1950, but many
of them were not recognized until recently after dis-
section of the male genitalia. From these twelve spe-
cies only eight can be considered to occur at present
in The Netherlands. H. pulchella, H. minutissima, H.
gracilis and H. belgica have been collected for the last
time at the beginning of this century and probably
these species were confined to large streams as the
Geul, Gulp, Voer (and Ratumse Beek) in the south-
ern (and eastern) part of the country. Their disap-
pearance is probably due to water pollution and, to a
minor extent, to regulation of streams. Some of these
species can, after improvement of water quality, re-
turn to The Netherlands from neighbouring streams
in Belgium and Germany. However, colonisation
power of Hydraena species seems to be limited. The
remaining running water species (H. melas, H. excisa,
H. assimilis and H. pygmaea) are all very rare and
under serious threat. Especially H. excisa, which occu-
pies an extreme western outpost in The Netherlands,
will not easily return after extinction. The chances for
survival of the other three species, which occur main-
ly in smaller streams or ditches receiving seepage-wa-
ter, seem to be better, as most localities are situated in
nature reserves. Hydraena species of stagnant waters
are widely distributed and are not under threat.
However, a decline of H. testacea and H. riparia in the
western parts of The Netherlands is suggested by the
available data.

ACKNOWLEDGEMENTS

This study would not have been possible without the ma-
ny specimens, set at my disposal, from private and museum
collections mentioned under material. Dr. G. N. Foster and
anonymous referees are thanked for commenting on a first

draft of this paper.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

REFERENCES

Balfour-Browne, F., 1958. British water beetles III. — Ray
Society, London, 210 pp.

Berthélemy, C., 1986. Remarks on the genus Hydraena and
revision of the subgenus Phothydraena (Coleoptera:
Hydraenidae). — Annales de Limnologie 22: 181-193.

Cuppen, H. P. J. J., 1983. Een oecologisch onderzoek naar
de macrofauna van een temporair kwelmoeras op de
Oost-Veluwe. I. De waterkevers. — Regionale Milieuraad
Oost-Veluwe, Apeldoorn, 17 pp.

Cuppen, J. G. M., 1981. Hydraena bohemica Hrbacek,
nieuw voor Belgié en Nederland (Coleoptera, Hydrae-
nidae). — Phegea 9: 61-64.

Cuppen, J. G. M., 1985. Rediscovery of Hydraena testacea
Curtis (Coleoptera: Hydraenidae) in Scotland. —
Entomologist’s Gazette 36: 181-182.

Cuppen, J. G. M., 1991. Genus Hydraena Kugelann. In: De
waterkevers van Nederland (Coleoptera) (eds B. Drost,
H. P. J. J. Cuppen, E. J. van Nieukerken & M. Schreijer).
— Natuurhistorische Bibliotheek Koninklijke Neder-
landse Natuurhistorische Vereniging 55: 280 pp.

Cuppen, J. G. M. & H. P. J. J. Cuppen, 1982. Hydraena
britteni Joy new for The Netherlands (Coleoptera: Hy-
draenidae). — Entomologische Berichten, Amsterdam 42:
45-48.

Cuppen, J. G. M. & K. Dettner, 1987. The larvae of the
predaceous water beetle Laccornis oblongus (Stephens)
(Coleoptera: Dytiscidae), with notes on ecology and dis-
tribution. — Aquatic Insects 9: 211-220.

Derenne, E., 1952. Les Hydraena de Belgique. — Bulletin et
Annales de la Société Entomologique de Belgique 88:
195-218.

Drost, M. B. P., 1989. Helophorus croaticus and H. pumilio
in The Netherlands, with description of their larvae
(Coleoptera: Hydrophilidae). — Entomologische Berich-
ten, Amsterdam 49: 1-7.

Everts, E., 1898. Coleoptera Neerlandica. I.: i-viii, 1-676. —
Nijhoff, ’s Gravenhage.

Everts, E., 1922. Coleoptera Neerlandica. III: i-xviii, 1-
667. — Nijhoff, ’s Gravenhage.

Foster, G. N., 1990. Atlas of British water beetles.
Preliminary edition. Part 6. — Balfour-Browne Club
Newsletter 48: 1-18.

Hebauer, F., 1980. Beitrag zur Faunistik und Okologie der
Elminthidae und Hydraenidae in Ostbayern (Coleo-
ptera). Mitteilungen der Miinchener Entomologischen
Gesellschaft 69: 29-80.

Horion, A., 1949. Faunistik der Mitteleuropäischen Kafer

10

II. — Vittorio Klostermann, Frankfurt am Main, 388 pp.

Hrbätek, J., 1943-44. O larvách rodu Hydraena
(Coleoptera, Hydrophilidae). — Sbornik Entomologic-
kého Oddeleni Narodniho Musea v Praze 21-22: 84-89.

Hrbätek, J., 1951. Revue des espèces du genre Hydraena
Kug. sur le territoire de la république Tchecoslovaque
(Col. Hydroph.). — Casopis Ceskoslovenske Spolecnosti
Entomologickä 48: 201-226.

Ienistea, M. A., 1978. Hydradephaga und Palpicornia. — In:
Illies, J., Limnofauna Europaea. Gustav Fischer Verlag,
Stuttgart, 532 pp.

Jach, M. A., 1988. Revisional notes on the Hydraena ripar-
ia species complex (Coleoptera: Hydraenidae). — Aquatic
Insects 10: 125-139.

Knie, J., 1974. Ein Beitrag zur Verbreitung und Okologie
von Hydraena excisa Kiesw. — Decheniana 127: 263-264.

Landin, J., 1976. Seasonal patterns in abundance of water
beetles belonging to the Hydrophiloidea (Coleoptera). —
Freshwater Biology 6: 89-108.

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wantsen uit de omgeving van Winterswijk (Coleoptera et
Heteroptera aquatica). — Entomologische Berichten,
Amsterdam 31: 3-12.

Nilsson, A. N., 1984. The distribution of the aquatic beetle
family Hydraenidae (Coleoptera) in northern Sweden,
with an addenda to the Elmidae. — Fauna Norrlandica 4:
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Belgique 65: 45-53.

Pretner, E., 1930. Ueber die angebliche Variabilitat der
Hydraena gracilis Germar. — Coleopterologisches Zen-
tralblatt 5: 81-105.

Tol, J. van, 1981. An introduction to The Netherlands. —
Nieuwsbrief European Invertebrate Survey - Nederland
10: 5-12.

Tolkamp, H. H., 1980. Organism-substrate relationships in
lowland streams: 1-211. Centre for Agricultural
Publishing and Documentation, Wageningen.

Valladares, L. F., 1989. Los Palpicornia acuaticos de la pro-
vincia de Léon. II. Hydraena Kugelann, 1794 y Limnebius
Leach, 1815 (Coleoptera: Hydraenidae). — Boletin de la
Asociacion Espafiola de Entomologia 13: 313-330.

Received: 12 November 1991
Revised version accepted: 10 September 1992

RYSZARD HAITLINGER

Department of Zoology, Wroclaw Academy of Agriculture, Poland

A NEW SPECIES OF THE GENUS
DIPLOTHROMBIUM BERLESE (ACARI,
PROSTIGMATA, JOHNSTONIANIDAE) FROM
POLAND, BASED ON THE LARVA

Haitlinger, R., 1993. A new species of the genus Diplothrombium Berlese (Acari, Prostigmata,
Johnstonianidae) from Poland, based on the larva. — Tijdschrift voor Entomologie 136: 11-13,
figs. 1-8. [issN 0040-7496]. Published 1 July 1993.

Diplothrombium ludwinae sp. n. (larva) is described from Poland and a key is provided to the
larval stages of the genus.

Dr. R. Haitlinger, Department of Zoology, Wroclaw Academy of Agriculture, ul. Cybulskiego
20, 50-205 Wroclaw, Poland.

Key words. — Acari, Johnstonianidae, new species, Poland.

The genus Diplothrombium Berlese was established from the U.S.A.(Newell 1957, Feider 1959, Robaux
in 1910. Since then, only four species based on larvae 1977, Southcott 1987). Larvae of Diplothrombium
have been described, viz. D. monoense Newell, D. cas- were found attached to aquatic beetles (Coleoptera)
cadense Newell, both from the U.S.A., D. moldavi- (Newell 1957) and Tipulidae (Diptera) (Feider

cum Feider from Romania and D. newelli Robaux 1959). Up to now, only one species was known from

all

visible

Figs. 1-8. Diplothrombium ludwinae sp. n. -- 1, idiosoma, dorsal view; 2, scutum; 3, idiosoma, ventral view; 4, palp; 5, palp-
tarsus; 6, leg I, tarsus-trochanter; 7, leg II, tarsus-trochanter; 8, leg III, tarsus-trochanter.

TIJDSCHRIFT voor ENTOMOLOGIE, VOLUME 136, 1993

Table 1. Measurements in mp of Diplothrombium ludwinae
sp. n., larva, holotype.

Length of idiosoma 904
Width of idiosoma 752
Length of scutum 108
Width of scutum 76
Distance between bases of AL (AW) 46
Distance between bases of PL (PW). 56
Length of anterior scutala (AL) 42
Length of posterior scutala (PL) 60
Distance between ASE (SBa) 10
Distance between PSE (SBp) 40
Length of anterior sensilla (ASE) 16
Length of posterior sensilla (PSE) 134
Distance between ASE ans PSE (ISD) 38
Distance between AL and PL (AP) 48
Distance between AL and ASE (AAS) 20
Ocular sclerite 34
Length of dorsal setae (DS) 78-90
Coxala I 56
Coxala II 58
Coxala III 40
fd 44
gd 36
Tal 104
Til 46
Ge I 44
TfI 52
BfI 46
Tre ll 40
Cx I 88
Ta II 84
Ti II 44
Ge II 40
Tf II 40
Bf II 40
Tr I 34
Cx II 76
Ta III 92
Ti Ill 62
Ge III 46
Tf Il 32
Bf III 62
Tr III 46
Cx III 86
Tale) > dal (A) 3.25
Til: Gel 1.04
Ti ll: Ge II 1.10
Ta III : Ge III 2.00
AW : ISD 1,211
ISD : AP 0.79
Til: AW 1.00
Ti II: AW 1.35

AW : AL 1.09

Europe. In the present paper a new species is de-
scribed from Poland, and a key to the larvae of
Diplothrombium is provided. Terminology for setae
and structures follows Southcott (1988). All measure-
ments are in micrometers (Um).

Diplothrombium ludwinae sp. n.
(figs. 1-8)

Type material. — Holotype larva: Poland, Zawoja-
Markowe, 800m (voi. Bielsko-Biala), 9.VI.1983, beech-co-
niferous forest from plants, leg. R. Haitlinger (in Museum

of Natural History, Wroclaw University).

Description

Idiosoma longer than wide, oval, anterior part of
idiosoma somewhat deformed. Dorsum with 27 bar-
bed setae arranged in five rows: 6, 7, 6, 5 and 3; each
setae placed on oval and small platelet (fig. 1). Eyes
small. Scutum longer than wide; behind bases of PSE
with transverse crista which divides scutum into une-
qual parts. Scutum with two pairs of scutalae (AL,
PL); posterior setae PL longer than AL; further with
two pairs of sensilla (ASE, PSE) from which posterior
sensilla PSE many times longer than ASE. Between
basis of ASE and posterior margin of scutum a longi-
tudinal crista. All scutalae and sensilla smooth.
Anterior margin with nasus (fig. 2). Ventral side of
idiosoma with 74 setae, each placed on small platelet;
posterior setae somewhat longer and placed on larger
platelets. Anus located in middle part of opisthosoma.
Two setae between coxae III.

Gnathosoma short, its base with two bifurcate se-
tae. Palpfemur and palpgenu each with one barbed
seta; palptibia with three setae, two of these are
barbed. Palptarsus long and with eight setae, five of
which are barbed (figs. 4, 5).

Legs short, length (including coxae, excluding
claws): I 420, II 358, III 426; coxa I with one slightly
barbed setae; medial coxala I bifurcate and separate
from coxa; coxa II with one barbed seta; coxa III with
one smooth seta. Tarsus II with enlarged solenidion
and small famala. Number of solenidions and other
setae on tibia, genu, telofemur, basifemur and tro-
chanter | e 111 2250246, @ey72So NI 2 Sor aan
lS or ONCS Or SOR ZN
2, Ne 1500 = IM Il Sor ©, Ge 2 So + 4, MI So + 3, Bir
QI

Measurements of holotype, sce table 1.

Remarks. — D. /udwinae sp. n. is similar to D. mol-
davicum Feider and D. cascadense Newell. It can be
distinguished from the first species by the absence of
a transverse bar on scutum above the posterior sensil-
la, barbed coxalae II, shorter legs I-III and tarsi I-III,
especially I and III; from the second species by the
shape of the posterior part of the scutum and the
smooth scutalae.

Etymology. — Name derived from Ludwina.

Key to species of Diplothrombium larvae

il. Scutum with three transverse bars, two of them
placed before the bases of posterior scutal sensilla
teek cl BL D. moldavicum Feider, 1959

— Scutum without transverse bars before the bases
o@posterorseutalisensi]t tn 2
2. Scutum with longitudinal and transverse bars be-
low the bases of posterior sensilla ..................... 3
— Scutum without transverse and longitudinal bars
below the bases of posterior sensilla .................. 4
3. Scutalae barbed, distance between bases of poste-
rior scutalae PW distinctly larger than the dis-
tance between bases of anterior scutalae; AW =
1.85 RENNES D. cascadense Newell, 1957
— Scutalae smooth, distance between bases of poste-
rior scutalae somewhat larger than the distance
between bases of anterior scutalae; AW = 1.22 …
PE NSA RAI a RANA REACH AVIO TARE D. ludwinae sp. n.
4. Medial coxala I separated from coxa, genua I with

8 solenidions ............... D. newelli Robaux, 1977
— Medial coxala I on coxa I, genua I with 15 sole-

DION NO D. monoense Newell, 1957
REFERENCES

Feider, Z., 1959. Prima specie a genului Diplothrombium

HAITLINGER: New species of Diplothrombium

(Acari) din R. P. R. si Europa sub forma de larva
(Diplothrombium moldavicum n. sp.). — Academia R. P.
R. Filiala Iasi Studii si Cercetari Stiintifice si Stiinte
Agricole 10: 261-268.

Newell, I. M., 1957. Studies on the Johnstonianidae (Acari,
Parasitengona). — Pacific Science 11: 396-466.

Robaux, P., 1977. Observations sur quelques Actinedida
(=Prostigmates) du sol d’Amérique du Nord. IX.
Nouvelles formes larvaires de trombidions (Acari). —
Acarologia 19: 258-271.

Southcott, R. V., 1987. The classification of the mite fami-
lies Trombellidae and Johnstonianidae and related
groups, with the description of a new larva (Acarina:
Trombellidae: Nothrotrombidium) from North America.
— Transactions of the Royal Society of South Australia
111: 25-42.

Southcott, R. V., 1988. Two new larval mites (Acarina:
Erythraeidae) ectoparasitic on north Queensland cicadas.
— Records of the South Australian Museum 22: 103-116.

Received: 20 December 1992
Accepted: 12 April 1993

13

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

BOOK REVIEW

Ossiannilsson, F., 1992. The Psylloidea (Homoptera) of
Fennoscandia and Denmark. — Fauna Entomologica
Scandinavica 26: 1-346, figs. 1-1415 + 1-9. [ISBN 90 04
09610 8]. Published by E. J. Brill, Leiden. Price NLG
180.00 (appr. USD 105.00).

The important series ‘Fauna Entomologica
Scandinavica’, with much more relevance to other
NW European countries than can be expected from
the title, continues with another volume on Homo-
ptera, viz. the Psylloidea. The author, F. Ossiannils-
son, is a well-known specialist of this group and an
author of Homoptera studies for more than fifty
years. He also contributed earlier to this series with
three volumes on the Auchenorrhyncha (1978-
1983).

The present volume includes for each species syn-
onymy, descriptions, including of larval stages, dis-
tributional data and biology. Especially from the biol-
ogy paragraphs, it is clear that the author is able to
summarize his knowledge, based on the literature and
on his own experience as well, in a concise and yet

easily readible form. Moreover, a wealth of original
data are provided for many of the species.

Terminology is clearly explained in the introducto-
ry chapters, while various keys guide the reader to the
names of families, subfamilies, genera and species.
Not less than 1424 figures illustrate all kinds of rele-
vant characters of the 98 species included.

Psyllids can cause, directly or indirectly, consider-
able damage to their host-plants, and some species are
specific to host-plants of economic importance (car-
rot, pear). The present work is, therefore, not only
strongly recommended to museums or amateur ento-
mologists, but also to „vraries of applied entomolo-
gists. Unfortunately, the publisher seems to discon-
tinue his policy of producing this series for a
reasonable price, although I certainly understand the
connection with the limited number of copies that
can be sold. Nonetheless, the price will inevitably af-
fect the possibilities for the individual entomologist
to purchase a personal copy.

[J. van Tol]

I. LANSBURY

Oxford University Museum, United Kingdom

STRONGYLOVELIA (VELIIDAE) AND

METROBATOPSIS (GERRIDAE) AND ASSOCIATED
PLEUSTON HEMIPTERA OF WEST NEW BRITAIN

I. Lansbury, 1993. Strongylovelia (Veliidae) and Metrobatopsis (Gerridae) and associated
pleuston Hemiptera of West New Britain. — Tijdschrift voor Entomologie 136: 15-22, figs. 1-
26 [issn 0040-7496]. Published 1 July 1993.

A brief survey of the pleuston bugs of the Von River, West New Britain are given.
Strongylovelia priori sp. n. is described (Veliidae: Haloveliinae) with distributional data from
other localities. Two species of Rhagovelia (Veliidae: Rhagoveliinae) are listed. Metrobatopsis
flavonotatus Esaki (Gerridae) is described and figured as its specific identity is uncertain. Other
gerrid genera and species are listed, viz. Ptilomera, Limnometra and Tenagogonus.

Dr. I. Lansbury, The University Museum, Parks Road, Oxford ox1 3pw, United Kingdom.
Key words. — Hemiptera; Strongylovelia, Metrobatopsis, New Britain; new species.

During a single visit to the Von River near Dami,
West New Britain, the pleuston species diversity was
found to be rather unusual as it included species char-
acteristic of both lentic and lotic habitats. The river
runs through a deeptly cut channel of what appears to
be volcanic debris and silt. Substrate varying between
coarse sand and gravel in slower shallow stretches and
much deeper extremely fast stretches with large stone
and boulders. The banks are steep, banded alternate-
ly with black layers and broader intervening pale
brown bands of silt. The habitat is deeply shaded by
overhanging secondary forest and in places a dense
sub-shrub layer.

Supplementary data from other regions of Papua
New Guinea are given for Strongylovelia priori sp. n.,
Metrobatopsis flavonotatus Esaki and other species of
Veliidae and Gerridae. Data for Rhagovelia (Veliidae),
Ptilomera, Metrobatopsis and possibly some species of
Tenagogonus are found on slow-fast lotic habitats,
whereas Limnometra species (all Gerridae) most com-
monly found on lentic habitats. Published data on
Strongylovelia give no indication of habitat preferences.

SYSTEMATICS
Veliidae, Haloveliinae

Strongylovelia priori sp. n.
(figs. 1-19, table 1)

Type material. — Holotype d : Papua New Guinea, West
New Britain, Balima River near Ulamo, 19.viii.1989, R. N.
B. Prior. — Paratypes (all Papua New Guinea, West New
Britain): Von River, Banaule near Dami, 21.11.1990, 129;
Tamari Creek, freshwater/saline habitat, 11.xii.1988, R. N.

B. Prior, 19. The holotype and a series of paratypes in
OXUM, other paratypes in RMNH.

Description

Adult apterous. Male 1.6 mm long, maximum
width 0.77 mm, female 1.58-1.76 mm long, maxi-
mum width 0.86-0.99 mm.

Coloration. — Female: Head black, latero-posterior
margins and cephalic trichobothria dark yellow
tinged pale red. Eyes silvery grey. Antennae black,
first segment ventrally pale yellow. Rostrum shining
dark brown to black. Pronotum black, anterior mar-
gin narrowly pale yellow. Mesonotum broadly pale
yellow, lateral margins black. Metanotum with two
(1+1) yellow-orange blotches narrowly separated
from yellow mesonotum. Anterior tergites uniformly
black, distal tergites faintly iridescent. Connexivum
posteriorly broadly pale yellow. Propleura narrowly
black posterior of eyes, mesopleura broadly pale yel-
low with upper margin and most of metapleura black.
Metasternum pale yellow. Sternites laterally yellow,
posteriorly uniformly blackish-brown, ventrally black
(fig. 3).

Legs: Front leg, trochanter and femur pale yellow,
distally femur narrowly annulated dark brown. Inner
margin of tibia anteriorly pale yellow, remainder and
tarsi dark brown. Middle and hind legs black, tro-
chanter and proximal part of hind femur pale yellow.

Male similar to female, but pale yellow pronotal
band narrower. Meso and metanotum and meso and
metapleura as in female. Tergites black, not iridescent
posteriorly. Connexivum and sternites black.

Structure. — Female subovate, length 1.82x greatest

15

Figs. 1-5. Strongylovelia priori sp. n., 1-2, 5 holotype d , 3-4 paratype ?.—1, dorsal habitus; 2, ventral aspect; 3, dorsal hab-

itus; 4, side view; 5, side view Scale line 0.5 mm.

width across thorax. Head length about 1.6x greatest
width. Eye width about 0.4x width of head between
eyes. Thoracic dorsum clearly raised above tergites
(fig. 4). Mesonotum and metanotum with sparse ad-
pressed pubescence. Pro and mesopleura with much

longer prominent curled black pubescence. Apical

tergites broad, covered with adpressed greyish pubes-
cence, distal tergites less pilose, shining greenish iri-

16

descent. Connexivum clearly raised above tergites,
apically sinuate, curving and more erect posteriorly.
Connexivum with long black curled hairs. Viewed
laterally, connexivum posteriorly with a cluster of
erect black hairs (figs. 3-4).

Female genitalia (figs. 18-19). Tergum 8 large, first
gonocoxae elongate. First gonapophyses distally with
scattered prominent spines. Second gonapophyses

ES

\
I
it
N
Ni)
N
MN

AIN
\
| I
|
|

3

Figs. 6-9. Strongylovelia priori sp. n. — 6, holotype d, anten-
na; 7, same, hind leg; 8, paratype female, antenna; 9, same,

hind leg. Scale line 0.5 mm.

connected by a lightly sclerotised bridge, distally with
fringes of fine hairs. Proctiger viewed laterally elon-
gate, subquadrate from dorsal aspect.

Male elongate, about 2x greatest width across tho-
rax. Head length slightly less than greatest head width
between eyes. Greatest eye width about 0.4x head
width. Tergites slightly raised, connexivum continu-
ous with tergites. Lateral margins of prothorax, ter-
gites and connexivum covered with long hairs.
Sternites not pilose with scattered minute spicules,
more prominent and in organised row on distal seg-
ment (figs. 1, 2, 5).

Male genitalia (figs. 14-17): Genital segment hid-
den within abdomen and moderately sclerotised,
proctiger prominent. Parameres symmetrical, viewed
laterally long and slender, from dorsal aspect sinuate.

Legs: Male front femur moderately incrassate, tibia
distally with a row of fine spinose hairs (fig. 10).

LL

= ZE

LANSBURY: Strongylovelia of West New Britain

Table 1. Proportions of leg segments of Strongylovelia prio-
ri sp. n.

Femur Tibia Tarsil Tarsi II
Male front leg 60 60 8 20
Female front leg 63 63 8 22
Male middle leg 130 108 44 25
Female middle leg 141 105 40 25
Male hind leg 87 ij 13 20
Female hind leg 80 72 13 15

Middle femur about half length of male (21:41) (fig.
12). Hind femur slightly more robust than middle fe-
mur, ventrally with numerous long spine-like hairs
(fig. 7). Female front femur slightly curved, tibia dis-
tally with a row of fine spinose hairs (fig. 11). Middle
femur about half length of female (fig. 13). Hind fe-
mur not noticeably thicker than middle femur, ven-
trally fringed with fine hairs (fig. 13).

Antennal segments 1-4 male (fig. 6) 26 : 37 : 63 :
52. First segment about 0.66x head length, second
segment very robust. Female segments 1-4 (fig. 8) 27
: 33 : 50 : 42. First segment just over half median —
head length.

Remarks

Strongylovelia priori sp. n. is similar in general ap-
pearance to S. formosa Esaki (1924) described from
two females from Taihoku, Northern Formosa. The
hind femora of formosa are almost entirely pale and
the pronotum and connexivum of the female are uni-
formly dark brown to black. Lundblad (1933) figured
the male of formosa from Central Sumatra, Singkarak.
The parameres of formosa sensu Lundblad are proxi-
mally very broad compared with priori. Lundblad
gives the relative lengths of male antennal segments 1-
4 as 50 : 53 : 92 : 67. He states ‘In his Figure 1, Esaki,
has shown the Ist segment too short, because it has
apparently been drawn from an antenna that has not
been detached. The condyle is therefore not visible.
In fact the Ist and 2nd segments agree so closely in
their length that no differences can be detected except
by measuring them’. Figures 6 and 8 of priori are
drawn from slide mounted partially cleared prepara-
tions.

Esaki described a second species, S. albicollis from
New Guinea, Erima, Astrolabe Bay, based on a apter-
ous female 1.5 mm long and a macropterous female 2
mm long. The apterous albicollis is distinguished
from priori by the broadly yellowish white pronotum
and the two yellowish white blotches on the black
mesonotum. The yellowish white coloration of part
of the connexivum is similar to priori. The variation
of coloration between male and females of

107

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 10-13. Strongylovelia priori sp. n. — 10, holotype à, front leg; 11, paratype female, front leg; 12, holotype male, middle

leg; 13, paratype female, middle leg. Scale line 0.5 mm.

Strongylovelia and iridescence of the distal tergites
have not previosuly been commented upon.
Andersen (1982) figures the apterous female of S. for-
mosa and forewing of macropterous form showing the
extreme reduction of the venation.

The longest series of Strongylovelia were collected
from shaded moderately slow lotic stretches of the
Von River, occurring with Metrobatopsis (Gerridae).
Because of their small size, they were impossible to see
on the surface of the river.

Etymology. — A noun in the genitive case. This elu-
sive haloveline is dedicated to Dr. R. N. B. Prior for
his invaluable assistance whilst in West New Britain
and for the collections he made for me during his ten-
ure at Kimbe, West New Britain.

18

Rhagoveliinae

Rhagovelia biroi Lundblad

Rhagovelia biroi Lundblad, 1936: 13-14: type series New
Guinea, Erima, Astrolabe Bay and Stephansort and East
New Britain, Herbertshöhe [Kokopo near Rabaul] in
HNHM [not examined]; Polhemus & Polhemus 1988:
165.

Material. — Papua New Guinea, West New Britain, Von
River, Banaule near Dami, 21.iii.1990, I. Lansbury, 17d
109, all apterous. This series were found principally along
the margins amongst fallen branches. It is moderately com-
mon at Tamari, Bialla and Bilomi River, Kimbe Bay, West
New Britain. Polhemus & Polhemus (1988) place biroi in
the ‘novacaledonica group’.

LANSBURY: Strongylovelia of West New Britain

Figs. 14-19. Strongylovelia priori sp. n., 14-17, holotype d genitalia; 18-19, paratype female — 14, segment 8 and ancilliary
structures, scale line 0.25 mm; 15, genital capsule dorsal aspect; 16, same, side view; 17, part of segment 8. (Abbreviations S8,
8th segment; Pr, proctiger; pg, pygophore; c, clasper (paramere)). — 18, side view of ovipositor; 19, dorsal aspect of oviposi-
tor. (Abbreviations t8, tergite 8; gx1, first gonocoxa; gol-go2, first and second gonapophysis; pr, proctiger). Scale lines 0.5

mm.

Rhagovelia papuensis Lundblad

Rhagovelia papuensis Lundblad, 1936: 28-30: type series
New Guinea, Erima, Astrolabe Bay and Stephansort and
East New Britain, Herbertshéhe [Kokopo near Rabaul]
in HNHM [not examined]; Andersen 1982: 152;
Polhemus & Polhemus 1988: 165.

Material. — Papua New Guinea, West New Britain, Von
River, Banaule near Dami, 21.iii.1990, I. Lansbury, 19 ap-
terous. Found in small numbers at Tamari Creek and
Balima River, Bialla where it was found in dense shade.
Andersen (1982) figures the apterous male of papuensis.
Polhemus & Polhemus (1988) include about twenty species
in the ‘papuensis group’, which includes also À. australicus
Kirkaldy, an australian endemic found in the N.
Queensland rain forest habitats.

Gerridae, Trepobatinae

Metrobatopsis flavonotatus Esaki
(figs. 20-26)

M. flavonotatus Esaki, 1926: 144-146: type series New
Guinea, Erima, Astrolabe Bay, in HNHM [not examined];
Hungerford & Matsuda 1959: 31-35 [lectotype d desig-
nated]; Matsuda 1960: 367-371, 628-629.

Material. — Papua New Guinea, West New Britain, Von
River, Banaule near Dami, 21.11.1990, I. Lansbury, 13d
19 apterous, 19 macropterous and immatures; New
Hanover, Lavonga Mission, small freshwater stream,
31.x.1989, R. N. B. Prior, 45 29 apterous and numerous

immatures.

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 20-22. Metrobatopsis flavonotatus Esaki. — 20, macropterous female; 21, apterous male, dorsal habitus; 22, apterous male,
side view. Scale line 1 mm.

20

Figs. 23-26. Metrobatopsis flavonotatus
Esaki, male. — 23, front leg; 24, antennae;
25, genital capsule side view; 26, genital
segment, ventral aspect.

LANSBURY: Strongylovelia of West New Britain

te
SS
TAS

26

Esaki (1926) and Hungerford & Matsuda (1959) de-
scribe the variation in colour pattern and general structural
features, the latter describing the male genitalia. The series
from West New Britain and New Hanover differ slightly in

the structure of the male genitalia.

Description. — Adult apterous. Male 2.34-2.64
mm long, maximum width 1.0-1.1 mm; females in-
cluding macropterous form with damaged wings (fig.
20) 2.6 mm long, 1.32 mm wide.

Coloration: Male. Head, pronotum and mesonotum
velvety black. Basal inner margin of head adjacent to
eyes reddish brown. Pronotum and mesonotum with a
prominent pale yellow longitudinal stripe, distal margin
of mesonotal stripe evanescent blue. Metanotum, ter-
gites, connexivum and metacetabulae velvety black with

bluish evanescent patches, very noticeable on metano-
tum and tergites. Genital segments black, proximally ir-
idescent blue, lateral margins pale yellow (figs. 21-22).
Pro and mesopleura pale yellow, distally the latter black
overlaid with iridescent blue. Lateral margins of ster-
nites blackish brown becoming paler ventrally. Genital
segment ventrally pale yellow, distally black. Process of
pygophore, suranal plate and paired processes shining
black. Antennae black, proximal third of first segment
pale yellow. The pale band is much more conspicuous
when the gerrid is alive, likewise the proximal pale yel-
low band on front femur. Trochanter of middle and
hind legs pale yellow, remainder of legs black.
Macropterous female (fig. 20) similar to male.
Pronotum with a yellowish patch antero-mesially.
Remnants of wings dark brown, venation black.

21

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Male structure. Antennal segments 1-4 (fig. 24)
39.5: 25: 11.5 : 14. The ratios agree with Hungerford
& Matsuda (1959); they are however, rather more ro-
bust than those of the the female, contrary to
Hungerford & Matsuda (1959). The inner median
and distal surface of first segment with an area of pit-
like structures. Front leg (fig. 23) femur proximally ro-
bust, narrowing distally, tibia with a very prominent
blunt pilose projection distally. Eighth segment long,
laterally slightly sinuate, but not constricted as figured
by Hungerford & Matsuda (1959), but does resemble
figure by Matsuda (1960). Segment dorso-anteriorly
with transverse striations (fig. 21). Ninth segment
(figs. 25-26) with a prominent curved process usually
ventrally but in some males the process is in a latero-
ventral position. Suranal plate with a pair of sharply
acuminate processes usually not visible (fig. 25).

Gerrinae

Tenagogonus (T.) kampaspe (Kirkaldy)

Gerris kampaspe Kirkaldy, 1900: 804: type series New
Guinea, Rigo and Kellesi in SNOW [not examined].

Limnometra kampaspe, Lundblad 1933: 371 [listed].

Tenagogonus kampaspe, Hungerford & Matsuda 1958: 386-
388 [redescription, locality Dilo Loria, vi, vii. 90];
Matsuda 1960: 212, 501 [species groups].

Material. — Papua New Guinea, Milne Bay Province,
Naura, lotic stony river, 29.ix.1989, R. N. B. Prior, 14
macropterous; West New Britain, Von River, Banaule near
Dami, 21.iii.1990, I. Lansbury, 32 apterous. Overall length
of male including wings 5.39 mm, apterous female between

5.8-6.0 mm long.

Limnometra monochroma Nieser & Chen

Limnometra monochroma Nieser & Chen, 1992: 21-23.

Material (type series). — Papua New Guinea, West New
Britain, Buluma nr Dami, rain water pit, 17.1.1989, R. N.
B. Prior, 26 19 (incl. 1d holotype) (OxUM); West New
Britain, Von River, Banaule near Dami, torrential habitat,

21.11.1990, I. Lansbury, 22 (OXUM).

Ptilomerinae

Ptilomera breddeni Hungerford & Matsuda
Ptilomera breddeni Hungerford & Matsuda, 1965: 451-454:

22

types New Guinea, SE Haveri; SE Paumenu River; Milne
Bay (SNOW); Andersen 1982: 183, 191, 211.

Material. — Papua New Guinea, West New Britain, Von
River, Banaule near Dami, 21.iii.1990, I. Lansbury, 16
macropterous, 105 109 apterous. Collected in small num-
bers on faster stretches and easily able to evade capture.

REFERENCES

Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera
Gerromorpha). — Entomonograph 3: 1-455, figs.

Esaki, T., 1924. On a new genus and species of the Gerridae
from Formosa. — Annals Entomological Society of
America 17: 228-230.

Esaki, T., 1926. The water-striders of the subfamily
Halobatinae in the Hungarian National Museum. —
Annales Musei Nationalis Hungarici 23: 117-164.

Hungerford, H. B. & R. Matsuda, 1958. The Tenagogonus -
Limnometra complex of the Gerridae. — University of
Kansas Science Bulletin 39: 371-457.

Hungerford, H. B. & R. Matsuda, 1959. Concerning the
genus Metrobatopsis Esaki with descriptions of a new spe-
cies (Heteroptera: Geriidae). — Bulletin Brooklyn
Entomological Society 44: 29-36.

Hungerford, H. B. & R. Matsuda, 1965. The genus
Ptilomera Amyot and Serville (Gerridae: Hemiptera). —
University of Kansas Science Bulletin 45: 397-515.

Kirkaldy, G. W., 1900. On some Rhynchota, principally
from New Guinea (Amphibicorisae and Notonectidae). —
Annali del Museo Civico di Storia Naturale G. Doria 20:
804-810.

Lundblad, O., 1933. Zur Kenntnis der aquatilen und semi-
aquatilen Hemipteren von Sumatra, Java and Bali. —
Archiv fiir Hydrobiologie, Supplement 12: 1-195, 263-
489, pls. 1-21, figs. 1-142.

Lundblad, ©., 1936. Die altweltlichen Arten der
Velidengattungen Rhagovelia and Tetraripis. — Arkiv for
Zoologi 28A: 1-63.

Matsuda, R., 1960. Morphology, evolution and a classifica-
tion of the Gerridae (Hemiptera-Heteroptera). —
University of Kansas Science Bulletin 41: 25-632.

Nieser, N. & P. Chen, 1992. Revision of Limnometra Mayr
(Gerridae) in the Malay Archipelago. Notes on Malesian
aquatic and semiaquatic bugs (Heteroptera), II. —
Tijdschrift voor Entomologie 135: 11-26, figs. 1-60, tab.
1

Polhemus, J. T. & D. A. Polhemus, 1988. Zoogeography,
ecology and systematics of the genus Rhagovelia Mayr

(Heteroptera: Veliidae) in Borneo, Celebes and the
Moluccas. — Insecta Mundi 2: 161-230.

Received: 17 March 1992
Accepted: 20 March 1992

I. LANSBURY

Oxford University Museum, United Kingdom

RHAGOVELIA OF PAPUA NEW GUINEA, SOLOMON
ISLANDS AND AUSTRALIA (HEMIPTERA-VELIIDAE)

Lansbury, 1., 1993. Rhagovelia of Papua New Guinea, Solomon Islands and Australia
(Hemiptera-Veliidae). — Tijdschrift voor Entomologie 136: 23-54, figs. 1-150, tables 1-26.
[ıssn 0040-7496]. Published 1 July 1993.

Three species of Rhagovelia are known from ‘New Guinea’. R. papuensis Lundblad and R. biroi
Lundblad are redescribed and figured. R. peggiae Kirkaldy is only known by the apterous female
from Moroka and remains unrecognised at present.

New species described from Papua New Guinea, R. priori sp. n.; R. hirsuta sp. n.; R. aureo-
spicata sp. n.; À. herzogensis sp. n.; R. crinita sp. n.; R. thysanotos sp. n. and R. caesius sp. n. No
records have so far been published of Rhagovelia from the Solomon Islands, R. browni sp. n.
described from Guadacanal; R. amnicus sp. n. and R. fulvus sp. n. described from Malaita.

I. Lansbury, Hope Entomological Collections, University Museum, Oxford, OX1 3PW,

United Kingdom.

Key words. — Papua New Guinea; Solomon Islands; Veliidae; Rhagovelia; key; new species.

The Rhagovelia (Veliidae) are highly specialised
pleuston bugs of lotic waters throughout most of the
tropics. Andersen (1982) describes the general mor-
phology and biology of Rhagovelia. Polhemus &
Polhemus (1988) discuss the relative merits of various
interpretations of the thoracic and abdominal struc-
ture and their interpretation is followed in this re-
view.

The Rhagovelia of the ‘northern’ region of Papua
New Guinea described in this review are from West
Sepik (Torricelli Mountains), Madang, Eastern
Highlands, Morobe, Oro or Northern, Milne Bay
and West Britain Provinces. Additionally series from
Malaita and Guadalcanal (Solomon Islands) are in-
cluded.

Lundblad (1936) described R. papuensis and R. bi-
roi from Madang and East New Britain Provinces.
Kirkaldy (1901, 1908) described R. peggiae and R.
australica from Papua New Guinea and Australia,
north Queensland, respectively. Polhemus &
Polhemus (1988) revised the genus from Borneo,
Celebes and the Moluccas adding 26 new species.
They divided the Rhagovelia of the region into eight
monophyletic infrageneric species groups using wing
venation, thoracic morphology of macropterous
forms and genitalia, and included some extralimital
species in their groups. Of the taxa included in this
review, R. papuensis, R. biroi Lundblad, R. amnicus sp.
n., À. browni sp. n., R. herzogensis sp. n. and R. caesius
sp. n. are known by both macropterous and the more
common apterous forms. The remaining species de-
scribed are only known by the apterous form.

The Papua New Guinea species are tentatively as-

signed to the ‘Polhemus’-groups (1988):

Papuensis-group: broadly characterised by the
greatly enlarged pronotum covering all but the poste-
rior margin of the mesonotum. Included species R.
papuensis Lundblad and R. priori sp. n. Additional
species from the Solomon Islands R. fulvus sp. n., R.
browni sp. n. and À. amnicus sp. n., and the australian
endemic R. australica Kirkaldy.

Bacanensis-group: Polhemus & Polhemus (1988)
suggest that this group is closely allied to the papuen-
sis-group. The forewing venation is similar to R. pa-
puensis. Included species R. herzogensis sp. n., and also
R. aureospicata sp. n., as the apterous males of À. her-
zogensis and R. aureospicata are similar. The females
are much easier to distinguish. The dark areas of the
dorsum of both sexes having prominent fields of
golden or silvery spicules.

Novacaledonica-group. Included species R. nova-
caledonica Lundblad, À. biroi Lundblad and possibly
R. hirsuta sp. n. may belong to this group. As defined
by Polhemus and Polhemus (1988) macropterous
forms have four closed cells and an adventitious fifth
cell on the forewing. The R biroi from Papua New
Guinea examined in this study agree in all respects
with Lundblad’s original description of the apterous
form, but the forewing venation differs considerably
from that figured by Polhemus & Polhemus (1988).

Rhagovelia crinita sp. n. and À. thysanotos sp. n. dif-
fer from other known species from Papua New
Guinea. The male front tibiae are explanate with a
cluster of long hairs / spines along the inner margin.
Neither species is known by the macropterous form.
Both have short pronotums compared with the me-

25

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

sonotum, the males are dorsally hairy and the genital
capsules are very large. The female distal tergites are
laterally variably pilose and the 8th and 9th abdomi-
nal segments are sharply depressed.

Rhagovelia caesius sp. n. Polhemus & Polhemus
(1988: 167) comment on this group in northern New
Guinea and mention another species from Negros in
the central Philippines. Data relating to material col-
lected by R. N. B. Prior suggests that R. caesius is
found in slow flowing habitats in shady areas. It is a
remarkable species which differs from other Papua
New Guinea Rhagovelia by the absence of prominent
projections on the hind femora of both sexes. Unlike
other species, caesius is sexually dimorphic. The males
dorsally are covered with long hairs, the females fine-
ly tumescent, lacking long hairs. Females are abnor-
mally flattened compared with males. The males rid-
ing on the females, such phoretic behaviour has not
been observed in other Rhagovelia species. There is
some doubt that caeszus can be included in the same
section of Rhagovelia s.s. as other species from this re-
gion of Papua New Guinea.

Kirkaldy (1901) described Rhagovelia peggiae,
‘Type’ apterous female from Central Province,
Moroko [sometimes Meroka] 1300 m, 7.xi.1893,
Loria. The type is thought to be in Genoa (MCSN). It
is described as being very hairy and pubescent.
Pronotum distinctly sutured from mesonotum, this
suggests that the mesonotum may be longer than pro-
notum. Anterior margin of prontum yellow other
than lateral margins. The hind femora are ‘blue-
greenish lustre’. The connexivum is much reflexed in-
wards, meeting about the apex of the 7th connexival
segment and continuing in contact up to the apex
[somewhat as in R. distincta Champion, except that
they do not diverge after meeting]. The description
suggests that peggiae shares some features with hirsuta
sp. n.

A key to species excluding R. peggiae is given which
will only work for apterous forms of both sexes. The
australian endemic À. australica Kirkaldy is included
in the key.

Key to the species of Rhagovelia of Papua New

Guinea, Solomon Islands and Australia

1. Pronotum covering all but a transverse narrow
band of mesonotum (figs. 39, 63, 69) .............. À

— Median pronotal length shorter than median
length of mesonotum (figs. 73, 84)

2. Pro-mesopleura and proepisternum bare lacking
fields of minute black spicules australica Kirkaldy

Areas of propleura, pro-meso and metasternum
and jugum or lateral margins of pro-meso and

metanotum with fields of black spicules ........... 3
Anterior lateral margins of pro-meso and meta-
notum with fields of black spicules ................... 4

Anterior lateral margins of pro-meso and meta-
notum lacking fields of black spicules
Pronotum anteriorly with a pale transverse band.
First antennal segment proximally pale, remain-
der black. Connexivum and tergites dark brown-
Black... en ee 5
Pronotum more or less uniformly coloured, pale
anterior band obsolete, antennae pale, connexi-
vann amel nensites pelle onsassonossosersoscens fulvus sp. n.
Sternites 2-5 with scattered black spicules, male
connexivum either slightly reflexed outwards or
almost erect (fig. 51). Female connexivum sinu-
ate EA) Re browni sp. n.
First two visible sternites with black spicules,
male connexivum anteriorly flattened, distally re-
flexed (fig. 39). Female connexivum reflexed over
res 5-7 (fig 47) cecconscconcecooaoes amnicus sp. n.
Males 2.88-3.48 mm long, male 7th sternite de-
pressed (fig. 34). Paramere truncate (fig. 34).
Female connexivum sinuate, distally convergent
(MIVD priori sp. n.
Males 3.48-3.76 mm long, male 7th sternite
evenly rounded (fig. 18). Paramere reniform (fig.
16). Female connexivum not convergent distally.

pirata papuensis Lundblad
Proepisternum, pro-meso and metasternum with
black spiculest tt Sie e 8

Proepisternum, pro-meso and metasternum lack-
ing black spicules
Pronotum anteriorly transversely orange, not
reaching inner margin of eyes. Eye length 1.5x
median pronotal length ............... biroi Lundblad
Pronotum anteriorly transversely orange, surpass-
ing inner eye margin. Eye length subequal to me-
dianspronorall Ener hirsuta sp. n.
Hind femora of both sexes lacking prominent
projections, not incrassate (figs. 141, 148). Body
size sexually dimorphic (males 2.32-2.52 mm
long, females 3.28-3.36 mm long. ..caesius sp. n.
Hind femora of both sexes variably incrassate
with one or more prominent ventral projections.
Males and females not sexually dimorphic ... .10

. Dark areas of dorsum with prominent fields of

adpressedheoldispieulesen ne re 11
Dark areas of dorsum lacking noticeable fields of
adpressed soldispicul Sp een 12

Figs. 1-11. Rhagovelia australica Kirkaldy. — 1, male dorsal habitus; 2, ibid female; 3, male external genitalia; 4, ibid female;
5-7 male, 8-10 female: 5, 9, hind leg; 6, 8, front leg; 7, 10, antennae; 11, male genital capsule. Figs. 1-4, scale line 1 mm, figs.

5-10 scale line 1 mm, fig. 11 scale line 0.5 mm.

24

LANsBURY: Papuan Rhagovelia

it
U)

25

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

11. Male front tibia not explanate medianly, inner
margin uniformly fringed with short hairs (fig.
102). Female connexivum not sinuate (fig. 95)...
II A arn treat aureospicata sp. n.

— Male front tibia explanate medianly, inner mar-
gin with a cluster of hairs/spines (fig. 119).
Female connexivum sinuate (fig. 118) ................
D tte Net a RE MS ce crinita sp. n.

12. Anterior pronotal margin transversely pale yellow
(figs. 106, 107). Male front tibia not explanate,
inner margin fringed with short hairs (fig. 109).
Female connexivum converging at 7th tergite
(Hp 107) ee herzogensis sp. n.

— Anterior pronotal margin transversely pale yellow
not reaching lateral margins (figs. 126, 132).
Male front tibia medianly explanate, inner mar-
gin with a cluster of hairs/spines (fig. 129).
Female connexivum sinuate, distally fringed with
Venyglomestiains| (hel 2)E rar. thysanotos sp. n.

SYSTEMATICS

Rhagovelia australica Kirkaldy
(figs. 1-11, tabs. 1-2)
Rhagovelia australica Kirkaldy, 1908: 783-784; Hale 1925:

5; Hale 1926: 203; Lundblad 1933: 286; Lundblad
1936: 25-26; Polhemus & Polhemus 1988: 165.

Material. — Australia: Queensland, Mt. Lewis NW of
Rumula, 1200 m., 6-15.vi.1966, G.A. & S.L. Samuelson
(BPBM) 32 6, 16 £ and 15 immature, all apterous.

Redescription

Adult apterous. — Males (fig. 1) 4-4.2 mm long,
maximum width 1.6-1.72 mm, females (fig. 2) 4.04-
4.4 mm long, maximum width 1.68-1.88 mm.

Coloration. — Male (fig. 1): dorsum bright yellow
and dark reddish brown-black. Head yellow to dark
reddish brown, basally almost invariably yellow.
Pronotum anteriorly narrowly black, mesially with a
broad dark brown-black transverse bar, usually nar-
rowly divided by a yellowish brown longitudinal
stripe. Anterior and posterior margins with scattered
brown pits. Adjacent to posterior pronotal margin, a
narrow black shining bar which may be the mesono-
tum. Metanotum yellow, mesially reddish brown-
black, posterior margin black. Tergites 1-4 either uni-
formly black or with varying areas reddish brown.
Tergites 5-7 yellowish brown, marginally variably
black. Eigth tergite black. Connexivum bright yellow
with margins dark reddish brown. Jugum, pro, meso

and metaepisterna, coxae, trochanters and part of fe-

Table 1. Proportion of antennal segments of Rhagovelia
australica Kirkaldy.

I Il II IV

Apterous male 60 35 37 29)
Apterous female 60 36 40 DIS

Table 2. Proportions of leg segments of Rhagovelia australi-
ca Kirkaldy.

Femur Tibia Tarsus I Tarsus II Tarsus III

d front leg 665) 6) 55 16 =

9 front leg 71 71 35 16 =

3 middle leg 113 86 6 37.5 45.5
2 middle leg 111 90 5 40 49
3 hind leg 88 96 4 8.5 18
2 hind leg 97 101 4 5 20

mora pale yellowish brown. Metasternum and 2nd
sternite black, sternites 3-8 reddish brown, distally
pale yellow. Pleural margins of sternites narrowly
margined black. Front femora (fig. 6) distally and
middle legs dark brown-black. Hind leg (fig. 5)
broadly dark reddish brown-black. Antennae (fig. 7)
dark reddish brown, Ist segment basally pale yellow,
ventrally more broadly pale.

Structure. — Male: Median pronotal length 2.23-
2.4x eye length. Pronotal width 1.35-1.44x median
length, posteriorly covering all but a narrow band of
‘mesonotum and between 6-9x length of metano-
tum. Tergites 1-6 subequal, 7th longer, 8th shorther
imam Jins 1-3 are 12.5 ¢ 152 De 128 2 12,1 es 12,11 =
13.7 : 22.7 : 18.5. Distal margins of 5-7 fringed with
short hairs, 8th pilose, genitalia distinctly hairy (fig.
3). Connexivum slightly raised, distal segment
fringed with short conspicuous hairs. Jugum clearly
distinct from antenniferous tubercles. Propleura, pro-
episternum and mesopleura without fields of minute
black spicules. Distal margins of propleura with a row
of black punctures, lateral margins with sparse semi-
erect hairs, upper margins of mesopleura with a field
of semi-erect brown hairs. Front femora with a vari-
able number of fine black spines along ventral mar-
gin, tibia sinuate. Hind trochanter with four or more
blunt nodules. Femora variably incrassate, proximal
ventral margin with a row of even-sized setae, medial-
ly with prominent acuminate and blunt projections,
distally becoming smaller. Inner margin of tibia with
a row of irregular spaced triangular setae. Male geni-
talia (fig. 11) capsule distally produced.

Figs. 12-25. Rhagovelia papuensis Lundblad. — 12-18, 23-24 male, 19-22, 25 female. 12, 13, 19, hind leg; 14, 20, front leg;
15, mid-femora; 16, genital capsule; 17, 22, antennae; 18, 21, external genitalia; 23, male macropterous; 24, male apterous;
25, female apterous. Figs. 12-15, 17-22 scale line 1 mm, fig. 16 scale line 0.5 mm, figs. 23-25 scale line 1 mm.

26

LANSBURY: Papuan Rhagovelia

27

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Coloration. — Female (fig. 2): Tending to resemble
male, dark brown-black pigmentation of tergites
sometimes more extensive than in males. Front femo-
ra (fig. 8) anterior dorsal third pale yellow, ventrally
pale coloration extending about two thirds femoral
length. Coxae and trochanters of all legs pale yellow-
ish brown.

Structure. — Female: Median pronotal length 2.27-
2.52x eye length. Pronotal width 1.37-1.46x median
length, posteriorly covering all but a narrow band of
‘mesonotum’. Pronotum at midline greatly exceeding
length of ‘mesonotum’ and between 8-10x length of
metanotum. Tergites 1-4 more or less subequal, 5-7
progressively slightly longer, 8th shorter than pro-
ceeding. Seventh tergite slightly depressed, proctiger
reaching beyond Ist gonocoxa (fig. 4). 1-8 are 12.6:
12,8 2 12,7 : 13352 1593 18:12 20.1 8 15.1, Nerziess
2-8 with conspicuous slightly flattened shining areas.
Pronotum, metanotum and tergites 1-6 with ad-
pressed dark brown hairs, especially noticeable on
pale areas of metanotum. Eighth tergite laterally pi-
lose. Connexivum anteriorly slightly reflexed out-
wards, converging and becoming more erect and pi-
lose posteriorly, upper margins shining black. Venter
similar to male. Front femora (fig. 8) with a variable
number of fine spines ventrally. Tibia slightly sinuate.
Hind femora (fig. 9) moderately incrassate, spinose
distad ventrally. Tibia straight, not spinose along in-
ner margin. Antennae (fig. 10) Ist segment curved,
remaining segments straight.

Remarks. — Described from Queensland, Kuranda,
apterous female type in BPBM. Hale (1925) lists
Malanda, Lundblad (1936) briefly commented on
the macropterous form from Kuranda. This species
has so far only been recorded from the Atherton
Tablelands region near Cairns, Queensland.

The apterous form is immediately recognisable by
its striking dorsal bright yellow-brown-black pattern
which is rather variable, especially the spread of the
dark pigmentation. The male genital capsule is distal-
ly turned upwards, parameres elongate almost par-
allel-sided (fig. 11). Female with two (1+1) clusters of
black hairs along the distal margin of the 8th seg-

ment.

Rhagovelia papuensis Lundblad
(figs. 12-25, tabs. 3-4)

Rhagovelia papuensis Lundblad, 1936: 28-30; Polhemus &
Polhemus 1988: 170-173.

Material. — Papua New Guinea: West Britain: Tamari
Creek, 300 m., 14.vii.1989, R. N. B. Prior, 126, 129 ap-
terous, 12 macropterous; Von River, Banaule, torrential
stream, 21.iii.1990, I. Lansbury 1 ® apterous; Bialla, Balima
River, 3.111.1989, in dense shade, R.N.B.P. 83, 132 apter-

28

Table 3. Proportions of antennal segments of Rhagovelia

papuensis Lundblad.
I II III IV

Apterous male 42 26 27

26.5
Apterous female 41 29 31 31
Macropterous male 42.5 25 30 27
Macropterous female 43 26 30 26

Table 4 Proportions of leg segments of Rhagovelia papuensis
Lundblad.

Femur Tibia Tarsus I Tarsus II Tarsus III

Apterous

d frontleg 47 49 3.5 13.5 -

2 frontleg 48 50 3 14 =

3 middle leg 77 55 3 23 36

9 middle leg 81 57 4 23 36
3
3

3 hind leg 70 66

® hind leg 70 71 6 16
Macropterous

3 frontleg 50 54 3 12.5 =

9 frontleg 51 52 4 15 =

3 middle leg 83 60 3 22 38
9 middle leg 81 55 4 23 39
d hind leg 76 72, 3 7 16.5
2 molles 70. 70 3 7 16

ous, 1d 19 macropterous.

Redescription

Adult apterous. — Males (fig. 24) 3.32-3.48 mm
long, maximum width 1.2-1.28 mm, females (fig. 25)
3.48-3.76 mm long, maximum width 1.2-1.4 mm.
Adult macropterous. Male (fig. 23) 4.12 mm, maxi-
mum width 1.52 mm, female 3.88 mm long, maxi-
mum width 1.48 mm.

Coloration. — Male (fig. 24): Dorsum head, most
of pronotum, meso, metanotum and basal tergites
black. Remainder of tergites dark brown, 7th tergite
basally faintly shining. Anterior margin of pronotum
broadly orange and tending to widen laterally.
Propleura faintly bluish black over orange coloration.
Posterior pronotal margin narrowly orange. Con-
nexivum broadly margined pale brown. Mesopleura
and metapleura bluish black evanescent. Metaste-
rnum and sternites 2-6 laterally black, ventrally
pitchy red, 7th sternite pale yellow. Coxae and tro-
chanters of all legs pale yellow. Front femora (fig. 14)
pale yellow, apically dark brown, tibia and tarsi of
front leg and all middle leg greenish black iridescent.
Hind femora (fig. 12) dark brown-black dorsally,
ventrally pale yellow, tibia and tarsi dark brown-
black. Antennae (fig. 17) basal segment broadly pale
yellow, remaining segments black.

Structure. — Male: Median pronotal length just
over 2x eye length and covering most of ‘mesono-

Figs. 26-27. Rhagovelia spp. dorsal habitus. — 26-27 priori
sp. n., 26, male apterous; 27, female apterous. Scale as for

fig. 25.

tum’. Pronotal width 1.4x median eye length.
Tergites 1-5 subequal, 6th slightly longer, 7th longer
and shining, 8th shorter:10 : 11:12:11:11:13:18:
14.5. Tergites and connexivum with adpressed brown
hairs, denser and longer along margins of 6th tergite
and tip of connexivum. Genital capsule (fig. 16).
Jugum barely separated from antenniferous tubercles.
Propleura, proepisternum, meso and metasternum
and jugum with fields of black spicules. Ventrites 2-4
carinate, laterally sternites bare, 6th with scattered
brown adpressed hairs. Front femora (fig. 14) finely
spinose, middle femora (fig. 15) dorso-ventrally flat-
tened, finely spinose. Hind femora (fig. 12) ventrad-
distally spinose, tibia sinuate inner margin distally
with prominent triangular projections.

Coloration. — Female (fig.25): Head and thoracic
coloration similar to male. Connexivum more broad-
ly margined pale yellowish brown. Antennae, front
and middle legs similar to male, hind leg (fig. 19)
dark brown on all surfaces.

Structure. — Female: Black denticules on propleu-
ra, pro-meso and metasternum more prominent.
Tergites 1-3 reflexed downwards, 1-4 pilose, 5-8 with
large shining areas, 10): 10:9): 11 : 14:15:16: 13.
Proctiger truncate and slightly depressed (fig. 21).
Connexivum partially raised converging along mar-
gins of tergites 1-3, more elevated distally and con-
verging at tergite 8. Ventrites with fields of shining

LANSBURY: Papuan Rhagovelia

semi-erect hairs covering median areas of sternites 2-
3 gradually decreasing in length and density distally,
lateral margins of sternites with similar hairs increas-
ing in length and density distally. First visible sternite
obtusely raised. Hind femora moderately incrassate
with large spine ventrad, tibia straight, not spinose
along inner margin.

Macropterous male (fig. 23). — Similar to apterous
male, pronotum lengthened caudad, posterior margin
orange yellow. Wings extending well beyond the end
of abdomen with four closed cells, two in distal half.
Wings brown, venation slightly darker. Hind femur
(fig. 13) not as incrassate as apterous male.

Macropterous female. — Similar to male. Venation
more conspicuous especially subcosta and radius, in-
ner margins near anal vein much paler.

Remarks. — Described from Papua New Guinea,
Astrolabe Bay, (Stephensort and Erima) and Ralum
in New Britain. Type series in HNHM. The enlarged
pronotum, ventral conspicuous fields of black spic-
ules and shape of the male parameres shows a close re-
lationship with species from the Solomon Islands.
The female connexivum is more or less elevated its
entire length (fig. 25) and the shining distal tergites
are visible. Rhagovelia papuensis males are consistently
smaller than À fulvus sp. n. and browni sp. n.
(Solomon Islands), papuensis is superficially similar to
priori sp. n.

Rhagovelia priori sp. n.
(figs. 26-38, tabs. 5-6)

Type material. — Holotype male: Papua New Guinea,
Oro Province, Popondetta, Ambogo Plantation, small
stream, 29.iv.1989, R. N. B. Prior (OXUM). Paratypes 36,
149, same data; 128 99 P. N. G., Oro Prov. Popondetta,
Iseveni, small stream, 2.v.1989, R. N. B. P.; 12 P. N. G.
Oro Prov. Popondetta, Sambogo River, 30.iv.1989, R. N.
B. P. (OXUM).

Description

Adult apterous. — Males (fig. 26) 2.88-3.48 mm
long, maximum width 1-1.6 mm, females (fig. 27)
3.2-3.48 mm long, maximum width 1.2-1.24 mm.

Coloration. — Male (fig. 26): Head most of thorax
dorsally black. Tergites greyish black, 7th tergite and
lateral margins of connexivum shining pale brown.
Anterior margin of pronotum pale yellow, laterally
rather wider. Propleura dark grey merging into red-
dish brown. Meso and metapleura greyish black. Pro,
meso and metasternum and sternites 2-3 bluish grey,
coxal cavities tending to be tinged reddish brown.
Sternites 3-5 reddish brown, 6th paler. Coxae and
trochanters pale yellow. Front femora with extensive
pale yellow coloration especially along inner margin.
Remainder of front and all middle legs dark shining

29

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

brown. Hind femur shining dark brown with lateral
pale areas same colour as trochanter along ventral
margin. Antennae (fig. 33) 1st segment basally pale
yellow, ventrally surface more broadly yellow, re-
maining segments black.

Structure. — Male: Median pronotal length 2x eye
length and covering almost all mesonotum. Pronotal
width 1.44x median length and 5-6x length of meta-
notum. Tergites 1-5 subequal, 6 and 8 slightly long-
er, 7th longest with a variably shaped shining area; 9 :
OPO O EDS ERD Nhoracıerdorsum
covered with fine golden hairs. Shining lateral con-
nexival margins and tergites 1-6 with longer ad-
pressed hairs. Connexivum either flat or slightly
raised distally with tufts of hairs. Jugum very close to
antenniferous tubercles. Propleura, pro, metaepister-
num and jugum with fields of minute black spicules.
Sternites anteriorly with a median keel with pale yel-
low hairs overlying fine adpressed hairs. Seventh ster-
nite depressed posteriorly, lateral margins obtusely
carinate and pilose, 8th with a conspicuous keel (fig.
34). Front leg (fig. 30) mid femora ventrally with two
rows of fine hairs (fig. 31). Hind femora variably in-
crassate (figs. 28, 29). Ventral margin variably spinose

Table 5. Proportions of antennal segments of Rhagovelia
priori sp. n.

I II III IV
Apterous male 371 22 26 26
Apterous female 37 DI 24 24

Table 6. Proportions of leg segments of Rhagovelia priori sp.

n.

Apterous Femur Tibia Tarsus I Tarsus II Tarsus III
3 frontleg 40 45 3 11 =

9 frontleg 41 44 3 12 =

3 middle leg 71 53 4 Dil 33

9 middle leg 72 Si, A 27 35

d hindleg 67 COS 5 14

® hindleg 59 60 2 5 14

with many fine setae, distally with triangular setae.
Tibia either straight or slightly sinuate with triangular
setae along inner margin, length increasing distally.
Genital capsule (figs. 32, 33), distal lateral aspect of
7-8th segments (fig. 34).

Figs. 28-38. Rhagovelia priori sp. n. — 28-35 male, 36-38 female. 28, 29, 36 hind leg; 30, front leg; 31, mid-femora; 32, 33,
genitalia; 34, 37, external genitalia; 35, 38, antennae. Figs. 28-31, 34-38 scale line 1 mm, figs. 32-33 scale line 0.5 mm.

30

Coloration. — Female (fig. 27): Head and pronotal
coloration similar to male. Mesonotum postero-late-
rad sometimes with brown shining areas. Tergites 5-7
sometimes shining tending to resemble À. papuensis
(fig. 25). Upper margins of connexivum narrowly
shining brown. Legs and antennae similar to male oth-
er than hind femora which tend to be paler ventrally.

Structure. — Female: Median pronotal length about
2x eye length. Mesonotal length variable, pronotum
about 5-7x length of mesonotum. Metanotum slight-
ly longer than mesonotum. Tergites 1-4 subequal, 5-
7 progressively longer, 8th short; 1-8 are: 10: 10: 10
: 10: 12:17:20: 8. Connexivum anteriorly reflexed
outwards with fields of hairs on inner margins. From
4th tergite abruptly turned inwards and converging
distally. Upper distal margins pilose and 8th tergite
densely pilose. Underside of thorax similar to male.
Anterior sternites with clusters of golden hairs me-
dianly. Proctiger reflexed downwards (fig. 37). Hind
femur (fig. 36) moderately incrassate with a promi-
nent curved spine in distal third. Hind tibia straight
without prominent setae along inner margin.
Antennae (fig. 38) first segment slightly more curved
than male (fig. 33).

Etymology. — A noun in apposition. This species is
dedicated to Mr. R. N. B. Prior who has collected
aquatic Hemiptera-Heteroptera from various local-
ities in Papua New Guinea, especially West New
Britain.

Remarks. — Rhagovelia priori sp. n. is superficially
much like papuensis. The apterous females of former
are easily distinguished by stronger convergent con-
nexiva with fields of hairs along anterior inner mar-
gins and prominent distal clusters of hairs. Males of
priori tend to be smaller than papuensis (2.88-3.48 :
3.32-3.48 mm). Genital capsule of priori distally acu-
minate, of papuensis distally convergent and rounded.
Parameres of priori slightly sinuate, of papuensis renif-
orm.

Rhagovelia amnicus sp. n.

(figs. 39-50, tabs. 7-8)

Type material. — Holotype male: Solomon Islands,
Malaita, Raiako Nafinua, in forest stream c 3 yards wide,
much shaded, fast running with shingle bottom, 27.v.1955,
E. S. Brown (BMNH). — Paratypes: same data as holotype,
186 409; same data, except ‘larger stream with a few slow
flowing areas’, 3d 72 (BMNH, OXUM).

Additional material. — Solomon Islands: Malaita: River
Aluta just west of Maelegwasu, in forest. A rather large rap-
id river with shingle bottom, 26.v.1955, E. S. B., 136 122
apterous, 1d 19 macropterous (BMNH, OXUM); same, in-
land east of Kwailasi. Small running stream shaded in forest,
25.v.1955, E. S. B., 196 132 (BMNH, OXUM); same, near
Kwailasi, large rocky stream, 25.v.1955, E. S. B., 106 62 (-
BMNH, OXUM).

LansBury: Papuan Rhagovelia

Description

Adult apterous. — Males (fig. 39) 3.32-3.96 mm
long, maximum width 1.16-1.44 mm, females (fig.
47) 3.16-3.84 mm long, maximum width 1.22-1.44
mm. Adult macropterous. Male (fig. 46) 4.12 mm
long, maximum width 1.58 mm, female 4.12 mm
long, maximum width 1.68 mm.

Coloration. — Male (fig. 39): Ground colour black,
anterior pronotal margin pale yellow, not reaching
lateral margins. Tergites dark brown-black, 7th ter-
gite shining. Connexivum narrowly shining pale
brown anteriorly. Legs mostly black. Coxae, tro-
chanters, proximal margins of front tibia and inner
proximal margin of-hind femur pale yellow. Jugum
brown, pro-meso and metasternum black. Sternites
dark reddish brown, laterally black. Antennae black,
proximal 0.2-0.25 of first segment pale yellow.

Structure. — Male: Median pronotal length 2.2x
eye length, posteriorly converging most of mesono-
tum. Pronotal width 1.47x median length.
Metanotum 2x length of mesonotum. First tergite
shorter than remainder, tergites 2-6 subequal, 7th
longest, 8th shorter and densely hairy. 1-8 are 10: 14:
SENS EME SE 2507 Wereitese2-o1and! basal
margin of connexivum with many pale hairs, 7th
sparsely pilose, distal margin fringed with longer
hairs. Connexivum anteriorly slightly flattened, be-
coming more reflexed upwards distally. Anterior lat-
eral pronotal margins, lateral margins metanotum,
and first two tergites with fields of black denticles.
Jugum barely separated from antenniferous tubercles.
Propleura, proepisternum and mesopleura with a sin-
gle row of deep black punctures, upper margins with
prominent erect hairs. Meso and metasternum with
scattered black denticles, overlying them long semi-
erect fine yellowish hairs. First two sternites with
black denticles. Anterior sternite bluntly acuminate.
Anterior sternites with long fine hairs, distal sternites
with short hairs which increase in density distally.
Seventh sternite depressed posteriorly, lateral margins
with tufts of hairs (fig. 39). Genital segment ventrally
carinate with a narrow fringe of fine hairs, laterally
smooth and shining (fig. 40) parameres different as-
pects (figs. 44-45). Front femora with a variable num-
ber of fine spines along ventral margin, tibia straight
(fig. 41). Hind trochanter with circa 5 blunt nodules.
Femur incrassate, proximal ventral margin with a row
of even-sized setae, medially with a prominent spine,
distally with irregular smaller spines in two rows.
Tibia sinuate, inner margin armed with triangular se-
tae, distally with a prominent bifurcate projection
(fig. 42).

Coloration. — Female (fig. 47): Resembling male,
pronotal anterior yellow band more intense. Tergites
5-8 with shining elongate areas. Propleura dark yel-
lowish brown. External connexival margins anteriorly

31

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 39-50. Rhagovelia amnicus sp. n. — 39-46 male, 47-50 female. 39, 47, dorsal habitus; 40, genital capsule; 41, tront leg;
42, 49, hind leg; 43, 50, antennae; 44, 45, parameres; 46, macropterous dorsal habitus; 48, external genitalia. Figs. 39-43, 46,
47, 49-50 scale line 1 mm, figs. 44-45 scale line 0.1 mm, fig. 48 scale line 0.5 mm.

broadly yellow, upper margins shining brown, distal- Structure. — Female: Median pronotal length 2.5x
ly dark brown. Sternites covered with bluish silvery eye length, pronotal width 1.4x median length, poste-
pubescence. Legs and antennae similar to male, but riorly covering most of mesonotum, the latter 0.1 1x
front tibia are more extensively yellow. pronotal length. Metanotum 2.2x length of mesono-

32

Table 7. Proportions of antennal segments of Rhagovelia
amnicus sp. n.

I II Il IV

Apterous male 43 28.5 29 27
Apterous female 43 26 27.5 25
Macropterous male 42 26 28 Dif
Macropterous female 43 26 28 26

Table 8 Proportions of leg segments of Rhagovelia amnicus
sp. n.

Femur Tibia Tarsus Tarsus Tarsus

Apterous

d front leg 53 53 3 13 =

2 front leg 49 49 4 14 =

3 middle leg 88 63 5 26 38

9 middle leg 79 57 4 25 37

d hind leg 81 64 4 5) 16

9 hind leg 70 65 DIS) 6 16

Macropterous

d front leg 52 56 4

? front leg 54 57 5}

3 middle leg 90 65 >) DS) 40
9 middle leg 85 66 5

3 hind leg 89 70 4

9 hind leg 75 75 3

tum. Lateral margins of pronotum, meso and meta-
notum with fields of black denticles. Tergites 1-3 de-
pressed, tergites 4-7 darker than preceeding, tergites
5-7 mostly shining. Tergites 1-2 with scattered long
black semi-erect hairs. 1-8 are: 11:9:8.5:12:11:
13 : 26 : 10. Connexivum anteriorly raised almost
erect, thickened and shining and sharply converging
along 3rd tergal margin, curving and strongly reflexed
over tergites 5-7, sometimes posteriorly completely
covering distal tergites. Upper distal margins densely
hairy, posteriorly with a prominent tuft of hairs,
proctiger reflexed downwards (fig. 48). Lower margin
of jugum, proepisternum and mesosternum with
fields of black denticles. Sternites faintly rugose, 7th
shining, laterally and distally fine yellowish pilose.
Hind femur moderately incrassate, ventrally spinose,
tibia not armed along inner margin (fig. 49).

Macropterous male (fig. 46). — Pronotum at widest
subequal to median length. Caudad depressed, hu-
meral angles slightly elevated. Forewings brown, ve-
nation black with four cells. Membrane of elytra just
surpassing genital segment. Hind femur incrassate.

Macropterous female. — Similar to male, mem-
brane rather longer than in male exceeding proctiger
and 8th segment.

Etymology. — Name derived from ‘amnis’ stream
of water, amnicus ‘inhabitant of stream’.

LANSBURY: Papuan Rhagovelia

Remarks. — This species is similar to À priori;
males differing by not having lateral margins of 7th
sternite laterally obtusely carinate. Genital capsule
truncate distally. Females have connexivum and distal
abdominal segments are much more pilose. The ante-
rior pronotal yellow band not usually reaching lateral
margins.

Rhagovelia browni sp. n.

(figs. 51-62, tabs. 9-10)

Type material. — Holotype d: Solomon Islands,
Guadacanal, Sorvohio River near Gold Ridge, rapid water
over boulders with calm pools, small shaded river,
20.11.1955, E. S. Brown (BMNH). — Paratypes: same data as
holotype, 64, 62 apterous; same data except, Gold Ridge,
1000-1500 feet, clean fast running water with boulders
mainly shaded by forest, 25.vi.1956, E. S. B., 174 69 ap-
terous, 14 29 macropterous; same data as preceeding,
30.vi.1956, E. S. B., 36 52 (BMNH, OXUM).

Additional material. — Solomon Islands, Guadacanal:
Suta, Sutakiki River, c. 2300 feet, 26.vi.1956, 46 99 apter-
ous, 1 2 macropterous; Sangava River near Suta. Strong sto-
ny trickling stream, mostly rather shaded, 27.vi.1956, 4d
5; Kokumbona River 13.v.1956, 148 59 apterous, 1d
macropterous; Tsarivonga River, rapid stony river with
quiet backwaters, 1000 feet, 22.iii.1955, 198 139 apter-
ous, 1d 19 macropterous; Tenaru River. Between Mission
and Tenaru Falls, mostly from comparatively calm water
along margin and in backwaters, 7.xi.1954, 4d 42 apter-
ous, 26 12 macropterous; Matanikao, pools in rocks
downstrem of Matanikao Caves, shaded 31.x.1954, 154
6% apterous; Honiara, Matanikoa River, 6d 29 apterous,
16 macropterous (all leg. E. S. Brown) (BMNH, OXUM);
Guadacanal, Gold Ridge 600 m. 22.vi.1956, J.L. Gressitt,
1d apterous; Santa Ysabel, Tamatahi-Molao Maringe Distr.
28.vi.1960, Jungle track C.W. O’Brien, 18 49 apterous,
2% macropterous (BPBM).

Description

Adult apterous. — Male (fig. 51) 3.88-4.12 mm
long, maximum width 1.36-1.44 mm, females (fig.
59) 4.12-4.24 mm long, maximum width 1.48-1.52
mm. Adult macropterous. Males (fig. 52) 4.64 mm
long, maximum width 1.72 mm, females 4.52-4.8
mm long, maximum width 1.72-1.76 mm.

Coloration. — Male (fig. 51): General dorsal color-
ation dark brown-black. Head black, anterior margin
of pronotum yellow, lateral margins bluish black or
reddish brown. Seventh tergite and genital segment
shining black. Jugum, coxae and trochanters yellow-
ish brown. Sternites dark reddish brown, 7-8th ven-
trites pale yellow. Pro, meso and metapleura reddish
brown. Lateral margins of sternites black. Antennae
black, basal fifth of lst segment pale yellow, dorsally
dark brown. Middle and hind legs dark brown-black.

Structure. — Male: Median pronotal length 2.36x
eye length. Pronotal width 1.44x median length, pos-
teriorly covering almost all mesonotum, metanotum
0.2x length of pronotum. Tergites 1-6 more or less

33

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

34

Table 9 Proportions of antennal segments of Rhagovelia
browni sp. n.

I Il II IV

Apterous male 52 31 35 30

Apterous female 47.5 28 32 28
Macropterous male 44 ITS 30 26
Macropterous female 48 30 32 30

Table 10. Proportions of leg segments of Rhagovelia browni
sp. n.

Femur Tibia Tarsus I Tarsus II Tarsus III

Apterous

d front leg 60 67 4 16.5 —

ironies DISS 5 18 -

3 middle leg 105 76 5 31 42
9 middle leg 91 67 5 28 41
3 hind leg OSG 4 8.5 20
9 hind leg 82 83 4 8 20
Macropterous

d front leg I Dee 19:5 =

2 front leg SMS 5 17 =

d middle leg 88 64 5 DI 40
9 middle leg 92 70 5 28 4]
3 hind leg DT Te 7 19
9 hind leg 84 86 4 9 20

subequal, dull, lateral margins finely pilose and distal
margins fringed with hairs. Seventh longest, shining,
transversely rugose, posterior margin fringed with
fine hairs. Genital segment (8th) shining, covered
with fine hairs, denser along lateral margins. 1-8 are
13:13:14.5:14:15:12: 23 : 20. Anterior lateral
margins of pronotum, meso and metanotum with
fields of black spicules. Connexivum either slightly
reflexed outwards or almost erect, upper margins
shining dark brown. Propleura anteriorly with a scat-
tered field of black spicules, posterior margin with
several short continguous rows of punctures. Meso
and metapleura with short black hairs directed cau-
dad. Jugum almost touching antenniferous tubercles.
Lateral margins of jugum, proepisternum, meso and
metasternum with prominent fields of black spicules.
Sternites 2-5 with scattered black spicules, 2nd ven-
trite obtusely carinate. Seventh sternite depressed to-
wards gentital segment. Latero-distal margins pilose.
Genital segment carinate and fringed with short hairs.
Genital capsule (fig. 53) parameres different aspects (-

LANSBURY: Papuan Rhagovelia

figs. 56-57). Front leg (fig. 54). Hind leg (fig. 55) fe-
mur incrassate, trochanter with 3-4 blunt nodules.
Femoral ventral margin anteriorly with two rows of
uniform sized setae, posteriorly with larger spinose
projections. Tibia sinuate with two rows of blunt
spines, medially with a large spine, distally with a
large spine.

Coloration. — Female (fig. 59): Resembling male,
pronotum black other than yellowish brown anterior
margin reaching inner margin of eyes. Tergites 5-8
shining. Upper margins of connexivum shining red-
dish brown. Pro-meso and metapleura varying
between dark reddish brown and bluish black.
Jugum, coxae and trochanters pale yellowish brown.
Meso and metaepisternum and sternites 2-6 dark red-
dish brown, lateral margins dark brown-black, 7th
sternite shining pale reddish brown. Antennae black,
Ist segment basally and ventrally pale yellow. Front
femur ventrally pale yellow, tibia black. Middle leg
dark reddish brown. Hind leg (fig. 61) femur proxi-
mally pale yellow, remainder of leg reddish brown.

Structure. — Female: Median pronotal length 2.4x
eye length. Pronotal width 1.5x median length, pos-
teriorly covering all but a narrow strip of mesonotum.
Metanotum 2x length of mesonotum. Tergites 1-3
subequal, 4-7 increasing in length, 8th shorter. 1-8
TR RS oS) o ASIN 5) 2 OE SOS hinne
areas of tergites 5-8 increasing in size distally and
faintly transversely rugose. Distal margin of 8th ter-
gite fringed with long hairs. Anteriorly connexivum
reflexed outwards, along margins of tergites 3-4 more
erect, distally slightly reflexed outwards forming a sin-
uate line, connexivum distally pilose. Lateral margins
of mesonotum with scattered black spicules, metano-
tum with larger fields. Propleura and thorax ventrally
similar to male. Sternites with black spicules, not car-
inate. Seventh sternite shining, laterally with fine yel-
lowish hairs. Genital segment more pilose, connexival
hairs almost reaching end of genital segment (fig. 62).
Hind femur moderately incrassate, spinose posterior-
ly, tibia straight, inner margin not spinose.

Macropterous male (fig. 52.). — Pronotum anteri-
orly yellow, lateral margins bluish black. Pronotal
width subequal to median length, humeral angles and
posterior tip depressed with scattered black punc-
tures, pronotal disc raised. Forewings brown, vena-
tion black with four closed cells, anteriorly inner mar-
gin of anal vein narrowly pale yellow. Membrane of
elytra surpassing end of abdomen and genitalia.
Coloration of legs and antennae similar to apterous

Figs. 51-62. Rhagovelia browni sp. n. — 51-58 male, 59-62 female. 51, dorsal habitus; 52, dorsal habitus macropterous; 53,
genital capsule; 54, front leg; 55, hind leg apterous; 56-57, parameres; 58, antenna; 59, dorsal habitus apterous; 60, hind leg
macropterous; 61, hind leg apterous; 62, external genitalia. Figs. 51-55, 58-61 scale line 1 mm, figs. 56-57 scale line 0.1 mm,

fig. 62 scale line 0.5 mm.

55

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

form. Hind leg (fig. 60) femur moderately incrassate,
tibia straight, inner margin with triangular setae.

Macropterous female. — Similar to macropterous
male, membrane slightly shorter just surpassing the
Ist gonocoxa.

Etymology. — This species is dedicated to the late
E. S. Brown, an enthusiastic hemipterist who collect-
ed extensively in the Solomon Islands.

Remarks. - Comparing females of À. browni sp. n.
it is rather similar to Rhagovelia fulvus sp. n., browni
has a sinuate connexivum whereas the connexivum of
fulvus is more or less straight. Rhagovelia browni is a

(ANN
AAN
A |

much darker species; parameres elongate compared
with the moderately reniform parameres of fulvus.

Rhagovelia fulvus sp. n.
(figs. 63-72, tabs. 11-12)

Type material. — Holotype d : Solomon Islands, Malaita,
Fulisano-Maelegwasu, 26.v.1955. Small spring near its
source, just west of main water-shed of island, E. S. Brown
(BMNH). — Paratypes: same data as holotype, 76 5 (BMNH,
OXUM).

Description
Adult apterous. — Male (fig. 63) 3.64-3.8 mm long,

Figs. 63-72. Rhagovelia fulvus sp. n. — 63-68 male, 69-72 female. 63, 69, dorsal habitus apterous; 64, genital capsule; 65 par-
amere; 66, front leg; 67, 70, hind leg; 68, 71, antennae; 72, external genitalia. Figs. 63, 66-71 scale line 1 mm, fig. 64, 72,

scale line 0.5 mm, fig. 65 scale line 0.1 mm.

36

Table 11. Proportions of antennal segments of Rhagovelia

fulvus sp. n.

I Il III IV
Apterous male 50 29.5 33 29
Apterous female 50 31 32.5 29
Table 12. Proportions of leg segments of Rhagovelia fulvus
sp. n.

Femur Tibia Tarsus I Tarsus II Tarsus III

Apterous

6 frontleg 55 63 3 16 -
9 frontleg 55 58 3.5 15 =
3 middle leg 94 70 5 26 39
2 middle leg 90 64 5 26 40
3 hind leg 92 85 3 8 19
Sihindiles 79786 3 9 16

maximum width 1.28-1.4 mm, female (fig. 69) 3.76-
4.04 mm long, maximum width 1.36-1.44 mm.

Coloration. — Male (fig. 63): Head dorsally dark
reddish brown. Pronotum either pale orange yellow
or pinkish brown. Anterior lateral and medianly rath-
er paler. Meso, metanotum and tergites 1-6 varying
between dark yellowish and reddish brown.
Connexivum and tergites 7-8 pale yellow. Propleura
pale reddish brown, meso and metapleura slightly
darker. Sternites dark reddish brown, 7th sternite
faintly shining, 7th and 8th sternites pale yellowish
brown. Coxae and trochanter pale yellow. Femur and
tibia of all legs pale yellow, tarsi tending to be dark
yellowish brown. First and 2nd antennal segments
pale yellow, remainder brown.

Structure. — Male: Median pronotal length 2.56x
eye length, covering almost all mesonotum. Pronotal
width 1.46x median length. Anteriorly pronotum
with a transverse row of punctures, lateral margins
with a small field of black spicules, sometimes spic-
ules much paler and difficult to see. Metanotum
about 2x length of mesonotum. Tergites gradually in-
creasing in length distally, 1-8 are 10 : 13:12:12:
13 : 14:19:17. Connexivum more or less uniform-
ly semi-erect. Lateral and posterior margins of prono-
tum with fine silvery adpressed hairs, remainder of
pronotum covered with shorter hairs. Posterior mar-
gin of metanotum, tergites and lower margin of con-
nexivum fringed long pale yellowish hairs. Lateral
margin propleura, margins of jugum, proepisternum,
meso and metasternum with fields of black spicules.
Jugum barely separated from antenniferous tubercles.
Sternites 2-6 with scattered black spicules and with
long fine adpressed silvery hairs becoming denser on
7th sternite. Front leg (fig. 66) moderately finely spi-
nose. Hind leg (fig. 67) femur incrassate, trochanter

LansBury: Papuan Rhagovelia

with a variable number of blunt nodules. Femur ven-
tro-anteriorly with short regular setae, medially with
2 prominent spines, distally. Tibia either straight or
slightly sinuate, inner margin armed with a regular
row of triangular setae. Antennae (fig. 68). Genital
capsule (fig. 64) paramere slightly reniform, paramere
from different aspect (fig. 65).

Coloration. — Female (fig. 69): Very similar to
male. Pronotum orange brown, anterior margin
sometimes paler. Pleura with a pinkish tinge. Tergites
dark yellowish brown with straggly yellow hairs, ter-
gites 5-8 with shining areas increasing in size distally.
Connexivum narrowly dark brown adjacent to ter-
gites, upper margins paler. Sternites laterally dark
brown, ventrally paler. Apices of femur narrowly an-
nulated black. First antennal segment pale yellow,
segments 2-4 pale brown.

Structure. — Female: Median prontal length 2.75x
eye length, covering all but narrow strip of mesono-
tum. Connexivum reflexed outwards, distally not
reaching distal margin of 8th tergite and densely pi-
lose. Pleural and ventral black spicules similar to
males second sternite with a denser field of black spic-
ules. Proctiger slightly raised, gonocoxal plates large
(fig. 72). Hind leg (fig. 70) femur not incrassate, ven-
tral margin with a prominent distal spine, tibia
straight, not armed along inner margin. Antennae
(fig. 71) first segment sometimes infuscated pale
brown, likewise segments 2-4.

Etymology. — Name refers to the tawny, reddish
yellow coloration.

Remarks. — Rhagovelia fulvus sp. n. is a very pale
species compared with amnicus and browni. The fe-
murs are almost entirely pale. The connexivum of the
female is shorter than in other species not reaching
the end of tergite 8.

Rhagovelia biroi Lundblad
(figs. 73-83, tabs. 13-14)

Rhagovelia biroi Lundblad, 1936: 13-14; Polhemus &
Polhemus 1988: 165.

Material. — Papua New Guinea: West Britain: Bialla,
Balima River, 3.iii.1989, R. N. B. Prior, 54 12 apterous;
Bialla, River Inu, 10.ii.1989, R. N. B. P., 154 172 apter-
ous; Bialla, River Tiaru, 10.1.1989, R. N. B. P., 106 79 ap-
terous; same data, near log in river, 25.1.1989, 3d 19 apter-
ous; Tamari Creek, 300 m., 14.vii.1989, R. N. B. P., 216
159 apterous; Dami near Kimbe, vii.1989, R. N. B. P., 2d
apterous; Bilomi River 9.iv.1989, R. N. B. P., 7d apterous;
Von River, Banaule Village near Kimbe, 21.11.1990, I.
Lansbury, 174 92 apterous, 1 ? macropterous (OXUM).

Redescription
Adult apterous. — Male (fig. 73) 3.2-3.48 mm long,
maximum width 1.16-1.28 mm, female (fig. 79)

SH

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1992

Figs. 73-83. Rhagovelia biroi Lundblad. — 73-78 male, 79-83 female. 73, 79, dorsal habitus apterous; 74, genital capsule; 75,
ventral aspect genital plate; 76, 82, antennae; 77, front leg; 78, 81, hind leg; 80, dorsal habitus macropterous; 83, external
genitalia. Figs. 73, 76-83 scale line 1 mm, figs. 74-75 scale line 0.5 mm.

3.24-3.48 mm long, maximum width 1.2-1.56 mm.
Adult macropterous. Female (fig. 80) 3.92 mm long,
maximum width 1.52 mm.

Coloration. — Male (fig. 73): Dorsum black, 7th
tergite with a triangular iridescent greenish black
patch. Pronotum anteriorly with an orange band not

38

reaching inner margin of eyes. Venter greyish black.
Coxal cavaties pale yellow. Antennae black other than
basal third of Ist segment. Basal third of front femur
pale yellow. Trochanters of front and hind legs pale
yellow, remainder of all legs greenish black iridescent,
most apparent on middle and hind femora.

Table 13. Proportions of antennal segments of Rhagovelia
biroi sp. n.

I II III IV

Apterous male 43 19 21 23
Apterous female 42 22 20 22
Macropterous female 46 22 17 DD)

Table 14. Proportions of leg segments of Rhagovelia biroi
sp. n.

Femur Tibia Tarsus] Tarsus II Tarsus III

Apterous

3 frontleg 48 53 4.5 13 -
9 frontleg 48 50 2.5 13 -
3 middle leg 83 56, 2 26 34
? middle leg 78 S20" 94 26 57
d hindleg 72 68 3 5 15
9 hindleg 72 75 2 6 16
Macropterous

9 frontleg 47 48 3 13 -
? middle leg 78 53. 1 4 26 37
9 hindleg 72 69 3 5) 16

Structure. — Male: Median eye length 1.5x median
prontal length. Pronotal width 4.8x median length.
Mesonotum large, 4.3x pronotal length. Metanotum
short, 0.3x pronotal length. Tergites 1-7 increasing in
length distally, 8th shorter. Shining area of 7th and all
of 8th with long brown hairs. 1-8 are7:9:9: 10.5:
11: 12:22:15. Dorsum clothed in fine pubescence.
Pronotum, meso-metanotum and Ist tergite with
short recumbent black setae. Remaining tergites with
longer greyish hairs. Connexivum slightly reflexed
outwards, upper margin narrowly shining brown.
Jugum clearly distinct from antenniferous tubercles.
Proepisternum with sparse minute black spicules dif-
ficult to see against background. Pro, meso and met-
asternum with scattered black spicules. Lateral mar-
gins of sternites 3-6 with two groups (1 + 1) of black
spicules either side of midline. Second sternite with a
dense area of black spicules. Venter clothed with long
recumbent greyish hairs, 7th ventrite more pilose,
medianly carinate with a fringe of semi-erect hairs.
Front leg (fig. 77) tibia flattened. Legs with self
coloured hairs and spines. Coxae with groups of
curved hairs. Hind leg (fig. 78) femora with a promi-
nent spine midway ventrad, further projections dis-
tally. Tibia slightly curved, inner margin armed with
small setae. Antennae (fig. 76) 1st segment with many
spines. Genital capsule (fig. 74) parameres elongate,
apically acuminate, Lundblad (1936) figure shows
apex more truncate. Ventral aspect of genital plate
(fig. 75).

Coloration. — Female (fig. 79): Very much like the

LANSBURY: Papuan Rhagovelia

male. None of tergites shining. Connexivum, upper
margin narrowly yellowish brown.

Structure. — Female: Median eye length 1.85x pro-
notal length. Pronotal width 5x median length.
Mesonotum large, 4x pronotal length. Metanotum
short 0.38x pronotal length. Tergites 1-7 increasing
in length distally, 8th shorter, 1-8 are 7:9: 10.5: 11
> 11.5: 14:18:10. Eighth tergite depressed, posteri-
or margin of 7th with a cluster of hairs. Connexivum
distally produced beyond 8th tergite and densely pi-
lose. Mesonotum and metanotum with minute black
shining spicules overlaid by long black hairs extend-
ing to Ist tergite along margins. Pro-meso and meta-
sternum with scattered black spicules. Seventh ster-
nite with dense areas of long golden hairs either side
of midline. Hind leg (fig. 81) femora ventrad-distally
with a bifurcate spinose projection, tibia straight, in-
ner margin unarmed. Antennae (fig. 82). Gonocoxa
and connexivum (fig. 83).

Macropterous female (fig. 80). — Coloration and
structure similar to apterous form. Pronotal width
subequal to median length, posteriorly slightly de-
pressed and transversely rugose. Pronotal margins ad-
jacent to prosternum narrowly yellow. Wings dark
reddish brown, membrane clearly surpassing the
proctiger. Veins not prominent reaching just beyond
end of connexivum. Subcostal, medial and anal veins
with minute black setae; basally with two closed cells
extending onto distal half.

Remarks. — Described from Papua New Guinea,
Astrolabe Bay, (Stephansort and Erima) and Raium
in New Britain. Type series in HNHM. Both sexes of
biroi are very similar, the hind femora are much alike.
The projecting distal connexival segments of the fe-
male are distinctive. The male parameres are long and
apically acuminate unlike other Rhagovelia so far re-
corded from ‘New Guinea’. The greatly reduced pro-
notal length compared with the longer mesonotum
distinguish biroi from species of the papuensis group
(priori, amnicus and browni etc.).

Rhagovelia hirsuta sp. n.
(figs. 84-93, tabs. 15-16)

Type material. — Holotype d: Papua New Guinea, Oro
Province, Kokoda, Mambare River, Mambe Estate,
12.ix.1988, sheltered pools at side of river. R. N. B. Prior
(OXUM). — Paratypes: Same data as holotype, 3d 59; East
Highlands Province, Isodenu Yonki, 1700 m. 13.x.1989,
Prior, 16 12 (OXUM).

Additional material. — Papua New Guinea, Oro Province,
Popondetta, Iseveni, small stream, 2.v.1989, Prior, 1d (OX-
UM); ‘Papua, Mafulu, 4000 feet, December 1933, L.E.
Cheesman’, 38 32 (BMNH).

Description

Adult apterous. — Male (fig. 84) 3.92-4.04 mm

59

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 84-93. Rhagovelia hirsuta sp. n. — 84-89 male, 90-93 female. 84, 90, dorsal habitus apterous; 85, front leg; 86, 91, hind
leg; 87, genital capsule; 88, mid-femora; 89, 92, antennae; 93, external genitalia. Figs. 84-86, 88-93 scale line 1 mm, fig. 87

scale line 0.5 mm.

long, maximum width 1.4-1.48 mm, female (fig. 90)
4-4.4 mm long, maximum width 1.52-1.64 mm.
Coloration. — Male (fig. 84): Dorsum generally
black. Tergites and most of connexivum black, upper
margins narrowly shining dark brown. Seventh ter-
gite with an elongate triangular shining area. Anterior
margin of pronotum orange brown. Lateral margins

40

of pronotum, prosternum, mesosternum and ster-
nites greyish blue tomentose. Proepisternum pale red-
dish brown, meso and metasternum dark grey mar-
gined reddish brown. Trochanters yellowish brown.
Basal 0.2 of front femur pale yellow, remainder of
front leg, middle and hind femora dark iridescent
green, remainder of legs black. Antennae black, basal

Table 15. Proportions of antennal segments of Rhagovelia
hirsuta sp. n.

I Il Ul IV

Apterous male 48 27 32 29
Apterous female 49 25 34 28.5

Table 16. Proportions of leg segments of Rhagovelia hirsuta
sp. n.

Femur Tibia Tarsus I Tarsus II Tarsus III

d frontleg 56 62 5 16 =
9 frontleg 57 62 5 17 -
3 middle leg 95 73 6 32 40
? middle leg 92 72 5 32 41
d hindleg 85 84 5 8 19
9 hindleg 75 88 4 7 19

0.2-0.3 of Ist pale yellow.

Structure. — Male: Maximum eye length clearly
longer than pronotum (18: 16). Pronotal width 3.4-
3.5x median length. Mesonotum 2.3x pronotal
length, metanotum 0.3x length of pronotum.
Tergites variable length, 1-8 are 12 : 12.5: 10.5: 10:
10 : 11.5 : 22 : 21. Eighth tergite densely pilose.
Pronotum laterally, meso and metaepimeron, lateral
margins of metanotum, tergites and lower margins of
connexivum with very long prominent semi-erect
hairs. Proepisternum, prosternum and lateral margins
of pronotum with fields of black spicules. Jugum al-
most touching the antenniferous tubercles. Second
sternite bluntly keeled, 7th slightly flattened, covered
with adpressed golden hairs. Eighth segment ventral-
ly concave and distally reflexed downwards. Legs, all
coxae with clusters of long golden hairs, hairs and
spines of all legs self coloured. Front leg (fig. 85). Mid
femora (fig. 88) slightly flattened and with fine hairs.
Hind leg (fig. 86) trochanter unarmed. Femus mod-
erately incrassate, antero-ventrad with a row of even-
sized setae, medianly and posteriorly with larger irreg-
ular sized spines. Tibia straight armed with small
setae along inner margin. Antennae (fig. 89) Ist seg-
ment not very spinose. Genital capsule (fig. 87) dis-
tally bluntly acuminate. Parameres large almost pad-
dle-like.

Coloration. — Female (fig. 90): Vertex, posterior
margin of pronotum, mesonotum, metanotum and
most of Ist tergite black. Anterior margin of pro-
notum pale orange band reaching inner margin of
eyes. Lateral margins of pronotum, visible tergites
and lateral margins of visible sternites evanescent
bluish grey. Connexivum apically narrowly dark
brown margined narrowly pale yellow. Prosternum
pale brown. Remainder of coloration similar to
male.

LANSBURY: Papuan Rhago velia

Structure. — Female: Eye length subequal to me-
dian pronotal length (17:18). Pronotal width 3.22x
median length. Mesonotum 2x median pronotal
length, metanotum about 0.5x pronotal length.
Nereitesyl-8rare 12.5: 12291071 3:17:19, 222014.
Eighth tergite and proctiger strongly reflexed down-
wards (fig. 93). Pronotum, lateral margins of meso
and metanotum and posterior margin of Ist tergite,
pro and mesosternum with scattered prominent black
hairs. Posterior margin of pronotum, all of meso-
metanotum and Ist tergite with adpressed short gold-
en hairs. Tergites 3-6 bluntly raised medianly with
discrete clusters of golden hairs increasing in length
and density on 6th tergite. Connexivum anteriorly re-
flexed outwards, from 3rd tergite converging and
meeting across 7th. Connexival margins of Gth with a
cluster of long hairs. Lateral margins of 8th segment
with two (1+1) clusters of curled hairs. Sternites cov-
ered with fine adpressed golden hairs, very dense on
7th sternite. Proctiger and ancilliary structures pilose.
Hind femur with a single prominent spine ventrad,
tibia straight, not armed along inner margin (fig. 91).

Antennae (fig. 92).

Etymology. — hirsuta refers to the hairy appearance
of this species.

Remarks. — This species tends to resemble R. aure-
ospicata sp. n. The female hirsuta differs in having the
connexivum converging posteriorly. The eye/prono-
tal ratio is subequal whereas the pronotum of aureos-
picata sp n. is longer than eye and the male middle fe-
mur has several stout setae proximo-ventrally whereas
male middle femur of hirsuta sp. n. is fringed with
fine hairs ventrally.

Rhagovelia aureospicata sp. n.

(figs. 94-104, tabs. 17-18)

Type material. — Holotype d : Papua New Guinea, Oro
Province, Kokoda, Mambare river, 24.ix.1989, R. N. B.
Prior (OXUM). — Paratypes, same data, 49 (OXUM).

Description

Adult apterous. — Male (fig. 94) 4.24 mm long,
maximum width 1.56 mm, female (fig. 95) 4.40-4.44
mm long, maximum width 1.60-1.72 mm.

Coloration. — Male (fig. 94): Dorsal ground colour
black. Anterior pronotal margin pale yellow.
Connexivum dark brown with a pale brownish yellow
band near upper margin. Prosternum pale yellow.
Mesosternum upper margins evanescent blue remain-
der pale yellowish brown. Metasternum evanescent
blue graduating into pale yellowish brown. Coxae
and trochanters yellowish brown. Inner proximal half
of front femur pale yellow, outer margin narrowly
pale yellow. Front tibia and middle legs iridescent

4]

TIDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

greenish brown. Hind femur mostly pale brown, dis-
tal and dorsal surfaces and hind tibia and tarsi dark
brown. Sternites 1-3 black, remainder yellowish
brown tending to merge into black along connexival

line. Antennae black, proximal third of Ist segment
pale yellow graduating into black.

Structure. — Male: Pronotum longer than eye
(25:18) pronotal width 2.44x median length.
Mesonotum 1.2x length of pronotum and 4.2x
length of metanotum. Lateral margins of prothorax
fringed with long hairs. Tergites 1-6 subequal, 1-8 are
PAPAS 2 12-3). 28): 50. Black dorsal ar-
eas covered with fine golden spicules other than 7th
tergite which is slightly flattened and dull shining,
posterior margin fringed with long hairs. Distal con-
nexival segment with very long black hairs which are
carried round posterior margin of 7th sternite.
Proepisternum without black spicules, pro-meso and
metasternum with scattered black semi-erect hairs.
Jugum broad almost touching antenniferous tuber-
cles. Seventh sternite shallowly depressed, covered
with fine brown hairs, ventrally segment depressed
adjacent to 7th sternite, segment sharply reflexed
downwards. Front leg (fig. 99) tibia slightly explanate
distally. Middle femur (fig. 100) proximo-ventrad
with four spines. Hind leg (fig. 98) femur moderately
incrassate, proximo-ventrad with a row of seven even
sized setae, medially with four large projections and a
variable number of smaller iregular setae. Tibia sinu-
ate, inner margin armed with stout projection distal-
ly and a row of small setae. Antennae (fig. 101) Ist
and 2nd segments moderately hairy. Genital capsule
(fig. 102), distal end of capsule curved upwards.
Parameres elongate, distally rounded.

Coloration. — Female (fig. 95): Head and thorax
similar to male. Tergites 5-7 with variable shining ar-
eas. Pale yellowish brown connexival band tending to
be slightly wider than of male. Metasternum black,
upper lateral margins of sternites black decreasing in
area to 7th sternite. Basal half of front femur pale yel-
low. Ventrally front femur, middle and hind femura
pale brown, dorsal surface of middle and hind femo-
ra iridescent green.

Structure. — Female: Pronotum longer than eye
(25:18) pronotal width 2.6x median length.
Mesonotum 1.2x length of pronotum and 6x length
of metanotum. Tergites variable in length, 1-8 are 13 :
SE1527145171725%: W522 2 WOsStiucturally the
thorax is same as male, the fringe of long hairs not
present. The density of gold spicules on pronotum is
not as dense as on mesonotum which has the spicules
arranged in a semi circular pattern. The spicules on

LANSBURY: Papuan Rhagovelia

Tabel 17. Proportions of antennal segments of Rhagovelia
aureospicata sp. n.

I Il III IV

Apterous male 50 34 42 34
Apterous female 50 32 38 3]

Tabel 18. Proportions of leg segments of Rhagovelia aureos-
picata sp. n.

Femur Tibia TarsusI Tarsus II Tarsus III

d front leg 63 64 5 119% =
9 front leg 60 Gl 5 17 -
3 middle leg 108 84 5 37 46
9 middle leg 93 15 5 35 42
d hindleg 105 055 10 DD
9 hind leg 79 95 4 9 20

the metanotum and tergites 1-2 are denser than on
male. Seventh tergite flattened and shining.
Connexivum anteriorly reflexed outwards becoming
more erect posteriorly. Proctiger and gonocoxal plates
strongly reflexed downwards (fig. 97). Hind leg (fig.
103) femur with a prominent spine distad ventrally,
tibia straight, inner margin unarmed. Antennae (fig.

104).

Etymology. — Aureospicata alludes to the golden
spicules on the dorsum.

Remark. — This species belongs to the group which
have the pronotum longer than the eye length, the
former being shorter than the mesonotum. The pro-
thorax is densely clothed with golden spicules and the
male mid-femur with proximo-ventrad group of setae
suggests a close relationship with herzogensis sp. n.

Rhagovelia herzogensis sp. n.

(figs. 105-116, tabs. 19-20)

Type material. — Holotype d apterous: New Guinea,
Morobe District, Herzog Mountains, Vagau c. 4000 feet, 4-
17.1.1965, M. E. Bacchus (BMNH). — Paratypes, same data,
36 39 apterous, 29 macropterous (BMNH, OXUM).

Description

Adult apterous. — Male (fig. 106) 4.08-4.2 mm
long, maximum width 1.4-1.48 mm, female (fig.
107) 4 mm long, maximum width 1.56 mm. Adult
macropterous. Female (fig. 105) 4.8 mm long, maxi-
mum width 1.72 mm.

Coloration. — Male (fig. 106): Head, thorax and

abdomen dorsally black. Pronotum anteriorly broad-

Figs. 94-104. Rhagovelia aureospicata sp. n. — 94, dorsal habitus apterous male; 95, ibid female; 96, external genitalia male;
97, ibid female; 98-102 male, 103-104 female. 98, 103, hind leg; 99, front leg; 100, mid-femora; 101, 104, antennae; 102,
genital capsule. Figs. 94-101, 103-104 scale line 1 mm, fig. 102 scale line 0.5 mm.

43

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

ly yellow. Inner lateral margins of connexivum with
yellow stripe. Seventh tergite slightly shining.
Anterior margin of mesonotum variably dark reddish
brown. Pro and mesopleura pale yellow, upper lateral
margins of metapleura black, remainder yellow.
Jugum and prosternum pale yellow, meso and meta-
sternum becoming progressively dark brown.
Sternites 2-4 dark reddish brown, 5-8 pale yellowish
brown. Lateral margins of sternites black, tapering
distally to 7. Coxae and trochanters pale yellow.
Front femur proximally yellowish brown, distally fe-
mur, tibia, middle and hind legs dorsally greenish
black shining. Underside of front femur and hind fe-
mur yellowish brown, remainder of legs dark brown
on ventral surfaces.

Structure. — Male: Median pronotal length longer
than eye length (20 : 18). Pronotal width 3x median
length. Mesonotum 1.6x median pronotal length and
6.4x length of metanotum. Tergites, 1-8 are 13 : 10:
Ile Wil 2 ils Ws D7 3 By, Whoyoer merrein Or 7dn were
gite slightly shining, rugose, 8th very pilose. Lateral
margins of pronotum, thorax, upper margins of con-
nexivum and tergites with scattered long black hairs.
Posterior margin of pronotum, meso and metanotum
and tergites 1-6 covered with silvery short hairs.
Connexivum reflexed slightly outwards with sparse
silvery pubescense. Distal connexival segment with
cluster of long curved hairs. Proepisternum without
black spicules, pro-meso and metasternum with scat-
tered brownish-black semi-erect spines. Jugum al-
most touching antenniferous tubercles. Second ster-
nite obtusely carinate, 3rd less so fringed with two
(1+1) rounded pilose elongated ridges. Eighth ster-
nite sharply depressed adjacent to posterior margin of
7th sternite. Front leg (fig. 109). Middle femur (fig.
110) proximo-ventrad with several stout setae, tiba
flattened. Hind leg (fig. 108) femur moderately in-
crassate, ventrally with two prominent projections
and several smaller setae, tibia slightly sinuate, inner
margin armed with setae. Antennae (fig. 111) seg-
ments 3-4 missing. Genital capsule (fig. 116) capsule
distally acuminate, parameres elongate. External gen-
italia and abdomen (fig. 112).

Coloration. — Female (fig. 107): Similar in general
appearance to male. Anterior yellow band on prono-
tum broader and tending to cover lateral margins pos-
teriorly. Reddish brown area of mesonotum more
prominent. Tergites 5-7 with shining areas. Front fe-
mur basal third pale yellow dorsally, ventrally pale
colour extending along femur. Middle and hind legs
dull greenish black ventrally pale yellowish brown.
Abdominal coloration similar to male.

Table 19. Proportions of antennal segment of Rhagovelia
herzogensis sp. n.

I II III IV

Apterous male 45 28 = =
Apterous female 45 30 39 31
Macropterous female 45 31 39 32.5

Table 20. Proportions of leg segments of Rhagovelia herzo-
gensis sp. n.

Femur Tibia TarsusI Tarsus II Tarsus III

Apterous

d front leg 61 67 4 16 =
9 front leg 60 60 5 16 >
3 middle leg 100 78 6 33 41
9 middle leg 94 68 5 32 42
d hind leg SOS 4.5 9 19.5
9 hind leg 74 88 3 7 20
Macropterous

9 front leg 55 60 4 15 =
9 middle leg 91 69 5 34 43
© hind leg 17 92 3.5 8 20

Structure. — Female: Median pronotal length long-
er than eye length (23:19.5). Pronotal width almost
3x median length. Mesonotum 1.17x median prono-
tal length and 6.75x length of metanotum. Tergites
Nes} eves W725) 2 WAS) MS ss 19224105, Tei
1-2, sometimes 3 with a slight median ridge, distal
shining tergites slightly flattened. Connexivum erect
anteriorly, converging posteriorly, either reflexed in-
wards slightly or curved over and almost touching
posteriorly obscuring shining area of 7th tergite.
Posterior half of pronotum with scattered silvery
hairs, mesonotum more densely pilose with hairs ar-
ranged in semi-circular pattern either side of midline.
Lateral margins of metanotum and tergites 1-4 with
decreasing areas of silvery hairs. Sternites medianly
clothed with fine semi-erect golden hairs. Hind leg
(fig. 113) femur slightly incrassate with a prominent
projection distally, tibia straight, inner margin un-
armed. Proctiger small and sharply reflexed down-
wards (fig. 114).

Macropterous female (fig. 105). — Similar to apter-
ous form, except pronotum greatly lengthened cau-
dad, anterior margin narrowly pale yellow, posterior
margin reddish brown. Wings with four closed cells,
wings brown, venation black. Wings extending well
beyond end of abdomen.

Figs. 105-115. Rhagovelia herzogensis sp. n. — 105, dorsal habitus female macropterous; 106, dorsal habitus male apterous;
107, ibid female; 108-112 male, 113-115 female. 108, 113, hind leg; 109, front leg; 110, mid-femora; 111, 115, antennae;
112, 114, external genitalia. Scale line 1 mm for figs 105-107, and 108-115 respectively.

44

LANSBURY: Papuan Rhagovelia

45

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 116. Rhagovelia herzo-
gensis sp. n. — Male genital
capsule. Scale line 0.5 mm.

Ss
SS

4,

S
NN

N
N
N

116

Etymology. — Specific name alludes to the type lo-
cality Herzog Mountains.

Remarks. — This species is closely allied to aureospic-
ata sp. n., male mid femura of both species bearing a
short row of small spines proximo-ventrally. Male gen-
ital capsules similar. Lateral pilosity is much denser of
aureospicata (fig. 94) compared with herzogensis (fig.
106). Female herzogensis has the connexivum converg-
ing posteriorly with clusters of long hairs (fig. 107),
connexivum of female aureospicata not convergent dis-
tally and lacking the clusters of long hairs (fig. 95).

Rhagovelia crinita sp. n.
(figs. 117-125, tabs. 21-22)

Type material. — Holotype d apterous: New Guinea,
Madang District, Finisterre Mts., c. 5500 feet, 30.x.-
15.xi.1964, M. E. Bacchus (BMNH). — Paratype: same data,
54 119 (BMNH, OXUM).

Additional material. — New Guinea: NE. Wau, Morobe
District, Mt. Missim, 1200 m. 19.iv.1966, G. Lippert, 16
apterous (BPBM).

Description

Adult apterous. — Male (fig. 117) 3.92-4.32 mm
long, maximum width 1.42-1.62 mm, female (fig.
118) 3.92-4.32 mm long, maximum width 1.42-1.66
mm.

Coloration. — Male (fig. 117): Head velvety black.
Pronotum anteriorly bright yellowish orange, pro-
pleura, proepisternum and jugum pale yellow.
Posterior half or more of pronotum, meso-metano-
tum, tergites 1-6 and inner lower margin of connexi-
vum greyish black. Tergites anteriorly narrowly black.
Dorsum covered with adpressed golden spicules
which tend to be denser on median areas of tergites.
Seventh tergite shining, 8th dark brown. Slightly ten-

46

NS

I

eral specimens tend to have anterior half of mesono-
tum reddish brown. Connexivum broadly yellowish
orange, apically narrowly black. Meso and metaepim-
eron dark reddish brown. Sternites 2-5 reddish
brown, black laterally, 6-8 tending to be pale yellow.
Proximal antennal segment narrowly pale yellow, re-
mainder of segments black. Coxae pale yellow. Inner
margin front femur and tibia pale yellow, distal outer
margins dark brown. Ventral margins of middle legs
pale brown, dorsally brown-black. Hind femur pale
yellow, dorsally dark brown, tibia and tarsi reddish
brown.

Structure. — Male: Median pronotal length sub-
equal to eye length (20:19). Pronotal width 3x me-
dian length. Mesonotum 1.6x pronotal length and
6.4x metanotum at midline. Tergites 1-8 are 13: 13:
13 : 12.5 : 14 : 16 : 25.5 : 20. Seventh tergite broad-
ly faintly shining appearing slightly transversely ras-
trate, 8th tergite densely pilose. Connexivum almost
erect, inner margins fringed with fine brown hairs,
distal lateral margins fringed with very long hairs.
Pronotum and propleura with scattered black hairs,
posterior margin of propleura with a few dark brown
pits. Jugum broad almost reaching antenniferous tu-
bercles. Thorax ventrally without fields of black spic-
ules. Meso and metapleura with scattered black hairs.
Second sternite and part of 3rd moderately carinate,
2-5 with scattered adpressed yellow hairs, 6th densely
hairy, 7th slightly flattened, laterally with two (1+1)
rows of long yellow hairs, 8th proximally acutely de-
pressed. Front leg (fig. 119) femur fringed with black
hairs, tibia proximally narrow, medially explanate
with a prominent fringe of stout setae/hairs decreas-
ing in length distally, inner margin distally acumi-
nate. Middle femur slightly flattened. Hind leg (fig.

120) femur moderately incrassate, ventral margin

LANSBURY: Papuan Rhagovelia

Figs. 117-125. Rhagovelia crinita sp. n. — 117, 119-122 male, 118, 123-125 female. 117-118, dorsal habitus apterous; 119,
front leg; 120, 123, hind leg; 121, 124, antennae; 122, genital capsule; 125, external genitalia. Figs. 117-121, 123-125 scale
line 1 mm, fig. 122 scale line 0.5 mm.

47

“TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

proximally with a row of regular sized setae, distally
with two prominent projections and several larger se-
tae. Tibia almost straight, inner margin armed with a
row of irregularly spaced setae. Antennae (fig. 121).
Genital capsule (fig. 122) capsule distally truncate,
parameres elongate, finely pilose.

Coloration. — Female (fig. 118): Broadly resembling
male. Mesonotum tending to be more widely anterior-
ly dark reddish brown merging to black laterally and
posteriorly. Tergites 3-8 becoming more yellowish
brown medianly, 5-7 medianly pale brown and shining
areas increasing in size distally. Some females have 3-8
becoming progressively more yellowish brown distally,
8 entirely yellowish brown. Connexivum and under-
side of thorax usually similar to male, metasternum
sometimes very dark reddish brown. Sternite 2-7 red-
dish brown, distally tending to be pale brown. Black
areas viewed from different angles appearing to bluish-
black. Antennae and legs similar to male.

Structure. — Female: Pronotum slightly longer than
eye (20:18). Pronotal width 3.7x median length.
Mesonotum 1.57x median pronotal length and 5.7x
metanotum at midline. Dark areas of pro-meso and
metanotum with scattered long black hairs. Tergites
128 are ASIN 2 1620102029 15. Terres
1-2 raised above following tergites. Connexivum an-
teriorly erect, reflexed over lateral margins of 3-7,
converging slightly posteriorly. Distally connexivum
appearing acuminate with a tuft of black hairs.
Sternites 1-6 evenly rounded. Distal margin of 7th
transversely depressed (fig. 125). Eighth segment re-
flexed downwards, proctiger small. Antennae (fig.
124). Front femur densely pilose, tibia laterally with
long hairs subequal to tibial width, distally with lon-
gitudinal depression. Middle femur slightly flattened.
Hind leg (fig. 123) femur not incrassate, distally with
a prominent projection, tibia straight, unarmed along
inner margin.

Etymology. — crinita alludes to the fringe of stout
hairs and spines along inner margin of male front tibia.

Remarks. — Rhagovelia crinita sp. n. and thysanotos
sp. n. are superficially rather similar. The male front
tibiae of both have a prominent fringe of stout hairs/-
spines along inner margin (figs. 119, 129). Distal
connexival segments of female thysanotos densely
fringed with long hairs, crinita females have smaller
tufts of hairs. The male genitalia of crinita differ from
thysanotos being smaller, parameres of crinita distally
bluntly acuminate (fig. 122) those of thysanotos are
bent over apically.

Rhagovelia thysanotos sp n.
(figs. 126-135, tabs. 23-24)

Type material. — Holotype d apterous: New Guinea,

48

Table 21. Proportions of antennal segments of Rhagovelia
crinita sp. n.

I II III IV
Apterous male 50 33 48 38
Apterous female 50 31 48 38

Table 22. Proportions of leg segments of Rhagovelia crinita
sp. n.

Femur Tibia Tarsus I Tarsus II Tarsus III

d front leg 70 13 17 =
9 front leg 67 68 6 17 =
d middle leg 110 83 5 41 43
9 middle leg 105 2 3 40 41
3 hind leg 100 UO 5 14 26
® hind leg 90 MOSS 13 24

Torricelli Mts., 1700 feet, 11.11.1939, G. P. Moore (BMNH).
— Paratypes: same data, 28 19 apterous, 1d macropterous
(BMNH, OXUM).

Description

Adult apterous. — Male (fig. 126) 3.84-3.92 mm
long, maximum width 1.4 mm. Female (fig. 132)
3.96 mm long, maximum width 1.52 mm. Adult
macropterous. Male 4.08 mm long, maximum width
1.48 mm.

Coloration. — Male (fig. 126): Head velvety black.
Pronotum anteriorly pale yellow reaching just beyond
inner margin of eyes. Lateral margins, propleura and
proepisternum bluish-black evanescent. Pronotum
posteriorly, meso, metanotum, tergites and most of
connexivum black. Upper margins of connexivum
narrowly dark brown, apex narrowly blackish-brown.
Seventh tergite faintly shining, 8th shining iridescent
green. Jugum and coxae pale yellow. Underside of
thorax and sternites greyish black, 6th sternite with
lateral margins shining brown. Antennae black, prox-
imal 0.2 of first segment pale yellow. Front leg, femo-
ra proximally pale yellow, remainder with tibia dark
iridescent green. Middle femur iridescent green, tibia
and tarsi rather darker. Hind femora dark brown with
a faint green iridescent patch on outer margin, tibia
and tarsi dark brown-black.

Structure. — Male: Eye length greater than median
pronotal length (18:16). Pronotal width 3.43x me-
dian length. Mesonotum 1.9x pronotal length and
4.3x length of metanotum at midline. Lateral bluish
margins of pronotum with scattered fields of minute
black pits. Posterior margin of propleura with an al-
most continuous row of similar pits. Tergites 1-8 are
MES 2 13:3 1.529) 75 OR 1322520) hornax and
tergites covered with short fine pale brown hairs.
Overlying these long black hairs which are particular-
ly dense along posterior margin of 7th tergite and

LANSBURY: Papuan Rhagovelia

i
\
N

LA

—=

Figs. 126-135. Rhagovelia thysanotos sp. n. — 126-131 male, 132-134 female. 126, 132, dorsal habitus apterous; 127, 134,
hind leg; 128, mid-femora; 129, front leg; 130, antenna; 131, 133, external genitalia; 135 male genital capsule. Figs. 126-134

scale line 1 mm, fig. 135 scale line 0.5 mm.

49

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

whole of 8th. Connexivum anteriorly slightly reflexed
outwards, posteriorly erect, inner and outer margins
pilose. Distal connexival segment fringed with very
long hairs. Eighth tergite distally with two (1+1) clus-
ters of shorter hairs. Pro and mesosternum with long
black hairs. Jugum broad almost touching antennifer-
ous tubercles. Second, 3rd and proximal part of 4th
sternites bluntly carinate and with a field of short fine
hairs. Fifth and 6th sternites laterally pilose, slightly
flattened, 7th more pilose, transversely depressed.
Front leg (fig. 129) femur moderately pilose. Tibia
proximally narrow, medianly explanate with promi-
nent fring of stout hairs decreasing in length distally,
inner margin distally acuminate. Middle femur (fig.
128) very slightly flattened. Hind leg (fig. 127) femo-
ra moderately incrassate, medianly almost parallel sid-
ed. Ventral margin anteriorly and posteriorly with al-
most regular rows of triangular setae. Anteriorly with
prominent projection, medianly with several smaller
projections. Hind tiba slightly sinuate, inner margin
armed with setae varying in size and density.
Antennae (fig. 130). Genital capsule ventrally pilose,
parameres elongate, distally slightly hooked (fig.
135). Entire genital segment very pilose (fig. 131).

Coloration. — Female (fig. 132): Broadly resembles
male, anterior margin of pronotum more brightly
coloured. Tergites appearing dull black with a faint
bluish evanescence, 7-8 with small shining bluish-
black discs, that of 8 covering most of segment.
Connexivum apically broadly dark brown. Thorax
ventrally and sternites bluish black. Legs and anten-
nae similar to male.

Structure. — Female: Eye length less than median
pronotal length (16:18). Median pronotal width 3.1x
median length. Mesonotum 1.3x median pronotal
length, posterior margin deeply emarginate, with a
row of black pits along margin. Metanotum 0.24x
length of mesonotum. Lateral margins of pronotum
with scattered long black hairs. Tergites 1-8 are 13 :
SENDER : 17.5 2 22,5% 15: Connie
slightly reflexed outwards anteriorly, converging at
4th tergite erect, posteriorly slightly sinuate converg-
ing and erect distally, with prominent clusters of long
dark brown-black hairs. External genitalia and procti-
ger sharply reflexed downwards, proctiger with lateral
clusters of hairs (fig. 133). Hind leg (fig. 134) femur
slightly incrassate, ventrally with two prominent pro-
jections and several smaller setae distally. Hind tibia
straight, unarmed along inner margin.

Macropterous male. — Elytra damaged one missing,
specimen in poor condition. Similar in coloration to
apterous male. Pronotum lengthened caudad, anteri-
or margin with transverse pale yellow band just reach-
ing inner margin of eyes. Wing reaching beyond end
of abdomen with four closed cells, basal two small,
extending into distal part of wing. Elytra yellowish

50

Table 23. Proportions of antennal segments of Rhagovelia

thysanotos sp. n. * = missing.

I II III IV
Apterous male 49 27 32 30
Apterous female 47 DD + =
Macropterous male 4] 21 si %

Table 24. Proportions of leg segments of Rhagovelia thysano-
tos sp. n.

Femur Tibia Tarsus I Tarsus II Tarsus III

Apterous

3 front leg 59 60 4

9 front leg 55 55 51 5 -

d middle leg 95 70 5 31 4]

9 middle leg 82 61 5) 29 40

3 hind leg 80 80 5 14 18
4

16.5 =

© hind leg 73 75 7 17
Macropterous

d front leg 52 52 4 13.5 =
3 middle leg 83 53 3 26.5 40
3 hind leg 75 65 4 7 17

brown, veins black, R and Sc with many dark brown
hairs. Hind leg similar to apterous male.

Etymology. — Thysanotos alludes to the fring of
stout hairs along the inner median margin of the male
front tibia.

Remarks. — Rhagovelia thysanotos sp. n. has features
in common with crinita sp. n. Males of both species
having a prominent fringe along inner margin of
front tibia. The anterior pale band of thysanotos does
not reach the lateral margins of pronotum and the
shining areas of the distal tergites are much reduced
compared with crinita. The male genital capsule of
thysanotos is relatively much larger than of crinita and
the parameres of the former are somewhat bent over
distally, almost spatulate. The condition of the type
series of thysanotos is very poor, only one male having
complete antennae. All specimens covered with detri-
tus/dust with signs of mite damage at some stage.

Rhagovelia caesius sp. n.

(figs. 136-150, tabs. 25-26)
Type material. — Holotype & apterous: Papua New

Guinea, Oro Province, Popondetta, Sambogo River, close to
stones, 30.iv.1989, R. N. B. Prior (OXUM). — Paratypes:
Same data, 648 289 apterous, 1d 19 macropterous (OX-
UM); Papua New Guinea, Milne Bay Province, streams near
Alotau, 27.iv.1989, Prior, 78 69 apterous (OXUM).
Additional material. — Papua: Kokoda, 1200 feet,
ix.1933, L. E. Cheesman, 7d 79 apterous; New Guinea,

LANSBURY: Papuan Rhagovelia

Figs. 136-150. Rhagovelia caesius sp. n. — 136, dorsal habitus macropterous female; 137, dorsal habitus apterous female; 138,
ibid male; 139-145 male, 146-150 female. 139, 146, front leg; 140, 147, distal end of tibia and tarsi of front leg; 141, 148,
hind leg; 142, 149, distal end of tibia and tarsi of hind leg; 145, 150, antennae. Figs. 136-138 scale line 1 mm, figs. 139, 141,
144-146, 148, 150 scale line 1 mm, figs. 140, 142-143, 147, 149 scale line 0.5 mm.

DI

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Table 25. Proportions of antennal segments of Rhagovelia
caesius sp. N.

I II III IV

Apterous male 38 19.5 17 19

Apterous female 48.5 30 23 23
Macropterous male 36 20 18 20

Table 26. Proportions of leg segments of Rhagovelia caesius
sp. n.

Femur Tibia Tarsus I Tarsus II Tarsus III

Apterous
3 frontleg 45 46 3
? front leg DE) 60 4.
3 middle leg 67 54 À 29 31
9 middle leg 85 67 5
3 hind leg 48 71 2
3

9 hind leg 60 88 9 19
Macropterous

d front leg 46 46 3 9 -
3 middle leg 68 23 28 34
3 hind leg 48 70 3 3 11

Morobe District, Lae-Bululo Road, 28.xii.1964, M.E.
Bacchus, 1d 29 apterous; Papua New Guinea, Morobe
Province, Buso, ix-xi.1979, J. Martin, 3d 32 apterous, 1d
12 macropterous (BMNH).

Description

Adult apterous. — Male (fig. 138) 2.32-2.52 mm
long, maximum width 1-1.08 mm. Female (fig. 137)
3.28-3.36 mm long, maximum width 1.56-1.64 mm.
Adult macropterous. Male 2.84-2.92 mm long, maxi-
mum width 1.24-1.28 mm. Female (fig. 136) 3.4
mm long, maximum width 1.64 mm.

Coloration. — Male (fig. 138): Head velvety black.
General dorsal coloration appearing greyish black.
Pronotum anteriorly with a rather indistinct orange
brown transverse band not reaching the inner margin
of eyes. Lateral margins of mesonotum sometimes
faintly orange brown. Pro, meso and meta pleura tu-
mescent black, viewed from various angles appearing
dark evanescent blue. Antenniferous tubercles shin-
ing black. Jugum and rostrum shining iridescent
greenish black. Median area of 7th sternite shining
pale brown. Coxae and trochanters shining yellowish
brown. Legs iridescent greenish black. Antennae
black, basal third of proximal segment pale yellow.

Structure. — Male: Eye length greater than median
pronotal length (13:9). Pronotal width 4.4x median
length. Mesonotum 3.6x median pronotal length, lat-
eral posterior margins emarginate. Metanotum about
0.12x length of mesonotum. Lateral margins of pro-
notum, all of mesonotum, metanotum, tergites and
connexivum with long semi-erect black hairs overly-

52.

ing short greyish hairs. Tergites 1-8 are 5.5 : 5.75 :
5.9:6:6:6:12.5:9. Connexivum usually flattened,
from side view tergites visible. Jugum clearly distinct
from antenniferous tubercles, proepisternum lacking
black spicules. Front coxae with clusters of short se-
tae. Front leg (figs. 139-140) femur moderately fine-
ly spinose, tibia curved explanate distally with a
prominent ridge of fine setae along inner distal mar-
gin (fig. 140). Middle and hind trochanters and cox-
ae with stout short yellowish hairs. Hind leg similar to
middle leg, not incrassate and spinose (figs. 141-142),
middle and hind femora slightly flattened, hind tarsi
pilose (fig. 142). Antennae (fig. 145) proximal seg-
ment with many long fine spines. Seventh abdominal
segment rounded, genital segment cylindrical and pi-
lose. Genital capsule (fig. 143) distally truncate, para-
meres elongate.

Coloration. — Female (fig. 137): Ground colour
appearing blackish suffused with faint greenish black
tumescence, pro-meso and metanotum sometimes
suffused pale yellowish brown. Pronotum anteriorly
with a faint transverse orange stripe merging into
greenish tumescence almost reaching inner margin of
eyes. Mesonotum and metanotum laterally pale
greenish black, mesonotum and metanotum with
field of minute black spicules. Connexivum and ter-
gites uniform greenish black. Antenniferous tuber-
cles, jugum and rostrum shining dark brown-black.
Pro-meso and metaepisternum silvery grey tumes-
cent. Venter greyish tumescent, 7th sternite shining
dull black. Coxae and trochanters variably pale-dark
yellow. Proximal margin of front femur narrowly pale
yellow, remainder of all legs greenish black iridescent.
Antennae black, proximal third of first segment pale
yellow.

Structure. — Female: Eye length greater than me-
dian pronotal length (17:13). Pronotal width 4.1x
median length. Mesonotum 3.3x pronotal length and
7.1x midline of metanotum. Posterior lateral margins
of mesonotum slightly emarginate, posterior margin
deeply emarginate. Tergites 1-8 are 6.5: 11 : 10 : 8.5:
9:10:11: 10. Connexivum usually flattened, some-
times elevated reflexed outwards. Dorsally thorax, ab-
domen and connexivum not pilose as in males.
Proepisternum without black spicules. Lateral mar-
gins of prothorax and abdomen with scattered erect
black hairs. Lateral margins of sternites 2-6 with
sparse golden short hairs. Seventh sternite black shin-
ing, laterally with fields of longer golden hairs increas-
ing in density along distal margin. Jugum distinct
from antenniferous tubercles. Front coxae and tro-
chanters with pale yellow spine-like hairs. Front leg
(fig. 146) femur and tibia curved with inner distal
margin of tibia slightly expanded (fig. 147). Middle
and hind legs similar, hind leg (fig. 148) tarsi elongate
and pilose (fig. 149). Antennae (fig. 15) proximal seg-

ment with many long fine hairs, 2nd segment sparse-
ly pilose.

Macropterous male. — Coloration not differing
greatly from apterous form. Pronotum long (1.08
mm). Pronotum at its widest raised very prominent-
ly. Forewings more or less uniformly brown extend-
ing well beyond the end of abdomen. Venation with
three closed cells proximally.

Macropterous female (fig. 136). — Coloration, re-
sembles apterous form. Pronotum about 1.52 mm
long, transverse orange band slightly more obvious
than in apterous form, posterior lobe transversely stri-
ated and finely pitted. Forewings basally greenish
grey, distally orange brown reaching just beyond end
of abdomen. Three closed cells in basal dark area of
wing.

Etymology. — Caesius alludes to the bluish grey ap-
pearance of the adult especially the female.

Remarks. — This species differs from others so far
known from New Guinea and Solomon Islands by
the absence of prominent spines/projections on the
hind femora of either sex. The sexual dimorphic body
size indicates phoretic behaviour. A pair from the
Sambogo River were taken ‘in cop’, the curved male
front femora fit in the lateral divergent margins of the
female mesonotum and the curved male tibia encircle
the pleural margins. The males also differ from the fe-
males in having the dorsum covered with long hairs,
whereas the females lack the long hairs dorsally and
their coloration is tomentose greyish blue. The fe-
males appear to be abnormally flattened compared to
the males, this aspect is exagerated by the flat connex-
ivum and broad prothorax in the majoriy of speci-
mens.

DISCUSSION

The Rhagovelia of the northern provinces of Papua
New Guinea have a distinctive endemic fauna. The
species so far recorded from the Oro Province
(Popondetta region - Kokoda) see map in Cheesman
(1943) includes priori sp. n.; hirsuta sp. n.; aureospi-
cata sp. n. and caesius sp. n. representing three species
groups, and does suggest that extensive sampling of
rain forest creeks and rivers in ‘limited areas’ will gre-
atly increase the Rhagovelia species known from
Papua New Guinea.

Rhagovelia caesius sp. n. is the most aberrant species
so far recorded from Papua New Guinea. The materi-
al included under this name has a distribution ex-
tending from Oro, Kokoda, 1200 feet to sealevel at
Alotau, Milne Bay Province. Further studies may
show that ‘caesius is a group of closely allied species.

Polhemus & Polhemus (1988) discuss at length the
zoogeography of Rhagovelia, commenting on the

LANSBURY: Papuan Rhagovelia

dearth of described species from the Philippines, New
Guinea, Solomons and other zoogeographically im-
portant areas. The Rhagovelia of New Guinea are
thought to have radiated extensively during the late
Tertiary and the existing Rhagovelia fauna is stated to
be endemic to New Guinea and to have originated
primarily since the Oligocene. Sources of New
Guinea Rhagovelia are thought to be New Caledonia
and relict forms from northern australian refugia.

Kikkawa et al. (1981) postulate that the continen-
tal margin of Australia extended to southern New
Guinea since the Lower Jurassic. Much of northern
Australia and New Guinea were subsea suring the ear-
ly Cretaceous. The northward movement of Australia
continued and the collision of crustal plates in the
mid Miocene created the present land mass of New
Guinea. By the Pliocene, most of New Guinea had
emerged from the sea. The major uplift of the present
mountain ranges with major faulting accompanied by
widespread volcanic activity in the late Miocene - ear-
ly Pliocene (7 Ma) and continues today. Recent stud-
ies of the north coast of New Guinea show 32 ter-
ranes (micro-tectonic plates), each of which has
‘docked’ to the north coast over the past 20 million
years, as the island was pushed northwards through a
complex subduction zone. An earthquake on the 15th
May, 1992 caused a 10 cm uplift near Sialum
(Morobe Province), which was felt very strongly
along the Madang coastline.

The total uplift may exceed 3000-4000 m. Smith
& Briden (1977) commenting on the tectonic folding
and uplift in central New Guinea and ‘fragmented
plates’ suggest the fragmented plates created the
‘Indonesian islands’ and may have joined the conti-
nents of Asia and Australia allowing the first wave of
Asian migrants to enter the Australian region.

The evidence of palaeogeography, climate and
geology present a very confusing set of scenarios to ac-
count for the diversity of some groups of water-bugs
in New Guinea. During the Pleistocene sea-level
changes, New Guinea was linked to Australia until
between 6500-8000 B. P. The aridity caused by the
lowered sea-levels may have inhibited the movement
of Rhagovelia between Australia and New Guinea and
species on either land mass may have survived in re-
fugic mesic habitats. The record of R. australica
Kirkaldy from Mt. Lewis is of more than passing
interest, as a number of relict forms have been record-
ed from Mt. Lewis. Other examples include the leaf-
hopper Myerslopia taylori Evans, a primitive tribe
hitherto known from Madagascar, New Zealand,
Chile and Juan Fernandez, further /gnambia, a flight-
less genus of dung beetles (Coleoptera) also known
from New Caledonia, and the stag beetle Sphaeno-
gnathus sp. (Coleoptera) unknown outside South
America.

53

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

The formation of the mountain ranges with high
rainfall has formed a wide variety of isolated lotic hab-
itats, which over a long period may have enhanced the
biodiversity of pleuston bugs in New Guinea.

ACKNOWLEDGEMENTS

The generous assistance of Mr. R. N. B. Prior, late-
ly residing in West New Britain for the extensive col-
lections he made on my behalf, especially from the
Oro Province. His help during a short visit to Kimbe,
W. N. B. was of great help to me. The British
Museum (Natural History), London (BMNH) and the
Bernice P. Bishop Museum, Honolulu (BPBM) kindly
loaned material for this review.

REFERENCES

Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera,
Gerromorpha). Phylogeny, adaptations, biogeography
and classification. — Entomonograph 3: 1-455.

Cheesman, L. E., 1943. Japanese Operations in New
Guinea. — The Geographical Journal 101 (3); 98-110.

Hale, H. M., 1925. Results of Dr. E. Mjoberg’s Swedish
Scientific Expedition to Australia 1910-1913. The
Aquatic and Semi-Aquatic Hemiptera. — Arkiv för
Zoologi 17A (20): 1-19.

Hale, H. M., 1926. Studies in Australian Aquatic

54

Hemiptera No. VII. — Records of the South Australian
Museum 3: 195-217.

Kikkawa, J., Monteith, G. B. & G. Ingrams, 1981. Cape
York Peninsula: Major region of faunal interchange. p.
1697-1736. —In: A. Keast (Ed.), Ecological biogeography
of Australia. Junk, The Hague.

Kirkaldy, G. W., 1901. On some Rhynchota, principally
from New Guinea, (Amphibicorisae and Notonectidae).
— Annali del Museo Civico di Storia Naturale G. Doria
20: 804-810.

Kirkaldy, G. W., 1908. Memoir on a few heteropterous
Hemiptera from eastern Australia. — Proceedings of the
Linnean Society of New South Wales 32: 768-788.

Lundblad, O., 1933. Zur Kenntnis der aquatilien und sem-
iaquatilen Hemipteren von Sumatra, Java und Bali. —
Arkiv fiir Hydrobiologie, Supplement 12: 1-195, 263-
489, pls. 1-21, figs. 142.

Lundblad, ©., 1936. Die altweltichen Arten der
Veliidengattungen Rhagovelia und Tetraripis. — Arkiv för
Zoologi 28A: 1-63.

Polhemus, J. T. & D. A. Polhemus, 1988. Zoogeography,
ecology, and systematics of the genus Rhagovelia Mayr
(Heteroptera: Veliidae) in Borneo, Celebes, and the
Moluccas. — Insecta Mundi 2: 161-230.

Smith, A. G. & J. C. Briden, 1977. Mesozoic and Cenozoic
Palaeo-continental maps. — Cambridge University Press;

Cambridge: 63 pp.

Received: 21 October 1992
Revised version accepted: 6 April 1993

WAYNE N. MATHIS' & TADEUSZ ZATWARNICKT

' Smithsonian Institution, Washington, D. C., USA
? Academy of Agriculture, Wroclaw, Poland

REVISION OF THE SHORE-FLY GENUS
CHLORICHAETA BECKER (DIPTERA: EPHYDRIDAE)

Mathis, W. N. & T. Zatwarnicki, 1993. Revision of the shore-fly genus Chlorichaeta Becker
(Diptera: Ephydridae). — Tijdschrift voor Entomologie 136: 55-76, figs. 1-53. [issN 0040-
7496]. Published 1 July 1993.

Chlorichaeta Becker, an Old-World genus of shore flies of the tribe Gymnomyzini, is revised.
Included are seven species of which C. africana (Kenya: Lake Victoria), C. mais (Seychelles:
Aldabra), and C. orba (Sri Lanka: Nilaveli) are newly described and C. gioiellae Canzoneri &
Rampini is determined to be conspecific and the junior synonym of C. villiersi Canzoneri &
Meneghini.

Correspondence: Wayne N. Mathis, Department of Entomology, NHB 169, Smithsonian
Institution, Washington, D.C. 20560, USA.

Key words. — Diptera; Ephydridae; Gymnomyzinae; Chlorichaeta; revision; new species; Old

World.

Shore flies of the genus Chlorichaeta Becker are
known only from the Old World, mostly where
warmer climates prevail, such as around the
Mediterranean Sea or in tropical regions of Africa and
the Orient. Only one species, C. albipennis (Loew), is
known to occur in the more temperate regions of the
Palearctic Region (Cogan 1984). The genus has nev-
er been treated on a worldwide basis, and that defi-
ciency, in part, prompted the research that resulted in
this revision.

There are few species in Chlorichaeta, and these are
poorly known except to specialists. The history of
previous work on the genus, as a consequence, is rela-
tively brief and straightforward. Becker (1922) pro-
posed Chlorichaeta towards the end of his career as a
dipterist, and in the same paper, he described C. tu-
berculosa, which became the type species of the genus
by monotypy. Becker’s first and finest published
work on shore flies was his monograph on Palearctic
species (1896), which included Gymnopa albipennis, a
species Loew (1848) described in his first paper on
shore flies. This species remained in Gymnopa or its
senior synonym, Mosillus Latreille, until Cresson
(1925) transferred it to Chlorichaeta. The genus com-
prised only these two species until the 1940’s when
Duda (1942) described Strandilus strandi in a pecu-
liar paper in which each new genus- or species-group
name, which numbered five and 19 respectively, was
named after Embrik Strand (1876-1947) except
where a homonym would result. Papp (1979) studied
Duda’s collection, now mostly housed in Berlin
(ZMHU), and found that the primary types of S. stran-
di (a lectotype was then designated) are conspecific

with Gymnopa albipennis with the latter being the
senior synonym. Cresson (1946), as part of a synopsis
of Afrotropical shore flies, described C. aerifer from
specimens collected in Angola. After another gap of
two decades, Canzoneri & Meneghini (1969a) de-
scribed C. villiersi from specimens collected in
Senegal and published the first figures of male termi-
nalia. In another publication, a well-illustrated faunal
treatment of shore flies occurring in Italy, Canzoneri
& Meneghini (1983) also provided a synopsis of C.
albipennis. Finally, Canzoneri & Rampini (1990) de-
scribed C. gioiellae from specimens collected in Sierra
Leone. Nothing is known about the immature stages,
and what little we know about the natural history
comes from brief annotations that were included as
part of taxonomic reviews or listings (Cresson 1946).
Apart from entries in regional catalogues (Cogan
1980, 1984, Mathis 1989) little other work has been
published on the genus.

In addition to revising the species of Chlorichaeta,
we recharacterize the genus within the context of phy-
logenetic studies we are conducting on the tribe
Gymnomyzini Latreille. Gymnomyzini include sev-
eral genus-group taxa (Mathis et al. 1993), of which
we have revised Placopsidella Kertész (Mathis 1986),
the western Palaearctic species of Athyroglossa Becker
(Mathis & Zatwarnicki 1990), and Mosillus Latreille
(Mathis et al. 1993). Other species revisions of gym-

nomyzine genera are anticipated.

METHODS

The methods used generally in this study were ex-

55

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

plained previously (Mathis 1986, Mathis & Zatwar-
nicki 1990). Because specimens are small, less than
3.50 mm, study and illustration of the male termina-
lia required use of a compound microscope.

Although we have followed the terminology for
most structures of the male terminalia that other
workers in Ephydridae have used (see references in
Mathis 1986, and Mathis & Zatwarnicki 1990a,
1990b), Zatwarnicki (1992) advocated usage of the
term dististylus rather than surstylus, largely based on
an hypothesis advanced by Hennig (1936). The mer-
its of Hennig’s hypothesis are still being debated, and
here we continue to use surstylus. In papers for which
Zatwarnicki is first author, however, dististylus will
be used.

Three ratios (two venational) are used commonly
in the descriptions and are defined here for the con-
venience of the user (ratios are averages of three spec-
imens):

1. Eye-to-cheek ratio: Genal height / eye height.
Measurements are taken from the head in lateral
view.

2. Costal vein ratio: the straight line distance
between the apices of R,,, and R,,,/ distance between
the apices of R, and R,,,.

3. M vein ratio: the straight line distance along M
between crossveins (r-m and dm-cu) / distance apicad
of crossvein dm-cu.

Although this study was based in large part on
specimens in the U. S. National Museum of Natural
History, numerous others were borrowed, particular-
ly type specimens of the species previously described.
To our colleagues and their institutions listed below
who lent specimens, we express our sincere thanks.
Without their cooperation this study could not have
been completed.

ANSP — Academy of Natural Sciences of
Philadelphia, Pennsylvania (J. K. Gelhaus and
Donald Azuma); BMNH — British Museum (Natural
History), London, England (Brian Pitkin); BBM —
Bernice P. Bishop Museum, Honolulu, Hawaii (N.
L. Evenhuis); Cas — California Academy of Sciences,
San Francisco, California (P. H. Arnaud, Jr.); CANZ —
Personal collection of Silvano Canzoneri, Venice,
Italy; MNHN — Muséum National d'Histoire
Naturelle, Paris, France (Loïc Matile); NMW —
Naturhistorisches Museum, Vienna, Austria (Ruth
Contreras-Lichtenberg); NMWL — National Museum
of Wales, Cardiff, Wales, United Kingdom (J. C.
Deeming); TAU — Tel Aviv University, Tel Aviv, Israel
(A. Freidberg); USNM — former United States
National Museum, collections in the National
Museum of Natural History, Smithsonian Institu-
tion, Washington, D. C.; ZMHU — Zoologisches
Museum, Humboldt Universität, Berlin, Germany

(H. Schumann).

56

SYSTEMATIC ACCOUNT

Genus Chlorichaeta Becker

Chlorichaeta Becker, 1922: 73. — Type species: Chlorichaeta
tuberculosa Becker, 1922, monotypy; Cogan and Wirth
1977: 322 [Oriental catalogue], Cogan 1980: 656
[Afrotropical catalogue]; 1984: 127-128 [Palearctic cata-
logue], Canzoneri & Meneghini 1983: 227-229 [diagno-
sis].

Strandillus Duda, 1942: 10. — Type species: Strandillus
strandi Duda, 1942 (= Gymnopa albipennis Loew, 1848),
monotypy; Papp 1979: 98 [synonymy].

Diagnosis

Small to medium-sized shore flies, length 1.50-
3.10 mm, mostly black, many surfaces subshiny to
shiny; dorsum, especially of thorax, somewhat micro-
tomentose, appearing subshiny to dull.

Head: Setation generally poorly developed; fronto-
orbital setae greatly reduced or lacking; ocellar setae
short, inserted laterad of anterior ocellus and widely
separated; both inner and outer vertical setae present,
short. Frons generally smooth, shiny, ocellar triangle
of mesofrons sometimes differentiated. Arista com-
paratively short, subequal to length of combined first
3 antennal segments, appearing bare or bearing sever-
al, very short hairs along dorsal surface, length of hairs
subequal with basal aristal width. Face symmetrically
pitted around midfacial protuberance, pits with sil-
very white microtomentum, making face appear spot-
ted; midfacial conical protuberance not prominent;
parafacials immediately adjacent to face with a verti-
cal row of shallow, horizontal grooves; portion of par-
afacial near anteroventral margin of eye with some sil-
very white microtomentum; gena generally bare and
high, height 2/3 to 3/4 eye height; posterior margin
sharply angulate and marginate.

Thorax: Mesonotum mostly black, shiny, fre-
quently with metallic luster, especially scutellum;
mesonotal surface appearing granulose to various de-
grees. Notopleuron with | seta inserted near postero-
ventral angle; apical scutellar setae arising from tu-
bercles; intra-alar seta 1; lacking row of prominent
setae between postalar seta and base of scutellum;
coloration of halter variable. Wing milky white; vein
R,,, long, with length of costal section II not quite
twice section III; vein CuA, not reaching wing mar-
gin; alula wide alular marginal setulae much shorter
than alular width. Fore femur much more swollen
than mid or hind femora, produced ventrally to a
somewhat pointed ridge near midlength, process
bearing a row of 4-5 stout setae along anterior half of
posteroventral margin, seta arising from ventral,
midlength point conspicuously larger; mid tibia
black, somewhat rounded (lacking conspicuously
flattened surfaces), lacking silvery-white microto-

mentum on anterior surface. Basitarsomere colour
variable, apical 2-3 tarsomeres usually blackish brown.

Abdomen: Dorsal surface of terga black, shiny,
more finely granulose than mesonotum, lacking
bands of microtomentum; terga well sclerotized, lat-
eral margins continued laterally and ventrally; sterna
of male relatively poorly developed, usually as small
sclerotized rectangular plates, Ist sternum of male
oriented perpendicular to plane of body, sterna 2-4
parallel to plane of body, 5th sternum as 2 sternites,
these oriented to form a V, with vertex anteriad,
sometimes fused at vertex; 5th tergum exposed but
shorter than 4th, usually triangular or trapezoidal,
with 2 dorsal pits toward posterior margin. Male ter-
minalia: Epandrium in lateral view narrowed dorsal-
ly, expanded laterally, in posterior view widest at lev-
el of cerci; cerci oval, bearing short setae; hypandrium
more or less U or V-shaped with posterolateral por-
tion relatively long (representing fused distal portion
of pregonite to hypandrium) and anteromedial por-
tion forming vertex variously developed; surstyli gen-
erally elongate, subequal in length to that of cerci,
broadly fused basally with ventral margin of epandri-
um, becoming tapered ventrally, pointed apically,
ventral portion generally curved anteriorly to some
degree; aedeagal apodeme more broadly developed in
lateral view toward end that attaches to hypandrium;
pregonite apparently fused with hypandrium; post-
gonite linear, rod-like, setulose; aedeagus with dis-
tinct basiphallus, distiphallus, and a sclerotized rod
(ejaculatory apodeme) attached by a membranous
duct to the base of the basiphallus; basiphallus well
sclerotized, simple, tube-like; distiphallus more com-
plex, with 2 sclerotized rods basally, apex membra-
nous, in some species bearing 1-2 paired extended lat-
eral processes and 1 unpaired ventral process, these
sometimes sclerotized, all processes arising near base

of distiphallus.

Distribution

Old World in tropical and warm temperate zones.
Afrotropical, Australasian, Oceanian, Oriental and
Palearctic (Japan, Lithuania, and Mediterranean) re-
gions.

Natural history

Species of Chlorichaeta apparently feed as general
scavengers in concentrated organic material, such as
manure of terrestrial vertebrates. Cresson (1945: 53)
reported that C. tuberculosa bred in rotten oil cake
manure in Madras, India, and that the dissemination
of the manure may account for the wide distribution
of this species. On atolls of the Aldabra Group, C.

mais, a new species described below, occurred in

MarHIs & ZATWARNICKI: Revision of Chlorichaeta

several habitats. We collected it most commonly on
tortoise droppings but also found it around pig pens
near the settlement (Picard Island) and occasionally
on rotting seaweed that had accumulated at the
strand line on beaches.

Adults are attracted to moisture on animals and are
frequently associated with livestock (camels, cattle,
donkeys, pigs). Suppurating sores on livestock are es-
pecially attractive (Cresson 1946: 244), and in the
Negev Desert of Israel, Amnon Freidberg and W. N.
Mathis observed that adults of C. tuberculosa were at-
tracted to moisture, especially perspiration on bare
legs. Cresson (1946) also noted that a specimen was
found on a coccid (Homoptera) on the trunk of a
China-tree in Angola (this occurrence may be inci-
dental and of little significance).

Diagnosis and discussion of phylogenetic
relationships

Chlorichaeta is similar and closely related to
Mosillus Latreille and a group of species related to
‘Gymnopa beckeri Cresson’. The genus is more dis-
tantly related to Placopsidella and possibly to
Chaetomosillus. The monophyly of Chlorichaeta,
Mosillus, and the species related to ; Gymnopa becker?
(hereafter called the Mosillus clade) is established by
the following synapomorphies:

1. The hypandrium, especially the lateral arms, is
fused with the pregonites, a character unique to this
clade.

2. The fore femur bears 5-8 stout setae, with the
first one or two being the largest. In most genera of
Gymnomyzini the fore femur is either unarmed or in
a few genera the fore femur is produced posteroven-
trally, usually forming a pointed, narrow ridge
(Chaetomosillus and Stratiothyrea) or there is 1-2 stout
setae (Athyroglossa).

3. The anterior margin of the hypandrium is pro-
duced anteriorly to form a Y-shaped sternite with the
two lateral arms. In other genera the hypandrium is
truncate to broadly rounded, but not produced ante-
riorly.

4. The ocellar setae are reduced or lacking. In oth-
er genera of Gymnomyzini the ocellar setae are dis-
tinct even though varying in size from small to well
developed.

Chlorichaeta is distinguished from related genera of
the tribe Gymnomyzini by the synapomorphies not-
ed for the clade and the following characters: body
mostly shiny black with reduced setation; face pitted,
appearing spotted, and with silvery white microto-
mentum in pits; mid facial conical protuberance
small; parafacials with a vertical row of furrows; apical
scutellar setae usually arising from basal tubercles;
wing lacteous; alula of wing broad, auriculate along

5%

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-5. Male genitalia of Chlorichaeta aerifer. — 1, epandrium and cerci, posterior view; 2, idem, lateral view; 3, internal
male genitalia, lateral view; 4, idem, ventral view; 5, hypandrium and postgonites, ventral view.

58

inferior margin, strongly convex; fore femur bearing a
large ventral seta at ventral prominence near midlength
and with 5-6 short but stout setae between largest seta
and apex along posteroventral margin; fifth tergum of
male and to a lesser degree in female with 2 small, dor-
sal depressions towards posterior margin.

Key to species groups and species of Chlorichaeta

1. Scutellum with brilliant, golden-bronze luster,
distinctly contrasted with blackish scutum; dis-
tiphallus short, angulate, lacking long, paired
processes at base (the 4erifer group) .................. y)

— Scutellum and scutum either essentially concol-
orous, mostly blackish, or scutum slightly golden
bronze and scutellum blackish; distiphallus elon-
gate, mostly straight or gently curved, bearing 2,
paired, basal processes (the albipennis group) ...3

2. Parafacials with 3rd silvery-white area narrowly
linear (sometimes narrowly connected dorsally
with 2nd spot) at anteroventral margin of eye;
posteroventral process of postgonite elongate,
HTT combo ey (COS
TE IONI. C. villiersi Canzoneri & Meneghini

— Parafacials with 3rd silvery-white area elongate,
lunate or shallowly triangular along anteroventral
margin of eye; posteroventral process of postgon-
ite relatively short (fig. 3) … … C. aerifer Cresson

3. Hind basitarsomere yellow, sometimes slightly
darker basally, mostly concolorous with tarso-

mers Dana ee 4
— Hind basitarsomere almost entirely dark colour-
edublackishi brown to brown... 5

4. Extended ocellar triangle granulose, contrasted
with shiny and smooth parafrons; silvery-white
microtomentum on parafacials as 3 patches: 2
spots near level of antenna (dorsal spot at level of
ventral margin of pedicel, ventral spot at level of
midheight of Ist flagellomere) and a more linear
patch at anteroventral margin of eye; mesonotum
distinctly microsculptured, appearing granulose,
with some metallic bronzish-gold luster .............

RO MI Ra C. mais sp. n.

— Extended ocellar triangle shiny and smooth, sim-
ilar to parafrons; silvery-white microtomentum
on parafacials as 2 patches: 1 spot at level of ped-
icel and a long linear patch at anteroventral mar-
gin of eye; scutum very lightly microsculptured,
scutellum more so, appearing granulose; mesono-
tum with some metallic bluish luster ..................

RR MR nn C. orba sp. n.

5. Scutellum bearing 3-5 large, marginal setae on
each side that arise from distinct tubercles; disc of
scutellum distinctly more microsculptured than
Seutumappearinetsranulose Renn

Matuis & ZATWARNICKI: Revision of Chlorichaeta

— Scutellum bearing 2 large, marginal setae on each
side that arise from tubercles (1-2 smaller setae
usually between larger setae); scutum and scutel-
lum about equally microsculptured . 6

6. Surstyli in lateral view distinctly and more or less
evenly curved anteroventrally, tapered gradually
and evenly to acutely pointed apex......................
BO o Sek SN GM se dors de C. tuberculosa Becker

— Surstyli in lateral view more shallowly curved an-
teroventrally and with apex slightly recurved and
somewhat spatulate … … C. africana sp. n.

THE aerifer GROUP

Diagnosis

This species group is distinguished from other taxa
of Chlorichaeta by the following combination of char-
acters (synapomorphies, which provide evidence of
the group’s monophyly, are indicated by an asterisk):
mesonotum appearing granulose, microsculptured,
shiny; *scutellum except along base with brilliant,
golden-bronze luster, distinctly contrasted with blac-
kish scutum, disc of scutellum distinctly rugose, with
rugae aligned longitudinally; larger 4-5 setae along
each side scutellum arising from tubercles; *distiphal-
lus short and lacking long, paired processes at base.

Both species of this group occur in the Afrotropical
Region, principally in countries along the west
coast.

Chlorichaeta aerifer Cresson
(figs. 1-6)

Chlerichaeta aerifer Cresson, 1946: 245. Holotype 9,
Angola: Makussi, July 23, 1931, J. Ogilvie (Cresson stat-
ed that the holotype is deposited in the BMNH , but on vis-
its there, the holotype was not found, nor did we find it
at the ANSP); Cogan 1980: 656 [Afrotropical catalogue].

Diagnosis

This species and C. villiersi are sister species, as es-
tablished by the synapomorphies noted for the species
group (see characters noted above and in key). It is
distinguished from C. villiersi by the short projection
of the postgonite and the more flared apex of the sur-
stylus. In specimens of C. villiersi the postgonal pro-
jection is long, finger-like, and the apices of the sur-

styli are parallel.

Description

Small to medium-sized shore flies, length 1.85 to
3.10 mm.

Head: Mesofrons, especially extended ocellar trian-
gle, entirely smooth, shiny, similar to parafrons; para-

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 6. Distribution map for
Chlorichaeta aerifer (dots)
and C. villiersi (diamonds).

20

facials with silvery-white to white microtomentum
arranged as 3 spots, Ist at level of pedicel, 2nd at lev-
el of first flagellomere, 3rd along anteroventral mar-
gin of eye, distinctly wider at middle, lunate or
sometime shallowly triangular. Gena very high, eye-
to-cheek ratio averaging 0.75.

Thorax: Mesonotum finely microsculptured, ap-
pearing granulose, scutellum more so; scutum black
with faint metallic bluish reflections; scutellum with
base 1/8 black, remainder brownish gold with metal-
lic reflections; marginal scutellar setae 4-6 on each
side that arise from tubercles. Halter white. Costal
vein ratio averaging 0.50; vein M ratio averaging
0.48. Fore basitarsomere reddish orange, other fore
tarsomeres becoming darker, apical 2 black; hind bas-
itarsomere black.

Abdomen: Male terminalia (figs. 1-5): Length of
epandrium+surstyli in posterior view 1.3 X width;
lateral margins of surstyli in posterior view distinctly
concave ventrally (fig. 1); surstylus in lateral view di-
gitiform, moderately rounded apically, distinctly
curved anteroventrally at basal merger with ventral
margin of epandrium (fig. 2); hypandrium in lateral
view sinuous basally, in posterior view broadly V-
shaped, emarginate ventrally; aedeagal apodeme only

60

slightly enlarged toward end that attaches to hypan-
drium (fig. 3); postgonite with a posteroventral, short
process (fig. 3), in posterior view becoming broader
toward apex, and apex concave; distiphallus distinctly
angulate, dorsal angle acutely produced, pointed;
bearing 1 short, lateral process that is broadly at-
tached toward base (figs. 3, 4), basoventral margin of
process irregularly serrate (figs. 3, 4).

Other specimens examined

Afrotropical: Angola. Luanda, 22 Aug 1949, Malkin
(18, 39; USNM). Cameroon. Kribi (beach), 28-29 Nov
1987, F. Kaplan (4d, 49; USNM); Kribi, Rt. N7 (beach),
28-29 Nov 1987, A. Freidberg (ld, 29; USNM). Nigeria.
Lagos (shore), 15 Dec 1987, A. Freidberg (26; USNM);
Lagos, Victoria Island (shore), 15 Dec 1987, F. Kaplan (36,
29; USNM). Sa Tomé and Principe. Sao Tomé (sandy be-
ach), G.R. Gradwell, D. Snow (66, 122; BMNH ).

Distribution (fig. 6)

Afrotropical. Angola (Luanda, Makussi), Came-
roon, Nigeria, and Sao Tomé and Principe.

MATHIS & ZATWARNICKI: Revision of Chlorichaeta

Chlorichaeta villiersi Canzoneri & Meneghini MUSEUM PARIS 2-X-61 [2 Oct 1961] A. Villiers [date and
(figs. 6-11) collector handwritten, blue]/sur Alternanthera [hand-
written]/HOLOTYPUS d Chlorichaeta villiersi sp. n. [gen-

Chlorichaeta villiersi Canzoneri & Meneghini, 1969a: 1481. der, name, and ‘sp. n.’ handwritten, red}. The holotype
Holotype d: ‘Sénégal Kayar [Kayar handwritten]/- is glued to a card, is in very poor condition (specimen

Figs. 7-11. Male genitalia of Chlorichaeta villiersi. — 7, epandrium and cerci, posterior view; 8, idem, lateral view; 9, internal
male genitalia, lateral view; 10, idem, ventral view; 11, hypandrium and postgonites, ventral view.

61

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

glued upside down, obscuring dorsum of head and tho-
rax; left wing, forelegs, and abdomen missing), and is de-
posited in the MNHN; Cogan 1980: 656 [Afrotropical cat-
alogue], Canzoneri 1981: 203 [list, Senegal]; 1982: 57
[list, Sierra Leone].

Chlorichaeta gioiellae Canzoneri & Rampini, 1990: 112. —
Holotype d : ‘Sierra Leone[,] W.Area-Spiaggia[,] Sussex -
9.XI.87 [9 Nov 1987] Leg. Rossi/ HOLOTYPUS d
Chlorichaeta gioiellae nov. det. Canzoneri S. [gender, spe-
cies name, and ‘nov.’ handwritten, red]’. The holotype is
double mounted (minuten in a rectangular block of plas-
tic foam), is in fair condition (left mid leg missing), and is
deposited in CANZ. Syn. n.

Diagnosis

This species and C. aerifer are closely related and
very similar but the former is distinguished by the
longer, finger-like projection of the postgonite.

Description

Moderately small to medium-sized shore flies,
length 2.10 to 3.00 mm.

Head: Mesofrons, especially extended ocellar trian-
gle, entirely smooth, shiny, similar to parafrons; para-
facials with silvery-white to white microtomentum
arranged as 3 spots, Ist at level of pedicel, 2nd at lev-
el of first flagellomere, 3rd along anteroventral mar-
gin of eye, distinctly wider at middle, lunate or some-
time shallowly triangular. Gena very high,
eye-to-cheek ratio averaging 0.65.

Thorax: Mesonotum finely microsculptured, ap-
pearing granulose, scutellum more so; scutum black
with faint metallic bluish reflections; scutellum with
base 1/8 black, remainder brownish gold with metal-
lic reflections; marginal scutellar setae 4-6 on each
side that arise from tubercles. Halter white. Costal
vein ratio averaging 0.49; vein M ratio averaging
0.47. Fore basitarsomere reddish orange, other fore
tarsomeres becoming darker, apical 2 black; hind bas-
itarsomere black.

Abdomen: Male terminalia (figs. 7-11): Length of
epandrium+surstyli in posterior view 1.5 X width;
lateral margins of surstyli only slightly concave ven-
trally (fig. 7), in lateral view digitiform, narrowly
rounded apically, curved anteroventrally but at same
curvature of epandrium (fig. 8); hypandrium in later-
al view sinuous basally, in posterior view broadly V-
shaped, rounded ventrally; aedeagal apodeme dis-
tinctly enlarged on 2/3 portion toward attachment
with hypandrium (fig. 9); postgonite with a postero-
ventral, relatively long, narrow, somewhat finger-like
process (fig. 9), in posterior view curved medially to-
ward apex, and apex acutely pointed; distiphallus dis-
tinctly angulate, dc sal angle acutely produced,
rounded (fig. 9), produced dorsolaterally at base, ap-
pearing wing-like in posterior view (fig. 10).

62

Other specimens examined

Afrotropical: Gambia. Bakau at tropic bungalow, (me-
adow rich in flowers at beach), 16-18 Nov 1977, L.
Cederholm, R. Danielsson, Hammarstedt, Hedqvist,
Samuelsson (296 , 47 ; BMNH , ZIL, USNM). Senegal. Dakar
N’Gor, sabbia nuda e talitr., 24 Jun 1973, A.G. Soika (34,
1; USNM); Dakar N’Gor, spiaggia sabb.-limo, 24 Jun
1973, A.G. Soika (19; USNM). Somone, spiaggia terr. crabs,
6 Jul 1973, A.G. Soika (46, 19; USNM).

Distribution (fig. 6)

Afrotropical. West Africa: Gambia, Senegal, Sierra
Leone.

Remarks

We directly compared the primary types of C. vil-
liersi and C. gioiellae and are of the opinion that they
are conspecific. The structures of the male terminalia
are virtually identical, and we are thus proposing the
synonymy noted above, with C. villiersi as the senior
synonym.

THE albipennis GROUP

Diagnosis

This species group is distinguished from other taxa
of Chlorichaeta by the following combination of char-
acters (synapomorphies, which provide evidence of
the group’s monophyly, are indicated by an asterisk):
mesonotum appearing granulose, microsculptured,
shiny; *distiphallus long and bearing long, paired
processes at base.

Chlorichaeta africana sp. n.
(figs. 12-17)

Type material. — Holotype d: “KENYA Lake Victoria
10km S Kisumu 19.X1.1986 [19 Nov 1986] A. Freidberg’;
13 paratypes (78, 6%; TAU, USNM) bear the same label data
as the holotype. Other paratypes are as follows: Cameroon.
Kribi (beach), 28-29 Nov 1987, F. Kaplan (26, 72; TAU,
USNM); Kribi, Rt. N7 (beach), 28-29 Nov 1987, A.
Freidberg (16, 49; USNM). Ghana. Kedzi-Keta (on coco-
nut), 29 Jul 1959 (Min. Agric. C.I.E. Coll. 16732 8377;
43,62; BMNH). Nigeria. Lagos, Victoria Island (shore), 15
Dec 1987, F. Kaplan (ld; USNM). The holotype is double
mounted (minuten in a block of polyporus), is in excellent
condition, and is deposited in the USNM.

Diagnosis

This species is very similar to C. tuberculosa in
having a black hind basitarsomere but is distinguis-
hed from the latter by characters of the male termi-
nalia, especially the shape of the surstylus, which, in
lateral view, is shallowly recurved apically and

MATHIS & ZATWARNICKI: Revision of Chlorichaeta

Figs. 12-16. Male genitalia of Chlorichaeta africana. — 12, epandrium and cerci, posterior view; 13, idem, lateral view; 14,
internal male genitalia, lateral view; 15, idem, ventral view; 16, hypandrium and postgonites, ventral view.

has a bluntly rounded apex.

Description

Small to moderately small shore flies, length 1.60-
2.60 mm.

Head: Mesofrons, especially extended ocellar trian-
gle, smooth, shiny, very similar to parafrons; parafa-
cials with silvery-white to white microtomentum ar-
ranged as 2 patches, dorsal patch much smaller, a spot
laterad of pedicel, ventral patch long, linear, length
equal to combined length of pedicel and flagellomere
1. Gena high, eye-to-cheek ratio averaging 0.60.

Thorax: Mesonotum microgranulose, scutum with
some faint metallic golden luster; scutellum with sim-
ilar microsurface as scutum, black, lacking metallic
luster, bearing 2 pairs of marginal setae that arise
from tubercles. Halter pale, whitish yellow to yellow.
Costal vein ratio averaging 0.71; vein M ratio averag-
ing 0.58. Fore tarsi black; hind basitarsomere entirely
dark coloured, blackish brown to brown.

Abdomen: Male terminalia (figs. 12-16): Length of
epandrium+surstyli in posterior view 1.65 X width;
lateral margins of surstylus in posterior view distinct-
ly though shallowly concave ventrally (fig. 12), base
of surstylus wide then distinctly narrowed at apical

63

Ty DSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 17. Distribution map for
Chlorichaeta africana.

1/3, apex less than greatest width of cercus; surstylus
in lateral view digitiform, base in same curvature as
epandrium, apex slightly recurved, narrowly spatulate
(fig. 13); hypandrium short, L-shaped in lateral view
with longer arm slender and more or less parallel with
basiphallus, lateral portion (= fused pregonite) more
or less as a parallelogram, more than twice width of
longer arm (fig. 14); aedeagal apodeme moderately
enlarged toward attachment with hypandrium (fig.
14); postgonite evenly lunate in lateral view (fig. 14);
distiphallus moderately long, slightly longer than bas-
iphallus, gently angulate in lateral view, lateral projec-
tions at base of distiphallus long, length equal to nar-
rowed portion of aedeagal apodeme (fig. 15),
basoventral margin of process distinctly and irregular-
ly serrate (figs. 14, 15).

Distribution (fig. 17)
Afrotropical. Cameroon, Ghana, Kenya, and Nigeria.

Etymology

The species epithet, africana, refers to the conti-
nent on which this species was collected.

Chlorichaeta albipennis Loew

(figs. 18-23)
Gymnopa albipennis Loew, 1848: 14. Syntypes from Italy.

Sicily: Messina [ZMHU, apparently lost].

Mosillus albipennis; Schiner 1864: 235 [generic combina-
tion].

Chlorichaeta albipennis; Cresson 1925: 235 [generic combi-
nation, review]; 1946: 244 [review], Cogan and Wirth
1977: 322 [Oriental catalogue], Cogan 1980: 656
[Afrotropical catalogue]; 1984: 128 [Palearctic cata-
logue], Canzoneri & Meneghini 1983: 229-230 [review,
Italy], Mathis 1989: 640 [Australasian/Oceanian cata-
logue].

Strandillus strandi Duda, 1942: 10. Lectotype d (designat-
edi ibys Pappy 197997) sande smc:
Duda/[Lithuania] Nidden [= Nida on Kurskaya Kosa
Peninsula] 4. 8. 40 [4 Aug 1940]/Lectotypus [red]’. The
abdomen of the lectotype has been removed, dissected
and is in a plastic microvial attached to the specimen pin.
The lectotype is deposited in the ZMHU. — Papp 1979: 97
[synonymy].

Mosillus subsultans of authors, not Fabricius (misidentifica-
tion); Miyagi 1977: 27.

Diagnosis

The species is similar to congeners of the a/bipennis
group but may be distinguished from them by the fol-
lowing combination of characters: scutellum bearing
3-5 large, marginal setae on each side that arise from
distinct tubercles; disc of scutellum distinctly more
microsculptured than scutum, appearing more granu-
lose; hind basitarsus black; characters of the male ge-
nitalia (see description below and figs. 18-22).

Figs. 18-22. Male genitalia of Chlorichaeta albipennis. — 18, epandrium and cerci, posterior view; 19, idem, lateral view; 20,
internal male genitalia, lateral view; 21, idem, ventral view; 22, hypandrium and postgonites, ventral view; 23, Distribution

map for Chlorichaeta albipennis.

64

Maris & ZATWARNICKI: Revision of Chlorichaeta

65

‘TijDSCH RIET VOOR ENTOMOLOGIE, VOLUME 136, 1993

Description

Small to moderately small shore flies, length 1.50
to 2.85 mm.

Head: Mesofrons, especially extended ocellar trian-
gle, shiny and smooth, similar to parafrons; silvery-
white microtomentum of parafacials as 2 patches:
dorsal spot much smaller, placed at level of pedicel
and a long linear patch at anteroventral margin of eye.
Gena high, eye-to-cheek ratio averaging 0.70.

Thorax: Mesonotum (colour and texture of scu-
tum and scutellum; scutellum distinctly granulose,
bearing 3-5 pairs of marginal setae that arise from tu-
bercles. Halter dark, brown to blackish brown (pale
coloured in specimens from Sri Lanka and Japan).
Costal vein ratio averaging 0.60; vein M ratio averag-
ing 0.58. Fore tarsi entirely black; hind basitarsomere
entirely dark coloured, blackish brown to brown.

Abdomen: Male terminalia (figs. 18-22): Length of
epandrium+surstyli in posterior view 1.6 X width;
lateral margins of surstylus in posterior view only
slightly concave ventrally (fig. 18), surstylus narrow,
apical 1/3 less than greatest width of cercus; surstylus
in lateral view digitiform, bluntly rounded apically,
curved anteroventrally but at about same curvature of
epandrium (fig. 19); hypandrium in lateral view L-
shaped with longer arm slender and more or less par-
allel with basiphallus, lateral portion (= fused pregon-
ite) more or less as a parallelogram, more than twice
width of longer arm (fig. 20); aedeagal apodeme dis-
tinctly enlarged toward attachment with hypandrium
(fig. 20); postgonite evenly lunate in lateral view (fig.
20); distiphallus short, less than length of basiphallus,
gently curved in lateral view, lateral projections at
base long, length equal to narrowed portion of aedea-
gal apodeme (fig. 20), basoventral margin of process
distinctly and irregularly serrate (fig. 21).

Other specimens examined

Oriental: India. Bombay, Juhu Beach, 7 Mar 1985, K.A.
Spencer (1 2; NMWL). Laos. Vientiane, 28 May 1965, P. D.
Ashlock (1; BBM). Sri Lanka. Colombo: Colombo, 14 Apr
1980, W. N. Mathis (12; USNM); Colpetty (103 Galle
Road), 3 Apr 1968, T. E. Halstead (138, 1725 CAs);
Negombo (6 km S), 7 May 1980, L. Jayawickrema, W. N.
Mathis, T. Wijesinhe (38, 52; USNM). Mannar: Mannar,
13 Apr 1968, T. F. Halstead (16, 39; CAS). Tricomalee:
Mutur, 2 May 1980, L. Jayawickrema, W. N. Mathis, T.
Wijesinhe (1d; USNM); Nilaveli (5 km N), 3 May 1980, L.
Jayawickrema, W. N. Mathis, T. Wijesinhe (346, 199;
USNM). Vietnam. Nha Trang, 17-26 Nov 1960, C.M.
Yoshimoto (1 d ; BBM). Palearctic: France. Herault: Etang de

Canet, stessa Straz., 27 Jul 1954, A. Giordani Soika (24;
BMNH ). Italy. Sicily: Capo San Vito, 29 Aug 1982, A.
Freidberg (12; USNM). Veneto: Lido Alberoni (Laguna di
Venezia), sentiero dune gulf, 21 Apr-4 Oct 1958, 1961, A.
Giordani Soika, Levrini (26, 29; USNM). Japan. Honshu:
Osaka, 29 Aug 1954, P. H. Arnaud, Jr. (226, 229; cas,
USNM). Okitsu (in cage), 27 Oct 1914, F. Muir (1 d ; BBM).
Russia. Primorski Krai (Maritime Territory): Glazkovka (20
km SW Valentin), 1-2 Sep 1986, A. Ozerov (3d, 29; ZM-
MU). Ukraine. Charkov: Charkov vicinity, 14 Sep 1883,
Jaroshevski (22; ZISP).

Distribution (fig. 23)

Oriental. India, Laos, Sri Lanka, Vietnam.
Palearctic. France, Italy, Japan (Hokkaido, Honshu,
Kyushu), Lithuania, Russia (Far East), Ukraine.

Remarks

Specimens of this species were frequently misiden-
tified as C. tuberculosa, and the two species are appa-
rently sympatric over some of their range (see records
from Italy, Canzoneri & Meneghini 1983).

The specimens from Sri Lanka and Japan have pale
coloured halteres but are otherwise similar, especially
characters of the male terminalia. Thus we attribute
the variation to be intraspecific.

Chlorichaeta mais sp. n.

(figs. 24-29)

Mosillus albipennis; Lamb 1912: 319 [misidentification,
Seychelles (Mahé)].

Type material. — Holotype d: ‘ALDABRA. Grande Terre:
Anse Mais[,] 17 March 1986[,] Wayne N. Mathis. The
holotype is double mounted (minuten in a block of plastic),
is in good condition, and is deposited in the USNM; 36 para-
types (128, 259) bear the same locality label as the holo-
type. — Other paratypes are as follows: Aldabra. Grande
Terre: Cinq Cases, 12-13 Mar 1986, W. N. Mathis (166,
149; USNM). Malabar: Passe Houareau, 14-18 Mar 1986,
W. N. Mathis (2d, 1 2; USNM). Picard: La Gigi, 19-20 Mar
1986, W. N. Mathis (18, 19; USNM); Settlement, 15-21
Mar 1986, W. N. Mathis (76, 21 2; USNM); trail to Jellyfish
Pool, 22 Mar 1986, W. N. Mathis (96, 11 9; USNM).

Additional specimens. — Cosmoledo. Menai Island:
Station, 26 Mar 1986, W. N. Mathis (1 2; USNM). Wizard
Island: 6 Mar 1968, B. H. Cogan, A. M. Hutson (ex;
BMNH). Astove. around coconut plantation, 5 Mar 1968, B.
H. Cogan, A. M. Hutson (4d, 379; BMNH ). Kenya.
Mombasa (150 km N), 4 Dec 1989, A. Freidberg, F. Kaplan
(1d; USNM). Madagascar. Antseranana: Nosy Be,
Ambatoloaka (beach), 4-7 Apr 1991, A. Freidberg, F.
Kaplan (56, 42; TAU, USNM); Nosy Tanikely, 6 Apr 1991,

Figs. 24-28. Male genitalia of Chlorichaeta mais. — 24, epandrium and cerci, posterior view; 25, idem, lateral view; 26, inter-
nal male genitalia, lateral view; 27, idem, ventral view; 28, hypandrium and postgonites, ventral view; 29. Distribution map

for Chlorichaeta mais.

66

Marais & ZATWARNICKI: Revision of Chlorichaeta

67

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

A. Freidberg, F. Kaplan (19; USNM); Nosy Komba, 6 Apr
1991, A. Freidberg, F. Kaplan (9d, 42; TAU, USNM);
Ramena, 10 Apr 1991, A. Freidberg, F. Kaplan (19; USNM);
Sambava Beach, 14 Apr 1991, A. Freidberg, F. Kaplan (49;
USNM). Fianarantsoa: Mananjary, 26 Oct 1988, R. Van
Epps (19; USNM). Toamasina: Fénérive (on beach), Dec
1955, B. Stuckenberg (1 2; USNM). Toliara: Berenty, 20 Apr
1991, A. Freidberg, F. Kaplan (1 2; USNM); Fort Dauphin (-
beach), 18 Apr 1991, A. Freidberg, F. Kaplan (4d, 89;
TAU, USNM); Fort Dauphin (50 km W, river), 20 Apr 1991,
A. Freidberg, F. Kaplan (16, 19; USNM); Morondava
(20°17°S, 44°16°E; sea level), 20 Jan 1985, F. M.
Harrington, J. D. Weintraub (49; USNM); Sud-Est Sainte-
Luce (10 m), Fort Dauphin, 22-24 Feb 1958, B.
Stuckenberg (1d; USNM). Seychelles. Mahé (12; BMNH ).
South Africa. Natal: St. Lucia Park, 7-8 Oct 1983, A.
Freidberg (1 2; USNM); Umkomaas (beach), 11 Oct-9 Dec
1954, 1983, A. Freidberg, B. Stuckenberg (664, 39 ; USNM);
Umhlanga Rocks, 4 Jan 1955, E. Callan (19; USNM);
Umhloti (mouth), 19 Jun 1962, B. & P. Stuckenberg (19;
USNM); Durban, The Bluff, Oct 1931, A. MacKie (16;
ANSP).

Diagnosis

This species is closely related to C. orba but is dis-
tinguished from that species by the granulose exten-
ded ocellar triangle and three versus two parafacial
microtomentose, silvery spots.

Description

Small to moderately small shore flies, length 1.60-
2.65 mm.

Head: Mesofrons, especially extended ocellar trian-
gle, granulose, contrasted with shiny, smooth para-
frons; silvery-white microtomentum of parafacials as
3 patches: 2 spots near level of antenna (dorsal spot at
level of ventral margin of pedicel, ventral spot at level
of midheight of Ist flagellomere) and a more linear
patch at anteroventral margin of eye (sometimes 2nd
spot and ventral, linear patch narrowly connected).
Gena high, eye-to-cheek ratio averaging 0.50.

Thorax: Mesonotum with faint to conspicuous
metallic bronzish-gold reflections; 2 marginal scutel-
lar setae on each side that arise from tubercles, several
smaller setulae between. Halter white. Costal vein ra-
tio averaging 0.63; vein M ratio averaging 0.45.
Basitarsomere of foreleg variable in colour, usually
dark, mostly blackish, at most very slightly lighter in
colour than other tarsomeres of foreleg, sometimes
yellowish; hind basitarsomere yellow, sometimes
slightly darker basally, mostly concolorous with tarso-
meres 2 and 3.

Abdomen: Male terminalia (figs. 24-28): Length of
epandrium+surstyli in posterior view 1.65-1.70 X
width; lateral margins of surstylus in posterior view
convex ventrally (fig. 24), broad, apical 1/3 wider
than cercus; surstylus in lateral view narrowly digiti-
form, narrowly rounded apically, shallowly curved
anteroventrally at about same curvature of epandrium
(fig. 25); hypandrium in lateral view very long and
sinuous, more or less in same conformation of basi-
phallus, lateral portion (= fused pregonite) more or
less as a parallelogram, only slightly wider than long
arm (fig. 26); aedeagal apodeme distinctly enlarged
toward attachment with hypandrium (fig. 26); post-
gonite evenly lunate in lateral view (fig. 26); disti-
phallus long, more than twice length of basiphallus,
sinuous in lateral view, apex recurved posterodorsally,
lateral projections at base of distiphallus long, length
equal to narrowed portion of aedeagal apodeme (figs.
26, 27), basoventral margin of process not serrate (fig.

27).

Distribution (fig. 29)

Afrotropical. Aldabra Group (Aldabra, Astove,
Cosmoledo), Kenya, Madagascar, Seychelles (Mahé),
South Africa (Natal).

Natural history

This species occurs in several habitats on atolls of
the Aldabra Group. We collected it most commonly
on tortoise droppings but also found it around pig
pens near the settlement (Picard Island) and occa-
sionally on rotting seaweed that had accumulated at

the high tide mark on the beach.

Etymology

The specific epithet, mais, alludes to the type loca-
lity of this species, Anse Mais, on the atoll of Aldabra
(Grande Terre). The species name is a noun in appo-
sition.

Remarks

The specimens from Aldabra, including the holo-
type, all have the fore basitarsomere mostly blackish,
similar in colour to the other tarsomeres of the fore-
leg. The states for this character are variable in speci-
mens from Madagascar and South Africa, however.
Some specimens have a pale, mostly yellowish basitar-

Figs. 30-34. Male genitalia of Chlorichaeta orba. — 30, epandrium and cerci, posterior view; 31, idem, lateral view; 32, inter-
nal male genitalia, lateral view; 33, idem, ventral view; 34, hypandrium and postgonites, ventral view; 35. Distribution map
for Chlorichaeta orba. The locality in Australia (Northern Territory) could not be precisely located.

68

MATHIS & ZATWARNICKI: Revision of Chlorichaeta

69

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

somere, but in others from the same series this struc-
ture is dark coloured, as in specimens from Aldabra.
We have dissected the male terminalia of the variable
series and see no differences and are thus of the opi-
nion that this variation is intraspecific.

Chlorichaeta orba sp. n.
(figs. 30-35)

Type material. — Holotype d': ‘SRI LANKA: TRI[NCOMAL-
EE]. Dist[rict]. Nilaveli (5 km N) 3 May 1980/Collectors:
W. N. Mathis[,] T. Wijesinhe[,] L. Jayawickrema’. Six par-
atypes (26, 49) bear the same label data as the holotype. —
Other paratypes are as follows: Sri Lanka. Colombo:
Ratmalana, Colombo (9 mi S), 7-13 Jan 1962, H.
Andersson, P. Brink, L. Cederholm (29; zit); Kalutara,
Colombo (25 mi SSE), 25 Jan 1962, H. Andersson, P.
Brinck, L. Cederholm (1 2; zi). Hambantota: Kirinda, 25
Apr 1980, W. N. Mathis (29; USNM). Uva: Heda Oya,
Bibile (29 mi SE), 7 Mar 1962, H. Andersson, P. Brinck, L.
Cederholm (16; ZIL).

Additional specimens. — Australasian Region: Australia.
Northern Territory: Wurruwardumanja, 15 Jan 1980, T.
Donovan (ld; ANIC). Papua New Guinea. Central
Province: Laloki (muddy river bank), 2 Jul 1984, J. W.
Ismay (16; USNM). Oriental Region: Philippines. Calicoan
(from dead land crab; sand near tent), 27 Jul 1945, F. F.
Bibby (83, 72; USNM).

Diagnosis

This species is similar to C. mais in having the hind
basitarsomere yellow but differs from the latter in the
following characters: extended ocellar triangle
smooth and shiny, similar to parafrons; parafacial
with 2 patches of silvery microtomentum; scutum
very lightly microsculptured, scutellum slightly more
so, appearing granulose; and mesonotum with some
metallic bluish luster.

Description

Small to moderately small shore flies, length 1.80-
2.60 mm.

Head: Mesofrons, especially extended ocellar trian-
gle, shiny and smooth, similar to parafrons; silvery-
white microtomentum of parafacials as 2 patches:
dorsal spot much smaller, placed at level of pedicel
and a long linear patch at anteroventral margin of eye.
Gena high, eye-to-cheek ratio averaging 0.47.

Thorax: Mesonotum with some metallic bluish re-
flections; scutum very lightly microsculptured, scutel-
lum more so, appearing granulose, bearing 2 pairs of
marginal setae that arise from tubercles. Halter yellow
to yellowish white. Costal vein ratio averaging 0.60;
vein M ratio averaging 0.55. Tarsi of foreleg with ba-
sal 2 tarsomeres yellow, apical 3 black; basitarsomere
of foreleg yellowish, contrasted with blackish apical

70

tarsomeres of foreleg; hind basitarsomere yellow,
sometimes slightly darker basally, mostly concolorous
with tarsomeres 2 and 3.

Abdomen: Male terminalia (figs. 30-34): Length of
epandrium+surstyli in posterior view 1.55 X width,
lateral margins irregularly tapered toward pointed
apex (fig. 30); surstylus in lateral view digitiform, nar-
rowly rounded apically, curved anteroventrally but
only slightly more curvature of epandrium (fig. 31);
hypandrium U-shaped in posterior view (fig. 34), in
lateral view L-shaped with anteromedial portion slen-
der, more or less parallel with basiphallus, tapered to-
ward apex, posterolateral portion (= fused pregonite)
as a parallelogram, more than twice width of antero-
medial portion (fig. 32); aedeagal apodeme distinctly
enlarged toward attachment with hypandrium (fig.
32); postgonite linear, lacking a process (fig. 32); dis-
tiphallus slightly longer than basiphallus, distinctly
angulate, dorsal angle acutely produced, rounded (fig.
32), bearing 1 paired slender, lateral processes at base
with anterodorsal orientation (fig. 33) and a single

medial process (fig. 32).

Distribution (fig. 35)

Australasian: Australia (NT), Papua New Guinea.
Oriental: Philippines, Sri Lanka.

Etymology

The specific epithet, orba, is derived from Latin
and means bereft of parents or an orphan.

Chlorichaeta tuberculosa Becker

(figs. 36-48)

Chlorichaeta tuberculosa Becker, 1922: 74. Lectotype 9
(designated by Cresson 1925: 234): ‘[Sudan. Khartoum
region] Sennar 18.-24.II. [handwritten] / Aegypt. Sudan
Ebner.1914/TYPE [red] / Ch. tuberculosa Beck. [handwrit-
ten] det. Becker’. The lectotype is double mounted (min-
ute nadel in plastic foam block), is in poor condition
(mesonotum damaged, right mid leg and Ist flagello-
meres missing), and is deposited in the NMW. — Cresson
1925: 234 [review]; 1945: 53 [review]; 1946: 245 [re-
view], Wirth 1955: 49 [list, Tanzania. Mugango (on Lake
Victoria)], Canzoneri & Meneghini 1969a: 1481 [list,
Senegal, figures of & genitalia]; 1969b: 167 [list, Zaire],
Cogan and Wirth 1977: 322 [Oriental catalogue], Cogan
1980: 656 [Afrotropical catalogue], Mathis 1989: 640
[Australasian/Oceanian catalogue].

Chlorichaeta albipennis of authors, not Loew (misidentifica-
tion). — Collin 1949: 202 [list, Egypt, Siwa], Tenorio
1980: 267-268 [list, Hawaii].

Diagnosis

This species is very similar to C. africana but is dis-
tinguished from the latter and other congeners of the

Maruis & Zarwarnickt: Revision of Chlorichaeta

b,

Figs. 36-41. Scanning electron micrographs of Chlorichaeta tuberculosa (Israel; scale length in parenthesis; bar scale for all pho-
tographs = fig. 36). 36, head, lateral view (0.27 mm); 37, idem, anterior view (0.30 mm); 38, frons, anterodorsal view (176
um); 39, antennae, anterior view (136 um): 40, face, anterior view (176 lim); 41, parafacial, anterior view (120 um).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 42-47. Scanning electron micrographs of Chlorichaeta tuberculosa (Israel; scale length in parenthesis; bar scale for all pho-
tographs = fig. 44). 42, mesonotum, dorsal view (0.40 mm); 43, scutellum, dorsal view (176 um); 44, left foreleg, anterior
view (150 um); 45, pleuron, lateral view (176 um); 46, notopleuron, lateral view (136 um); 47, 5th tergum of abdomen, dor-
sal view (75 Um).

72

albipennis group by the following combination of
characters: scutellum bearing 2 large, marginal setae
on each side that arise from tubercles; scutum and
scutellum about equally microsculptured and colou-
red; hind basitarsomere almostly entirely brownish
black to black; and surstylus in lateral view distinctly
and more or less evenly curved anteroventrally, tape-
red gradually and evenly to acutely pointed apex.

Description

Small to moderately small shore flies, length 1.65-
2.70 mm.

Head (figs. 36-41): Mesofrons, especially extended
ocellar triangle, smooth, shiny, very similar to para-
frons; parafacials with silvery-white to white microto-
mentum arranged as 2 patches, dorsal patch much
smaller, a spot laterad of pedicel, ventral patch long,
linear, length equal to combined length of pedicel
and flagellomere 1. Gena high, eye-to-cheek ratio av-
eraging 0.52.

Thorax (figs. 42-46): Mesonotum mostly black,
shiny, some specimens with faint, bronzish luster on
scutum; scutum and scutellum with similar texture,
both at most lightly granulose, bearing 2 pairs of mar-
ginal setae that arise from tubercles. Halter pale, whit-
ish yellow to yellow. Costal vein ratio averaging 0.64;
vein M ratio averaging 0.58. Fore tarsi black; hind
basitarsomere entirely dark coloured, blackish brown
to brown.

Abdomen (fig. 47): Male terminalia (figs. 48-52):
Length of epandrium+surstyli in posterior view 1.5 X
width; lateral margins of surstylus straight to concave
towards apex in posterior view (fig. 48), in lateral
view distinctly and evenly tapered to narrowly round-
ed apex, curvature in lateral view anteroventrad, dis-
tinctly more so than curvature of epandrium (fig. 49);
hypandrium short, curved in lateral view with antero-
medial portion more or less parallel with basiphallus
and tapered toward apex, dorsal portion (= fused
pregonite) as a parallelogram, more than twice width
of anteromedial portion (figs. 50, 52); aedeagal apo-
deme distinctly enlarged toward attachment with hy-
pandrium (fig. 50); postgonite evenly lunate in later-
al view (fig. 50); distiphallus longer than basiphallus,
gently curved along basal 2/3 in lateral view, recurved
apically, lateral projections from base long, length
equal to narrowed portion of aedeagal apodeme (figs.
50, 51), at base of process with a short membranous
lobe that bears a few finger-like serrations.

Other specimens examined

Afrotropical: Angola. Lobito Bay (on trunk of Melia aze-
darach), 19 Jul-10 Aug 1931, T. D. A. Cockerall, J. Ogilvie
(12, lex; ANSP). Ethiopia. Lake Langano (7°27'N,

Maruis & ZATWARNICKI: Revision of Chlorichaeta

39°11’E), 8 Feb-13 Dec 1973, 1989, R. Baker, A.
Freidberg, F. Kaplan (23; BMNH , USNM). Ghana. Cape St.
Paul, 22 May 1958 (19; BMNH ). Kenya. Tsavo National
Park, 4-7 Apr 1968, B. Cogan, A. Hutson (1d; BMNH ).
Malawi. Ruo ChiKonje (on cattle), 16 Dec 1915, R. C.
Wood (136, 69; BMNH , ANSP). Mozambique. Pafuri (hip-
po carcass), 21 May 1983, L. Braack (24, 49; USNM).
Nigeria. Cotonon, in Dahomey (70 mi W Lagos), 5 Jun
1914, W. A. Lamborn (19; ANsP). South Africa. Kruger
National Park (22°27°S, 31°17’E, across river from Pafuri,
on impala carcass), L. Braack (2d, 19; USNM). Sudan.
Erkowit, 17 May 1943, D.J. Lewis (16, 19; BMNH ).
Khartoum (sores on the backs of camels), 21 Feb-1 Mar
1932, 1933, H. W. Bedford (23, 59; BMNH , ANSP). Tuti
Island (attracted to human sweat), 20 Mar 1932, H. W.
Bedford (1d, 42; BMNH ). Wad Medani (on dead goat), 27
Mar 1940, D. J. Lewis (19; BMNH ). Yemen. Aden, 3 Dec
1951 (16; USNM); Kirsh, Wadi ‘Aqqan (3000 ft), 17 Mar
1940, P. W. R. Petrie (1d; BMNH ). Socotra Island: Ras
Shóab, Feb 1899, O. Simony (1d; ANSP). Australasian/
Oceanian: Guam. Agana (sea cucumber), 3 Aug 1945, G. E.
Bohart, J.L. Gressitt (19; ANSP); Antigua, 2 Aug 1945, G.
E. Bohart, J. L. Gressitt (1 2; USNM); SE coast, 9 May 1945,
G. E. Bohart, J. L. Gressitt (1d, 19; USNM); Pt Oca, May
1945, G. E. Bohart, J. L. Gressitt (19; USNM). Hawaii.
Hawaii: N Kona, Makalawena Tract, 12 Feb 1970, D. E.
Hardy (1d, 19; USNM). Maui: Lahaina (2 km S; on herbs
and grass along sea shore), 23 Oct 1981, L. & I. Cederholm
(26,39; BMNH ). Oahu: Fort Kam, 4 Jul 1944, D. G. Hall
(22; USNM). Oriental: India. Madras, Sidhout (bred in rot-
ting oilcake manure), 9 Mar 1934, C. S. B (32; BMNH ,
ANSP). Sri Lanka. W. Province: Ratmalana, Colombo (9 mi
S), swept on dry grassland, or on herbs on sandy beach, 7-13
Jan 1962, H. Andersson, P. Brinck, L. Cederholm (64,89;
ZIL); Kalutara, Colombo (25 mi SSE), swept on dry grass
on sandy beach, 25 Jan 1962, H. Andersson, P. Brinck, L.
Cederholm (16, 29; ZIL); Beruwala, Colombo (34 mi
SSE), 25 Jan 1962, H. Andersson, P. Brinck, L. Cederholm
(16; ZIL). Thailand. Thonburi Province: Mueng District,
Manop, 7 May 1959 (12; USNM). Rajburi, Banpongnoung,
29 Jun-3 Jul 1952, R. E. Elbel (12; USNM). Vietnam.
Saigon, Cho Binh Tay, 15 May 1973, R.E. Parsons (1d,
12; USNM). Palearctic: Afghanistan. Khourmalik (between
Farah and Dilaram, 800 m, under stones), 6 Sep 1957, K.
Lindberg (2; USNM). Algeria. Kairouan, 7 Aug, F. Santchi
(1d; ANSP). Egypt. Egypt (1d; ANSP). Giza Province:
Manshiet Radwan, 28 Apr 1965, K.V. Krombein (16;
USNM); Siwa, 15 Jul 1935, J. Omer-Cooper (7d, 92; BMNH
); Wadi el Natrun (N Cairo; sores on donkey), 20 Jul 1950,
C. W. Sabrosky (34, 52; USNM); Wadi Ghoderat Kosaima
(tamarix), 1934, Priesner (16; USNM). Saqgra, 10 May
1966, P. M. Marsh (16, 22; USNM). Sinai: Bir Tmade, 8
Sep 1977, D. Simon (2d; TAU); Dahab, 7 Apr-23 May
1973, 1981, A. Freidberg, M. Kaplan, W. N. Mathis (2d,
39; TAU, USNM); ‘En Qsaib, 15 Mar 1982, I. Yarom (19;
TAU); Wadi Firan (oasis), 9 Apr-29 July 1950, 1971, 1973,
A. Freidberg, C. W. Sabrosky (1d, 42; TAU, USNM); 1950,
C. W. Sabrosky (19; USNM); Mt. Abas, 14 Aug 1971, J.
Kugler (1d, 19; Tau); Muyat-Hara, 12 Aug 1971, J.
Kugler (1d, 22; TAU); Ofira (sewage area), 22 Mar-21 May
1981, A. Freidberg, W. N. Mathis (49d, 169 ; TAU, USNM);
Neviot, 14 May 1981, T. Furman (1d; TAU); Ras Burka, 20
May-5 Sep 1976, 1981, A. Freidberg (1d, 12; TAU); Sinai
Mountains, St. Katharina, 18 Jul 1974, F. Kaplan (1°;
TAU); Sinai Mountains, Wadi Tlach (1500 m), 15 Jul 1974,

Me

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 48-52. Male genitalia of Chlorichaeta tuberculosa. — 48, epandrium and cerci, posterior view; 49, idem, lateral view; 50,
internal male genitalia, lateral view; 51, idem, ventral view; 52, hypandrium and postgonites, ventral view.

A. Freidberg, F. Kaplan (29; TAU); Wadi Mesech, 14 Apr
1981, T. Furman (2d; TAU); Wadi Watir, 6 Apr-5 Aug
1973, 1975, A. Freidberg, M. Kaplan (15d, 129; TAU).
Khor Gwob, 25 Nov 1930, R. C. M. Darling (16, 29;
BMNH ). Greece. Macedonia, 1933, R. C. Shannon (1%;
USNM); Larissa, Platamon, 5-9 Sep 1965, B. H. Cogan (46,
1®; BMNH ). Israel. Arad, 21 Apr 1981, F. Kaplan, A.
Freidberg (35, 19; TAU); Dead Sea, 11-18 Oct 1939, E.
Rivnay (56,39; ANSP, TAU); ‘En ‘Avedar, 16-29 Mar 1980,
1988, A. Freidberg, W. N. Mathis (66, 99; TAU, USNM);
‘En Bogéq, 30 Jul-20 Sep 1971, 1977, A. Freidberg, J.
Kugler (16, 19; TAU); ‘Ein Fashkha (Dead Sea at ‘Enot
Zugim), 2 Jul 1946, O. Theodor (58, 22; AU); ‘En Gedi,
21 Mar 1980, A. Freidberg, W. N. Mathis (89 ; USNM); ‘En
Mor, 31 Mar-30 Oct 1981, 1984, 1986, A. Freidberg (7,

74

59; AU); Hameshar, 16 Mar 1988, A. Freidberg (2d;
AU); Hazeva, 21 Apr 1981, A. Freidberg (ld; TAU);
Jerusalem (Mt. Scopus), 9 Sep 1935, O. Theodor (2d;
TAU); Kefar Yeroham, 22 Jul 1962, J. Kugler (3d, 12;
TAU); Mikhmoret, 28 Aug 1980, C. Nagy (1 2; TAU); Nahal
‘Iddan, 22 Mar 1980, A. Freidberg, W. N. Mathis (20,19;
USNM); Nahal Ramon, 15-30 Oct 1975, 1984, A. Freidberg
(48,6%; TAU); Nahal Zibor, 17 Mar 1988, A. Freidberg, F.
Kaplan (4d, 39; rau); Ne’ot ha Kikkar, 21 Mar 1980, A.
Freidberg, W. N. Mathis (1d; USNM); Nizzanim, 4-12 Oct
1983, I. Nussbaum (4d; TAU); Qalya, 8 Mar 1976, A.
Freidberg (1 2; AU); Taba, 28 Apr 1974, A. Freidberg (1 2;
AU); Wadi Yamin, 19 Oct 1983, I. Nussbaum (1 ®; TAU).
Italy. Sardinia: Rio Geremeas, 23 Jul 1952, Th.G. Aitken
(93, 72; USNM). Oman. Al Khoudh, 11 Nov 1988, M. J.

MarHis & ZATWARNICKI: Revision of Chlorichaeta

Fig. 53. Distribution map for Chlorichaeta tuberculosa.

Ebejer (1d, 1 2; NMWL); Al Seefa, 4 Dec 1987, M. J. Ebejer
(19; NMWL); Ghuzayn, date culture, 10 Apr 1985, P. Ardö
(18; ZIL); Nizwa-Tanuf road, 12 Apr 1985, P. Ardö (19;
ZIL); Wadi Daigah, 23°06 N, 58°52’E, 27 Jan 1989, M. J.
Ebejer (22; NMWL).

Distribution (fig. 53)

Afrotropical. Angola, Cape Verde Islands, Ethio-
pia, Ghana, Kenya, Malawi, Mozambique, Nigeria,
South Africa, Sudan, Tanzania (no specimens seen),
Yemen (including Socotra). Australasian/Oceanian.
Guam, Hawaiian Islands. Oriental. India, Sri Lanka,
Thailand, Vietnam. Palearctic. Afghanistan, Algeria,
Bulgaria (no specimens seen), Canary Islands (no
specimens seen), Egypt, Greece, Israel, Italy, Oman.

ACKNOWLEDGEMENTS

We are grateful to the curators and their respective
institutions for the loan of specimens that were bor-
rowed in conjunction with this study (a listing of
acronyms, collections, and curators are noted in the
introduction of this paper). H. B. Williams prepared
the distribution maps, and Susann Braden and Victor
Krantz assisted with the preparation of the scanning
electron micrographs. We are also grateful to David
Challinor, former Assistant Secretary for Research,
Smithsonian Institution, and Stanwyn G. Shetler,
Deputy Director of the National Museum of Natural
History, for financial support to conduct field work
and study primary types through grants from the
Research Opportunity Fund. Curtis W. Sabrosky

hand carried the holotype of C. villiersi from MNHN
to Washington, D. C.

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Becker, T. 1922. Wissenschaftliche Ergebnisse der mit
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Canzoneri, S. 1981. Ricerche condotte dal Prof. A.
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di Storia Naturale di Venezia 31: 201-221.

Canzoneri, S. 1982. Ephydridae e Canaceidae della Sierra
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Canzoneri, S. & D. Meneghini. 1969a. Ephydridae [Dipt.]
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Canzoneri, S. & D. Meneghini. 1983. Ephydridae e
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Canzoneri, S. & L. Rampini. 1990. Appunti sugli
Ephydridae e Canaceidae della Sierra Leone (Diptera,
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Cogan, B. H. 1980. 71. Family Ephydridae. Pp. 655-669. —

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Afrotropical Region. British Museum (Natural History),
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Cogan, B. H. & W. W. Wirth. 1977. Family Ephydridae.
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Collin, J. E. 1949. Results of the Armstrong College
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the Indian Ocean in 1905, under the leadership of Mr. J.
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Ephydridae), I: A revision of the shore fly genus
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western palearctic species of Athyroglossa (Diptera:
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on Ephydridae (Diptera). — Proceedings of the Biological
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1993. Studies of Gymnomyzinae (Diptera: Ephydridae),
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Hardy & M. D. Delfinado, eds., Insects of Hawaii.
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451 pp.

Wirth, W. W. 1955. East African Ephydridae (Ins.,
Diptera). — Mitteilungen aus dem Zoologischen Museum
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based on phylogenetic reconstruction (Diptera:

Cyclorrhapha). — Genus 3 (2): 65-119.

Received: 25 January 1993
Accepted: 8 April 1993

THOMAS PAPE! & HIROSHI SHIMA?

‘Zoological Museum, Copenhagen, Denmark

Biological Laboratory, College of General Education, Kyushu University, Japan

A NEW GENUS OF TACHINIDAE FROM

THE PHILIPPINES (DIPTERA)

Pape, T. & H. Shima, 1993. A new genus of Tachinidae from the Philippines (Diptera). —
Tijdschrift voor Entomologie 136: 77-81, figs. 1-5. [issN 0040-7496]. Published 1 July 1993.
A new genus of Tachinidae, Aporeomyia gen. n., including only the type species Aporeomyia
antennalis sp. n., is described from the Philippines. It is placed in the subfamily Tachininae, and
possible phylogenetic relationships to other tachinine genera are discussed.

Correspondence: Dr. T. Pape, Zoological Museum, Universitetsparken 15, DK - 2100

Copenhagen, Denmark.

Key words. — Tachinidae, Philippines, new genus, new species.

The present study deals with two male specimens
belonging to the Tachinidae family-group and show-
ing a tripartition of the first flagellomere. Although
possibly sexually dimorphic with the female first fla-
gellomere unpartitioned, this feature makes the spe-
cies easily recognizable (fig. 3). Tripartition of the
first flagellomere occurs within the Tachinidae fami-
ly-group in the family Tachinidae and in some unde-
scribed Australian—New Guinean taxa of uncertain
familial affiliation (see discussion below). However,
the configuration of the tripartition is never fully
identical to that seen in the present species. We take
this opportunity to describe and name the taxon, and
as is discussed in detail below the species does not fit
very well into any of the named genera, for which rea-
son a new genus is described.

TAXONOMIC PART

Aporeomyia gen. n.

Type species: Aporeomyia antennalis sp. n.
Etymology. — Gender: Feminine. Composed from
the type locality, Mt. Apo of Mindanao, and the

Greek, oreos, oros = mountain, and myia = fly.

Diagnosis. — Male: Head profile narrowed ventral-
ly. Proboscis and palpus short. Antenna very large
and with first flagellomere tripartite from base. Arista
inserted on upper lobe at a point about 0.3 from base
to tip. Antennal scape and pedicel raised above level
of lunule. Prosternum bare. Subscutellum slightly
swollen. Wing vein R, setulose in distal part, vein M
almost straight and withour a bend distal to cross-
vein dm-cu, and vein Cu+A, extended to wing mar-

gin. Lower calypter narrow and with median margin
diverging from scutellum. Hind tibia with postero-
ventral apical seta.

Aporeomyia antennalis sp. n.
(figs. 1-5).

Type material. — Holotype d: PHILIPPINES, Mindanao,
Mt. Apo, Agko - Lake Binado, 1,350-2,300 m, 5.viii.1978,
A. Nakanishi & O. Yata. — Paratype d, data as holotype,
terminalia dissected and stored in a microvial pinned with
the specimen. Both types are deposited in the Biological
Laboratory, Kyushu University, Fukuoka, Japan.

Etymology. — From the Latin, antenna = feeler, -is
= adjectival suffix meaning with or having. The name
refers to the first flagellomere, the peculiar shape of
which makes the species easily recognizable.

Description. — Male. Head brown in ground
colour, without pollinosity; frontal vitta dark brown;
face pale yellowish; scape and pedicel reddish, first
flagellomere brown; palpus pale yellowish. Head pro-
file short and high, strongly narrowed ventrally; frons
about 0.4x as long as height of face profile; vertex
wide, 0.47-0.50x head width; frontal vitta weakly
widened anteriorly, about 3x as wide as fronto-orbital
plate at middle; parafacial in profile very narrow; ge-
na slightly less than 0.5x eye height, genal dilation
obliquely occupying about lower 0.3 of gena; mouth
opening reduced, about 0.75x as long as wide. Inner
vertical seta strong, slightly less than eye height; 2 fine
divergent postocellar setae; ocellar seta fine, procli-
nate, subequal in length to postocellar seta; no recli-
nate and proclinate orbital setae; 3-4 frontal setae,
lowest seta nearly level with base of pedicel; 2-3 irreg-
ular rows of short fine hairs on anterior portion of

77

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-5. Aporeomyia antennalis. — 1, right wing, dorsal view; 2, epandrium (in part), cerci and right surstylus; posterior
view; 3, male head; 4, epandrium, surstylus and cercus, lateral view; 5, hypandrium, pre- and postgonite and aedeagus, later-

al view.

fronto-orbital plate descending to upper 0.25 of para-
facial; vibrissa fine and short, slightly more than 0.3x
length of face; facial ridge with several fine short hairs
just above vibrissa; occiput with short black hairs.

78

- Antenna long, distinctly longer than face; first flagel-

lomere tripartite, each lobe weakly widened apically,
middle lobe slightly longer than others. Arista short,
about 0.4x as long as upper lobe of first flagellomere,

inserted on basal 0.3 of upper lobe of first flagello-
mere; second aristomere about 2x as long as wide.
Mouthparts strongly reduced; proboscis very short; la-
bella small; palpus short, subequal in length to pedicel.

Thorax brown on dorsum except for pale brownish
postpronotal lobe, pale brownish on pleura except for
brownish posterior 0.5 of anepisternum and anterior
0.3 of katepisternum, without pollinosity. Dorsum
with rather sparse short fine hairs except for bare area
between rows of dorsocentral and intra-alar setae of
postsutural scutum, pleura without hairs other than
strong setae; 1 postpronotal seta; 2 notopleural setae;
no acr seta; 1-2 + 2-3 dc setae; 0 + 1 ia seta; 1 supra-
alar seta; proepisternum bare, with 1 fine proepister-
nal seta; 1 fine proepimeral seta; 2 + 2-1 katepisternal
setae; katepimeron and katatergite bare; scutellum
with 2 pairs of marginal setae, strong subapicals and
short apicals; discal scutellar setae absent. Metatho-
racic spiracle rather small, anterior and posterior
fringes subequal and not standing out from spiracular
rim.

Wing hyaline, evenly tinged with pale brown; tegu-
la and basicosta dark brown; lower calypter pale
brownish yellow. Vein R,,, joins costa near wing apex,
giving a long third costal sector slightly more than 4x
length of second; vein R,,; joins costa slightly beyond
wing apex; vein M straight from crossvein dm-cu to
wing margin; last section of vein CuA, short, slightly
more than 0.25 of crossvein dm-cu. Costa setulose
dorsally and ventrally; vein R, setulose dorsally on
distal half; vein R,,, ventrally with 1-2 setulae at base,
dorsally with several setulae at base as well as setulae
from just_proximal to crossvein r-m and almost to
wing margin. Halter large, about 0.7x as long as low-
er calypter.

Legs pale brownish yellow; pulvilli whitish. Fore
tibia with 1 fine p seta; mid tibia with 0 ad, 2 pd and
1 v setae. Claws and pulvilli very short.

Abdomen brownish in ground colour, without
pollinosity; anterior 0.4 of syntergum 1+2, anterior
0.3 of terga 3-4 and anterior 0.25 of tergum 5 pale
brownish; mid dorsal stripe dark and narrow; synter-
gum 1+2 rather long, only faintly excavated at base;
sterna 2-3 exposed. Hairs short, fine, dense and re-
cumbent on dorsum, longer and sparser on venter;
tergum 2 with 2-3 lateral discal, 2 lateral marginal
and 2 median marginal setae, all setae strong; tergum
3 with 2 rather fine lateral discal, 2 strong median and
lateral discal setae; terga 4-5 each with 1-2 fine lateral
discal setae and a row of strong marginal setae; sterna
without strong setae.

Male genitalia. Cercal bases fused medially, apical
prongs well separated and pointed; surstylus narrow
and longer than cerci; epiphallus present; distiphallus
rather short, with tapering unpaired dorsomedian
process.

PAPE & SHIMA : New genus of Tachinidae

Length. Body, 3.7-4.5 mm; wing, 3.4-4.3 mm.

Female. — Unknown.

Biology. — Unknown.

Distribution. — Oriental
(Mindanao).

Region: Philippines

SYSTEMATIC DISCUSSION

Aporeomyia antennalis is unquestionably a member
of the Tachinidae family-group or Oestroidea, the
monophyly of which is discussed at length elsewhere
(Griffiths 1972, McAlpine 1989, Pape 1992).
Families of the Tachinidae family-group are in the
main well defined, but many genera showing a com-
bination of a slightly swollen subscutellum, oval or
tongue-shaped lower calypteres, small metathoracic
spiracles, and an obtuse (or missing) bend of vein M
have been, and are still being, shuffled around
amongst the tachinoid families, e. g., Mimodexia
Rohdendorf (currently Rhinophoridae, Tschorsnig
1985a), Bezzimyia Townsend and Malayia Malloch
(Tachinidae or Rhinophoridae, Pape 1992), and
Morinia Robineau-Desvoidy (currently Calliphori-
dae, Rognes 1991). Aporeomyia antennalis possesses
these four character states, which may be apomorphic
groundplan states of the Rhinophoridae, and it has
accordingly a rhinophorid appearance. The structure
of the aedeagus, however, provides no support for a
rhinophorid affiliation. In the Rhinophoridae, the ae-
deagus is generally slender, always equipped with a
long epiphallus, and usually with paired dorsolateral
processes. In Aporeomyia the aedeagus is rather short
with a short and compact epiphallus, and with an un-
paired, tapering dorsomedian process (‘Fortsatz des
Dorsalsklerits of Tschorsnig [1985a, b]). Pape
(1986: 27) recognized a single monophyletic group
within the Rhinophoridae, defined by an unpaired
dorsomedian process, but this group is part of a more
inclusive group sharing a tripartite (and apomorphic)
acrophallus not found in Aporeomyia. The ground-
plan of the possibly monophyletic Sarcophagidae +
Tachinidae is characterized by an unpaired dorsome-
dian process (Pape 1992), but Aporeomyia does not
share any synapomorphies with any of the sarcophag-
id subfamilies and cannot be included in this family.
It is our experience that the male terminalia in gener-
al provide much more substantial evidence of phylo-
genetic relationships than the above-mentioned rhi-
nophorid-like features. Thus, we consider an
assignment to the Tachinidae as most probable, even
though we have no clearcut evidence in support of
this.

Fitting Aporeomyia antennalis into the subfamilial
classification of the Tachinidae with no information
on the structure of female terminalia, female repro-
ductive system, and immatures (egg, first instar larva)

79

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

has to be preliminary. Of the four subfamilies gener-
ally applied, e. g. by Tschorsnig (1985b) and Wood
(1987), perhaps only the Dexiinae and the Phasiinae
are monophyletic taxa. Indeed, it has been stressed
that at least the subfamily Tachininae very probably is
not monophyletic (‘sehr wahrscheinlich nicht
monophyletisch’; Tschorsnig [1985b: 119]) or even
‘almost certainly polyphyletic’ (Crosskey 1980: 822-
823).

Aporeomyia antennalis does not possess the aedeagal
hinge characterizing the groundplan of the Dexiinae,
nor any of the derived character states listed for the
Phasiinae by Tschorsnig (1985b: 121). Also, it does
not possess a setose prosternum as found in many
(but not all) Exoristinae. Aporeomyia antennalis will
key out to subfamily Tachininae in Crosskey’s (1976:
11) comprehensive (given as ‘alternative’) key to sub-
families of Oriental Tachinidae, but mainly because it
repeatedly fits the entry: “Forms without such combi-
nation of characters present simultaneously’. Note
that Pape (1992) used a broad concept of the
Tachininae, including all species which embryonate
their eggs in the uterus. With only males of
Aporeomyia available, nothing can be deduced on fe-
male reproductive biology. We have chosen to place
Aporeomyia in the Tachininae as this does not conflict
with the general morphology of this subfamily
(whether in the traditional sense or the broader sense
used by Pape [1992]).

Assigning antennalis to a genus of its own deserves
an explanation as monotypy in itself conveys little, if
any, information. Often - as here - monotypic genera
are erected to accommodate aberrant species which
present insufficient and conflicting evidence for in-
clusion within already named genera.

The tripartite first flagellomere — in so far as this is
of very rare occurrence in the Tachinidae — may be
taken as evidence of phylogenetic relationship.
Trischidocera sauteri Villeneuve is the only other
Oriental species with a tripartite first flagellomere,
and the head profile of Aporeomyia antennalis more-
over resembles that of 7. sauteri to the extent that
both have the antennal insertion at or slightly above
the level of upper eye margin, an almost horizontal
frons, a short lower head margin and very short
mouthparts (compare Crosskey 1976: fig. 50 with the
present fig. 3). Many of these features, however, may
be considered constraints imposed by the very long
and tripartite first flagellomere. It is important to
note that the first flagellomere of A. antennalis and T.
sauteri reveals some structural differences: Firstly, 7.
sauteri has the first flagellomere basally bipartite with

the lower lobe (i. e., that closest to the facial plate) -

further bipartitioned at a point about one fifth from
its base, while in A. antennalis the three lobes of the
first flagellomere all originate from a common point

80

at the base — or perhaps with the middle and upper
lobes sharing a very short common base. Secondly,
the arista is terminal in 7. sauteri but inserted at 0.3
from the base in A. antennalis. Thus, for the triparti-
tion to be homologous, i.e., for the two configura-
tions to be connected in a transformation series, it is
necessary to assume some fairly dramatic reorganiza-
tions of the first flagellomere. It may thus be at least
as parsimonious to put up a transformation series
between the bipartite first flagellomere of Diglossocera
bifida Wulp (Crosskey 1976: fig. 51) and the tripar-
tite flagellomere of A. antennalis. This is not to say
that dramatic reorganizations are impossible; only
that tripartition per se is difficult to use as a character
(state) and consequently as an argument for phyloge-
netic relationship.

Whether or not Zrischidocera sauteri and T. atratu-
la (Malloch) are distinct species (see note by Crosskey
1976: 210) has no relevance for the present discus-
sion, but when Chao & Zhou (1987) described 7.
yunnanensis, which has a markedly different head pro-
file and a simple, and much shorter, first flagellomere,
the genus seems defined mainly by the unique posi-
tion of the arista apically on the first flagellomere.
The displacement of the arista towards the distal part
of the first flagellomere in A. antennalis is in itself
evidence of phylogenetic relationship with Trischido-
cera, but a similar, although less pronounced, dis-
placement occurs in the Oriental genera Ger-
mariochaeta Villeneuve (Crosskey 1976: fig. 39) and
Lophosiosoma Mesnil, which constitute the tribe
Germariochaetini. This tribe is interesting as the low-
er calypteres of its members are narrow and diverging,
as is shown by Villeneuve (1937: fig. 1) for
Germariochaeta clavata Villeneuve, and thus repre-
sent a possible synapomorphy with Aporeomyia an-
tennalis. No further evidence, however, corroborates
a sister-group relationship between Aporeomyia an-
tennalis and the morphologically very different
Germariochaetini, and the latter taxon is often con-
sidered closely related to Triarthria Stephens and its
relatives, or to the Minthoini (Crosskey 1976).

A slightly swollen subscutellum, the small meta-
thoracic spiracle, the narrow, diverging lower calyp-
teres, the reduced bend of vein M, and the extension
of the first anal vein towards the wing margin are
character states shared with the Oriental Malayia.
This genus was listed by Crosskey (1976: 163) as a
tribally unplaced genus within the Tachininae to-
gether with Zrischidocera. At least one of the two spe-
cies included in Malayia possesses an aedeagus with
separate dorsolateral processes, which have the ex-
treme tip free of the aedeagal wall (the only male spec-
imen of Malayia known is the single M. fuscinervis
Malloch reported by Dear & Crosskey [1982: 134];
examined by TP). The aedeagus of Malayia thus pro-

vides a character state that, in combination with the
rhinophorid-like structure of metathoracic spiracle,
subscutellum, wing venation, and lower calypteres,
actually is strong indication that Malayia should be
transferred to the Rhinophoridae. Similarities also ex-
ist between Aporeomyia antennalis and the Australian-
New Guinean taxon containing a dozen undescribed
species and tentatively put in a separate (unnamed)
tachinoid family by Colless & McAlpine (1991).
Apart from possessing all the rhinophorid-like fea-
tures mentioned above, these undescribed species
have the antennal arista displaced towards apex,
sometimes inserted very close to the tip (Colless &
McAlpine 1991, fig. 39.38: c, d). Bi- and tripartition
occurs in some of the undescribed species but is here
of a very different appearance. Also, as in Malayia, the
aedeagus has paired dorsolateral processes, which here
is considered evidence that these species may possibly
not belong to the Tachinidae.

In conclusion, we have not been able to find evi-
dence in the morphology of Aporeomyia antennalis for
a reliable inclusion within any named genus.
Tripartition of the first flagellomere gives the species
a superficial resemblance to the similarly equipped
Trischidocera sauteri, but differences in configuration
of the lobes weakens a hypothesis of homology. Other
character states suggest a hypothesis of a sister-group
relationship to the Germariochaetini, but several of
the characters discussed are notoriously much prone
to evolutionary homoplasy.

ACKNOWLEDGEMENTS

Dr. H.-P. Tschorsnig, Stuttgart, Mr. Stig
Andersen, Copenhagen, and Dr. Verner Michelsen,
Copenhagen, kindly read the manuscript and their
suggestions are highly appreciated. We are also grate-
ful to Mr. A. Nakanishi, Hyogo Natural History
Museum, and Dr. O. Yata, Kyushu University, for

making the material available.

REFERENCES

Chao, C.- M. & S. Zhou, 1987. New species of tachinid
flies from Hengduan Mountains of China (Diptera:

PAPE & SHIMA : New genus of Tachinidae

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with English summary].

Colless, D. H. & D. K. McAlpine, 1991. Diptera (Flies). —
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of Australia. Vol. 2. Melbourne University Press, vi +
543-1137.

Crosskey, R. W., 1976. A taxonomic conspectus of the
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of the British Museum (Natural History) Entomology
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Crosskey, R. W., 1980. Family Tachinidae. — Pp. 822-882
in R. W. Crosskey (ed.), Catalogue of the Diptera of the
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Dear, J. & R. W. Crosskey, 1982. A taxonomic review of the
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Griffiths, G. C. D., 1972. The phylogenetic classification of
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340 pp.

McAlpine, J. F., 1989. Phylogeny and classification of the
Muscomorpha. — Pp. 1397-1505 in J.F. McAlpine (ed.),
Manual of Nearctic Diptera. Vol. 3. Research Branch,
Agriculture Canada, Monograph No. 32, vi + 1333-
1581.

Pape, T., 1986. A phylogenetic analysis of the woodlouse-
flies (Diptera, Rhinophoridae). — Tijdschrift voor
Entomologie 129: 15-34.

Pape, T., 1992. Phylogeny of the Tachinidae family-group
(Diptera: Calyptratae). — Tijdschrift voor Entomologie
135: 43-86.

Rognes, K., 1991. Blowflies (Diptera, Calliphoridae) of
Fennoscandia and Denmark. — Fauna Entomologica
Scandinavica 24: 1-272.

Tschorsnig, H.-P., 1985a. Die Struktur des männlichen
Postabdomens der Rhinophoridae (Diptera). —
Stuttgarter Beiträge für Naturkunde (A) 375: 1-18.

Tschorsnig, H.-P., 1985b. Taxonomie forstlich wichtiger
Parasiten: Untersuchungen zur Struktur des männlichen
Postabdomens der Raupenfliegen (Diptera, Tachinidae.).
— Stuttgarter Beiträge für Naturkunde (A) 383: 1-137.

Villeneuve, J., 1937. Myodaires supérieurs de Chine. —
Bulletin de la Musée Royal d'Histoire Naturelle Belgique
13(34): 1-16.

Wood, D. M., 1987. Tachinidae. — Pp. 1193-1269 in J. F.
McAlpine (ed.), Manual of Nearctic Diptera. Vol. 2.
Research Branch, Agriculture Canada, Monograph No.
28, vi + 675-1332.

Received: December 1992
Accepted: 26 January 1993

81

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

BOOK REVIEWS

Scoble, Malcolm J., 1992. The Lepidoptera: Form, function
and diversity. — Natural History Museum Publications,
Oxford University Press. 404 pp., 321 figs., 4 colour
plates. [ISBN 0 19 854031 0]. Price £ 45.

This book is the third volume in a series, issued by
the Natural History Museum of London, the former
British Museum (Natural History), providing over-
views of the insect orders. Unlike the two previous
volumes (on Hymenoptera and on Hemiptera),
which are primarily based on the British fauna, “The
Lepidoptera’ treats the order on a global scale.

More than half the book is devoted to general
chapters, dealing extensively with morphology, im-
mature stages, hearing sound and scent and the eco-
logical importance of Lepidoptera. The remaining
part of the book deals with the lepidopteran super-
families and families. For all families a short descrip-
tion of adult and immature stages is given and biolo-
gy, phylogenetic relationships and classification are
briefly discussed. For some larger families, the sub-
families are treated separately. An extensive list of ref-
erences and an index conclude the book.

The four colour plates show adults in resting pos-
ture, various types of larvae, larval habits and eggs. An
adult example of most families is shown in 17 black
and white plates, with photos of mounted specimens.

Scoble has succeeded in bringing together a wealth of
information on the whole order, including much recent
work, yet the texts remain concise and clear. Especially
the general chapters give much detailed information on
lepidopteran structure and function. The family treat-
ments differ considerably, undoubtedly because for
some families much more data is available than for oth-
ers. What is missing compared to the earlier volumes are
references to identification guides. However, this is
quite understandable on the global scale, because the
number of references would have been endless. A slight-
ly weak point is the lack of illustration of the diversity of
Lepidoptera: the systematic chapters are completely
without line drawings and some of the black and white
photos of adults are of insuficient quality, hardly giving
an impression of the enormous diversity. A few more
colourplates would have helped a lot.

It is tempting to compare this volume to two other
recent treatments of the order, both centered on the
Australian fauna: I. F. B. Common’s Moths of
Australia (1990) and Nielsen and Common’s chapter
on Lepidoptera in the new edition of “The insects of
Australia’. Undoubtedly both are far better illustrated
than the present volume, especially Common’s book
with its many colour plates, and in general the family
descriptions have more detail. On the other hand,
dealing particularly with the australian fauna, these

82

treatments miss some information on non-australian
families, and Scoble’s general chapters have much de-
tail not found in the other books. One could wonder if
it is wise to produce such similar treatments within this
short period, with the inevitable amount of overlap.

One final point is purely esthetical: I personally do
not like the rather coarse design of the cover (a black
zygaenid on red), nor the graphic design (typeset in
palatino with a slightly too narrow spacing).

In conclusion: a valuable overview of Lepidoptera,
especially recommended for general and applied en-
tomologists, teachers and students. The specialist can
find an easy entry into the literature of a specific fam-
ily or subject, in addition to the other recent treat-
ments of the group. The general chapters give the
most extensive recent review of Lepidoptera form and
function.

[E. J. van Nieukerken]

Fibiger, Michael, 1993. Noctuidae Europaeae. Volume 2,
Noctuinae II. — Entomological Press, Soro. 230 pp., 11
colour plates, textfigs, many maps. [ISBN 87-89439-02-
6]. Price DKK 680 excl postage; subscribers to volumes
1-12 receive 10% discount. Distributed by Apollo Books,
Kirkeby Sand 19, DK-7771 Stenstrup, Denmark. Fax +
4562263780.

This is the second volume of this series, which
started in 1990, and concludes the treatment of the
subfamily Noctuinae.

The series Noctuidae Europaeae will provide in 12
volumes a monograph which will enable an accurate
identification of all European species of Noctuidae
and supply information on bionomics and distribu-
tion. Each volume will treat one or more subfamilies,
each with their own authorships. The text is in eng-
lish and french, printed alongside.

In this volume 116 species are dealt with. For each
species a short diagnosis is given, together with notes
on bionomics, distribution and taxonomy. An area
map gives a general impression of the distribution.
The adults are superbly illustrated in the colour
plates, which were prepared by the well known pho-
tographer David Wilson. Usually both sexes, and
some variation is shown. It should be mentioned that
the genitalia of the species from the volumes | and 2
will be shown in volume 3.

Again, Fibiger has prepared a wonderful piece of
work: thorough texts, much new information, which is
all brought together in an attractive book. We should
admire an amateur with a full time job who produces
such a fine piece of scientific work. We must hope that
the author and his coworkers find time to continue at

least with the same speed: if volumes will come out

with the same interval of time as between volume one
and two, the whole series will be completed in 33 years!

[E. J. van Nieukerken]

LOUIS M. ROTH

Museum of Comparative Zoology, Harvard University, Cambridge, M. A., U. S. A.

REVISION OF THE COCKROACH GENUS
CTENONEURA HANITSCH (BLATTARIA,

POLYPHAGIDAE).

Roth, L. M.,

1993. Revision of the cockroach genus Ctenoneura Hanitsch (Blattaria,

Polyphagidae). — Tijdschrift voor Entomologie 136: 83-109, figs. 1-21. [iss 0040-7496].

Published 1 July 1993.

Most of the 15 known species of Ctenoneura are redescribed and 12 new taxa are described.
The genus is principally Malaysian, Indonesian, and Asian. A key is given to distinguish the

adults.

Dr. L. M. Roth, 81 Brush Hill Road, P. O. Box 540, Sherborn, MA 01770, U. S. A.
Key words. — Crenoneura; Blattaria, Polyphagidae; cockroaches; taxonomy; new species.

There are 15 described species of Ctenoneura
(Princis 1963: 101, 1971: 1138). I have seen the
types of ten. I have not examined Bey-Bienko’s spe-
cies but have included his references and diagnostic
features and included them in my key. Twelve new
species are described.

The following museums and their curators or col-
lection managers kindly loaned me specimens, or
house types which I have not examined:

EASC - Institute of Entomology, Academy of Science, China.

HDEO - Hope Department of Entomology, University
Museum, Oxford, England; Dr. George C. McGavin &
Mr. I. Lansbury.

MCZ - Museum of Comparative Zoology, Harvard
University, Cambridge, MA, U.S.A.

NRSS - Naturhistoriska Riksmuseet, Stockholm, Sweden;
Dr. P. Inge Persson.

RMNH - National Museum of Natural History
(Rijksmuseum van Natuurlijke Historie), Leiden, The
Netherlands; Mr. J. van Tol.

zits - Zoological Institute, Lund, Sweden; Dr. Roy
Danielsson.

ZINR - Institute of Zoology, Academy of Science, Russia.

SYSTEMATIC PART

Genus Ctenoneura Hanitsch

Ctenoneura Hanitsch, 1925: 100; Princis (1953) 1954: 207-
208 (key to species); 1963:101. — Type species:
Ctenoneura major Hanitsch, selected by Princis 1950:
204.

Etymology. — The name refers to the comb-like ar-
rangement of the cubitus branches on the hind wing
(Hanitsch 1925: 100).

Diagnosis. — Head, pronotum and tegmina not pu-
bescent. Tegmina and wings fully developed extend-
ing well beyond the end of the abdomen, the former
horny, venation clearly defined, discoidal sectors usu-
ally oblique (fig. 4C), rarely longitudinal or curved
anteriorly (figs. 1F, 2B). Hind wings completely over-
lapping each other; usually with (fig. 9c) an interca-
lary vein (rarely absent; fig. 1E) between the radial
and branched media rami; cubitus vein with three to
eight parallel, curved, comb-like branches (some of
which may be forked) (figs. 1E, 9C), the anal area
folded over the rest of the wing, but not fan-like.
Anteroventral margin of front femur with a row of
minute piliform spinules, terminating with a single
large spine (Type C,); pulvilli absent, tarsal claws
symmetrical, usually simple, rarely serrated, arolia, if
present, minute, fleshy, unsclerotized. Supraanal
plate of both sexes transverse, short. Male: Abdomi-
nal terga appear to be unspecialized, but many species
have a white, membranous area which could be a ter-
gal gland on the supraanal plate (slides of the termi-
nalia were not prepared). Subgenital plate usually
strongly asymmetrical with modified hind margin
(e.g., fig. 6B), rarely symmetrical, unmodified (fig.
1B); usually with only one style on the left side (e.g.,
fig. 5B), rarely with two (fig. 1B), or styles absent (fig.
14E). Female: subgenital plate valvular.

I did not study the male genitalia because for most
of the species there were very few duplicate specimens
to prepare slides. One of the most diagnostic charac-
ters is the shape of the male’s subgenital plate and this
structure is so convex that it is greatly distorted when
flattened on a slide. Polyphagid genitalia are much
more complex structures than those of Blattellidae
and Blaberidae, and possibly might not show specific

83

Ty DSCHRIET VOOR ENTOMOLOGIE, VOLUME 136, 1993

differences. I did not think it worth mutilating
unique specimens for this purpose.

Distribution checklist by species of Ctenoneura

aberrans Hanitsch: Mentawai; Sumatra ................ 86
Geiten er Bienko Chun 87
annulicornis Princis: Borneo ................................. 88
GUGLIE en 90
binmanıca Paineisy Burman 91
brunnea Hanitsch: Sumatra ................................. 92
Erassısbylaspan ARAN 107
VJ Anes hs aanva kn 94
geantedsprn Derale ven 87
PESA Primal IS Ua 96
kennen PE AVIRA RE 92
CONA DA AIRIS DANS ALAN EEE 104
luma sp -1).2y eral LI sees Anse PO ee 101
VGA SAW 96
niscaAbeBienko Ch Aree E 9D
mjoebergi Princis: Kalimantan; Sarawak … … … … … 97
mnd CISSÉ IS arava kn 106
DATASGULIGAIS PRE AS ab a 103
DOTLTLAIS PAM AE RE RE AE 100
Prop IBOATS D> NES aa PEER 89
seutiensprns Sabah AEN ee 102
CINA) BIEN ACI RR 107
SPES outen A EE O ARE RE en 107
DEA SOA GD, es SUN ATOS ER RO 98
rip CES ars p Nn RS AD AE BR EEE PRE 106
tuberculata Princis: Sumatra … nnn 97
DIODES J Oe See SA DATES TI TI 99
PTE PAD |X EINEA OE {Chi bee ene 94

Geographical distribution checklist of Ctenoneura
species

Borneo (Kalimantan; Sabah; Sarawak): annulicor-
nis, fulva, kinabaluana, major, mjoebergi, murudensis,
parascutica; poringa; propannulicornis scutica; tripro-
cessa; uncata

Burma: birmanica

China: acuticerca; misera; simulans, yunnanea

Java: kemneri

Malay Peninsula (Pahang; Perak): crassistyla, gigan-
tea; luma; spinastyla

Mentawai I.: aberrans

Sumatra: aberrans, biguttata; brunnea; hanitschi;
tuberculata

South Vietnam: sp.

Key to species of Ctenoneura

le Hind wing without a intercalary vein (figs.
IE, DO) wAroliavabsentsamen sets RP ne 2
= Hind wing with an intercalary vein (fig. 9C).

84

8(6)

Arolia minute or absent. Male subgenital
PIACE ASY MANCHI CAR E 4
Tarsal claws serrated. Cubitus vein of hind
wing with three branches (fig. 1E). (male &
female). Male subgenital plate symmetrical
with two similar styles (fig. 1B) … … aberrans
Tarsal claws simple. Cubitus vein of hind
wing with four to six branches (figs. 2C, 2D)

Pronotal disk reddish brown, lateral zones
darker brown, the edges yellow (fig. 2A). (sex
TRIO WIN) re N gigantea
Pronotal disk yellowish brown, lateral regions
yellow-hyaline. (male) ................... acuticerca
Subgenital plate with a setal tuft on the left
posterolateral corner (figs. 3A, 3C) ............ 5
Subgenital plate without a setal tuft on the
le porter COMENT sesame 6
Hind margin of subgenital plate with two U-
shaped excavations on the right side (fig.
3A). Antennae with first four segments
brown, succeeded by three whitish antenno-
meres, remaining segments brown. (male) ...
TICE PET ATES, MIE MES TU te bee? annulicornis
Hind margin of subgenital plate unmodified
on the right side (fig. 3C). Antennae dark
brown, distal end with three white antenno-
meres succeeded by four dark terminal seg-
ments. (male) ...................... propannulicornis
Subgenital plate hind margin deeply excised,
with a large, round swelling on the left side
Ges, WDA, NSD) scocsssnescecocosoaene 7
Subgenital plate hind margin excised or not,
without a round swelling on the left side ..8
Left style small, arising near the apex of the
left lobe of the subgenital plate, apexes of
both lobes not setose (fig. 12B). (male) … …
RNN ee ee leo tuberculata
Left style elongate, spine-like, arising near the
base of the left lobe of the subgenital plate,
apexes of both lobes setose (figs. 13A-D).
(Male) e ER ANA spinastyla
Pronotum shining black medially, dark red-
dish brown laterally, and with a pair of
orangish maculae on either side of the ante-
rior margin (fig. 4A). Right of subgenital
plate with a pair of curved, spine-like pro-

cesses directed towards the left (figs. 4B, 4E).

(MITE eaters biguttata
Pronotum and subgenital plate not as above.
pia PE es dini Bora Lu 9

Hind margin of subgenital plate not excavat-
ed, with an elongate process on the right
side; style, if present, not visible from below
OL the rear (Gies EMA) (a MEN

10(9).

11 (10).

ANDP

1312).

14 (13).

15 (14).

16 (15).

17 (16).

Hind margin of subgenital plate excavated,
left style, if present, visibile from below or
TE on 10
Left side of subgenital plate with a hook-
like structure, style absent (figs. 14C-14E).
Antennae with four brown basal segments
succeeded by yellowish antennomeres to
about the middle, those on the distal half
brown with some white terminal or preter-
minallones. (MAC)... uncata
Subgenital plate without a hook-like struc-
ture on the left side, style present. Antennae
MOMBASA bover me el. ital
Subgenital plate with a long, slender process
on the left side, its apex directed towards the
right, bearing a small cylindrical style near its
middle (figs. 11B-11D). (male) .....mjoebergi
Subgenital plate not as above
Hind margin of subgenital plate widely exca-
vated, with a colourless, membranous area
bearing a small, cylindrical style on its basal
end (figs. 15A-15C). (male)
Hind margin of subgenital plate not as above
13
Hind margin of the subgenital plate divided
principally into two or three, dissimilar
lobes, with a small cylindrical style arising on
the left lobe near its middle, or posterolateral
corner, or on the inner margin of the excava-
UOMO, th occ ome eRe n. 14

above
Hind margin of subgenital plate with a deep
V-shaped excavation, with a slender style on
the left side; a slender, thorn-like filament is
located on the dorsal surface of the plate,
protrudes beyond the apical margin of the
right lobe (figs. 15E-15G). Supraanal plate
inigpnali(fiea15D). (male) 222277... luma
Subgenital and supraanal plates not as above
ee Lal 15
Subgenital plate with two overlapping lobes
on the right side, the dorsal one upturned
and partially covering the right side of the su-
praanal plate, the ventral lobe with minute
spicules near its distal margin (figs. 5B, 5C).
(Malo) nnt. birmanica
Subgenital plate not as above
Left lobe of the subgenital plate much nar-
rower than the right one, the style originat-
ing on the posterolateral corner (fig. SE).

(male) tte ea 2 misera
Hind margin of subgenital plate not 25 above
i iedee ga 17

Hind margin of subgenital plate with a pair
of narrowly separated lobes directed posteri-

20 (19).

23(22).

24 (23).

Rorn: Revision of Ctenoneura

orly, a style originating near the middle of
the left one (figs. 8B-8D). Arolia absent.
mal a Se Re fulva

Hind margin of subgenital plate not as
above ANON RIONI CR RO 18

‚Style on the left lobe of the subgenital plate

originating along the inner margin of the ex-
cavation (fig. GB). (male) ............... brunnea
Style originating near the posterolateral cor-
nemoftheleftlobets. al. nnd... 19
Hind margin of the subgenital plate with a
small process that curves dorso-sinistrad (fig.
10B-10D). Pronotum subrectangular, the
disk with a sublozenge-shaped reddish
brown macula narrowly margined with yel-
low, broad lateral zones yellow-hyaline (fig.

IVA) (male) ee secs hanitschi
Subgenital plate and pronotum not as above
Ne En 20

Right lobe of subgenital plate narrow, elon-
gate, extending well beyond hind margin of
the shorter, wider left lobe (figs. ID, 9E).
Pronotum suborbicular, reddish brown (fig.
9A). Cubitus of hind wing with eight
branches (4e 20) na major
Right and left lobes of subgenital plate not as
above (figs. 7C, 7F) :
Pronotal edging yellow, disk reddish brown,
lateral regions brown-hyaline. Cerci brown.
(MAO ee er. ur kemneri
Pronotal disk brownish black, lateral regions
yellow. Cerci yellow. (male) ......... yunnanea
Subgenital plate trigonal, hind margin with
three slender processes that are curved dorsad,
and a long tapering filament that protrudes
between the left and middle processes; style ab-
sent (figs. 19A-19E). Cubitus vein of hind
wing with three branches (fig. 19F)
FO ee ae nk, lee rade à triprocessa
Subgenital plate not as above. Style present.
Cubitus vein of hind wing with more than
three branches. Style present
Subgenital plate with a broad plate whose
distal margin is excavated and upturned; left
style is located in a pale, membranous area
(figs. 20A-20E). Cubitus vein of hind wing
with eight branches. (male) ........ murudensis
Subgenital plate not as above. Cubitus vein
of hind wing with less than eight branches .
A ee ie eenden 24
Hind margin of subgenital plate deeply exca-
vated with a very large, stout style in the left
corner (figs. 21A-21C). (male) … crassistyla
Subgenital plate and left style not as above …

85

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

25 (24). Subgenital plate broadly excavated, with a
short, broad, process without close lobes on
either side, that extends dorsad, its distal end
curving anteriorly towards the supraanal plate
(figs. 18A-18D). (male) ............ kinabaluana

— Subgenital plate narrowly divided medially,
with a dark spine-like process that curves
dorsad between the two lobes ................. 26

26 (25). Margins of the excision smooth, practically
touching the spine-like process (figs. 16A-
KEEN male) hs. ER eee scutica

= Margins of the excision are irregular and on
the left side the region is unsclerotized (fig.
AB) E (MIO DE parascutica

REDESCRIPTIONS, AND DESCRIPTIONS OF NEW i
SPECIES

Ctenoneura aberrans Hanitsch
(fig. 1)

Ctenoneura aberrans Hanitsch, 1928: 37, pl. 2 figs. 8,9 (2);
1929a: 294, Bruijning 1948: 41, 148, fig. 54; Princis,
1963: 101.

Material examined. — Holotype, (2), Mentawelali,
Siberut, no. 91, 23.ix.1924, H. H. Karny; Type Orth.
342'/,, in HDEO. — Paratype. Mentawai. HDEO: same locality
and collection data as holotype, 1d (reported as ‘sex 2’), no.
80, 22.ix.1924, Type Orth. 342°/,. — SUMATRA. HDEO: Wai
Lima, 16, 12, 2 (abdomens missing).

Redescription. — Female: Head covered, interocu-
lar space distinctly wider than the distance between
antennal sockets. Pronotum with lateral and anterior
margins convex, hind margin straight (fig. 1A).
Tegmina and wings extending beyond end of abdo-
men, the former with a few well marked raised veins,
few almost longitudinal veins in discoidal sector (fig.
1F). Hind wings broad, mediastinal vein straight,
simple, radial vein slightly sinuous, costals not distin-
guishable; media vein, simple, cubitus vein with three
curved parallel branches; intercalary vein (between
the radial and media veins) absent (fig. 1E). Front fe-
mur Type C;; pulvilli and arolia absent, tarsal claws
symmetrical, distinctly serrated. Supraanal plate
trigonal, apex broadly rounded (fig. 1D). Subgenital
plate trigonal, valvular, completely hidden under the
supraanal plate.

E

1mm

F 1mm

Fig. 1. Ctenoneura aberrans Hanitsch. — A. female (holotype) pronotum; B, C. male (paratype) subgenital (ventral) and su-
praanal (dorsal) plates, respectively; D-F. female holotype: D. supraanal plate (dorsal); E, F. hind wing and tegmen respec-

tively (a = radius; b = media; c = cubitus).

86

Fig. 2. Ctenoneura
gigantea sp. n., hol-
otype. — A. prono-
tum; B. right teg-
men; C, D. left and
right wings.

Rorn: Revision of Ctenoneura

2mm

Male (previously undescribed): Differs from the fe-
male as follows: Supraanal plate transverse, very short,
shallowly trigonal, apex rounded (fig. 1C). Subgenital
plate essentially symmetrical, with two widely separ-
ated, similar styles, interstylar margin unmodified,
convexly rounded (fig. 1B).

Colour. — Head dark chestnut brown, shiny; an-
tennae relatively pale with yellowish tinge. Pronotum
reddish brown throughout (fig. 1A). Tegmina hyaline
amber, humeral and part of costal vein regions darker
(fig. 1F). Wings with a dark blotch in the costal vein
region (fig. 1E). Abdomen dark brown. Cerci brown.
Legs pale testaceous.

Measurements (mm) (2 in parentheses). — Length,
3.8-4.3 (3.6-4.0); pronotum length x width,, 1.1-1.3
x 1.5-1.8 (1.1-1.3 x 1.7-1.8); tegmen length, 4.0-4.5
(4.2-4.6); interocular space, 0.8 (0.8).

Comments. — This is the smallest species in the ge-
nus and is also unique in having a symmetrical sub-
genital plate with two styles, and serrated tarsal claws.

Ctenoneura acuticerca Bey-Bienko

Ctenoneura acuticerca Bey-Bienko, 1957: 896, 903, 912 (ty-
pe 2, Mt. Santaishan, Yunnan, China; in EASC) [not exa-
mined] (Russian, English summary); Princis 1963: 101.

Description (from Bey-Bienko). — Female:
Pronotum smooth. Tegmina with five oblique
branches of the radius which run into the median
third of the costal margin, anal veins not quite dis-
tinct. Radius of hind wings with three slightly pro-
nounced oblique branches, intercalary vein absent

between radius ‘and media, cubitus divided into six
branches. Arolia absent. Supraanal plate large, trian-
gular, apex emarginate. Subgenital plate with valvular
part lozenge shaped. Apex of cerci with a large spine.

Colour. — Brownish yellow, pronotal disk yellow-
ish brown, lateral parts yellow, transparant. Wings
with distal part yellowish. Legs yellow. Cerci brown-
ish yellow.

Male: Unknown.

Measurements (mm). — Body length, 8.5-8.7; pro-
notum length, 2.7; tegmen length, 10.2-10.5.

Comments. — According to Bey-Bienko, this spe-
cies is related to aberrans (probably because of the ab-
sence of an intercalary vein, and arolia).

Ctenoneura gigantea Sp. n.

(fig. 2)

Type material. — Holotype, sex unknown (abdomen miss-
ing), Gunung Kledang, Perak, 2646’, xi.1916 (with a
handwritten label, C. pauciramosa..., ms. name); in
RMNH.

Description. — Sex unknown: Head exposed, inter-
ocular space greater than the distance between ocellar
spots and antennal sockets. Pronotum parabolic (fig.
2A). Tegmina and wings fully developed, the former
with raised veins, and longitudinal discoidal sectors
(fig. 2B). Hind wing with about three poorly defined
costal veins, intercalary vein absent, media vein with
one branch, cubitus vein with four branches (one
forked near the base on the left wing), and six branch-
es (two of them joined distad on the right wing) (figs.

87

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 3. Ctenoneura spp. — A-B, C. annulicornis Princis, male holotype, A. subgenital plate (ventral); B. subgenital plate (rear
view); C-E. C. propannulicornis sp. n., male holotype: C. subgenital plate (ventral); D. supraanal and subgenital plates (rear
view); E. supraanal and subgenital plates (dorsal; the distal region of the supraanal plate is deflexed and the margin is distort-

ed)

2C, D). Front femur Type C;; pulvilli absent, tarsal
claws very long, simple symmetrical, arolia absent.
Abdomen missing.

Colour. Head reddish brown, the region from the
occiput to ocellar spots darker; antennae blackish.
Pronotal disk reddish brown, lateral zones darker
brown, lateral edges yellowish (fig. 2A).

Measurements (mm). — Length, ?; pronotum
length x width, 2.9 x 3.7; tegmen length, 10.4; inter-
ocular space, 1.3.

Etymology. — The specific name refers to the large
size of the species.

Comments. — I am describing this species even
though its abdomen is missing, because its size is so
much greater than any of the known species. Also the
parabolic shape of the pronotum, the absence of an
intercalary vein on the hind wing (known only in
aberrans and acuticerca), and the complete absence of
arolia, are good distinctive characters when coupled
with its size.

88

Ctenoneura annulicornis Princis

(figs. 3A-B)

Ctenoneura annulicornis Princis, (1953) 1954; 206, fig. 3.
Holotype, d, Borneo, Long Navang, Mjöberg (NRSS)
[examined]; Princis 1963: 102.

Ctenoneura fulva (nec Hanitsch, 1925); Hanitsch 1933: 235
[in part; only the Long Navang d |; Princis 1963:102.

Redescription. — Male: Head hidden, interocular
space greater than the distance between ocellar spots
and antennal sockets. Pronotum suboval. Tegmina
and wings extending well beyond end of abdomen.
Tegminal veins not raised. Hind wing with interca-
lary vein, media vein with one forked branch, cubitus
with six branches. Front femur Type C;; pulvilli and
arolia absent, tarsal claws symmetrical, simple.

- Abdominal terga unspecialized. Supraanal plate

transverse, narrow, symmetrical. Subgenital plate
asymmetrical with a setal tuft on the left posterolater-
al corner, and with three sclerotized structures and a

Fig. 4. Ctenoneura
biguttata Hanitsch,
male holotype. — A.
pronotum; B. su-
praanal and subgeni-
tal plates (rear view);
C, D. left tegmen
and hind wing; E.
supraanal and sub-
genital plates (dor-
sal).

Rove: Revision of Ctenoneura

well developed cylindrical style, also on the left side,
the right side with a pair of U-shaped excavations
(figs. 3A, B).

Colour. — Head dark reddish brown; antennae
with first four segments brown, succeeded by three
whitish antennomeres, remaining segments brown.
Pronotal disk dark reddish brown, lateral zones hya-
line. Tegmina hyaline light brown. Adomen brown,
broad lateral zones on the sterna pale. Legs light
brown.

Female: Unknown.

Measurements (mm). — Length, 5.7; pronotum
length x width, 1.7 x 2.2; tegmen length, 6.3; interoc-
ular space, 0.6.

Ctenoneura propannulicornis sp. n.

(figs. 3C-E)

Type material. — Holotype, d, Malaysia, Sabah, Ulu
Rurun, 115°40’30”E 4°22’N, 1500 m, 21-22.xii.1986, J.
Huisman (RMNH). — Paratype. Sabah, same locality and col-
lector as the holotype, ML, 1d, 20.xii.1986 (RMNH).

Description. — Male: Head hidden, interocular
space greater than the distance between the antennal
sockets. Pronotum subelliptical. Tegmina and wings

05 mm

fully developed extending well beyond the end of the
abdomen, discoidal sectors of the former oblique.
Hind wing with distinct costal veins, the proximal
ones thickened distad, intercalary vein present, media
vein with one branch, cubitus vein with five branches
(one of them may be forked). Front femur Type C;;
pulvilli absent, tarsal claws simple, symmetrical, aro-
lia minute. Abdominal terga unspecialized. Supraanal
plate transverse, hind margin broadly, shallowly exca-
vated, with a large, dense group of curved setae on the
left corner, style absent, right side of plate unmodi-
fied (figs. 3C-E).

Colour. — Head dark brown; antennae dark brown,
the distal end with three white antennomeres, suc-
ceeded by four dark terminal segments. Pronotal disk
dark brown, the lateral regions infuscated-hyaline.
Tegmina dark brown-hyaline, humeral region yel-
lowish. Hind wing infuscated, darkest in the costal
vein and apical regions, lightest in the mid portion.
Abdominal terga brown, lateral edges darker, supraa-
nal plate with a white macula anteromedially.
Abdominal sterna light brown, subgenital plate dark-
er. Cerci brown, apical segment lighter. Coxae, femo-
ra, and tibiae brown, tarsi lighter.

Female: Unknown.

Measurements (mm). — Length, 5.8-6.3; prono-

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 5. Ctenoneura spp. — A-D, C. birmanica Princis, males: A. pronotum (holotype); B-D. paratype: B. subgenital plate
(ventral); C, supraanal and subgenital plates (dorsal; the dotted line indicates the margin of the supraanal plate that is hid-
den by one of the lobes of the subgenital plate); D. hind wing. — E, C. misera Bey-Bienko, male holotype, subgenital plate

(ventral) (redrawn from Bey-Bienko, 1970).

tum length x width, 1.9 x 2.3-2.4; tegmen length,
7.1-8.3; interocular space, 0.6.

Comments. — I place this species near annulicornis
(hence the specific name), because both taxa have a
dense tuft of setae in the left corner of the subgenital
plate. Differences in the shape of the right side of the
plate, absence of the left style (in propannulicornis),
and antennal colouration, separate the two species.

Ctenoneura biguttata Hanitsch

(fig. 4)

Ctenoneura biguttata Hanitsch, 1932a: 6. Holotype, d,
Sumatra, Deli, Fulmek, 1921-26, Brastagi, Urwald,
vili.1924, R. Ebner; Type Orth. 375 (HDEO), [exami-
ned]; Bruijning 1948: 41, 149; Princis (1953) 1954: 207
(footnote), 1963: 103.

Redescription. — Male: Head exposed, eyes wide

90

apart, interocular space greater than distance between
antennal sockets. Pronotum subcircular (fig. 4A).
Tegmina and wings extending beyond end of abdo-
men, veins and cross veins of both distinct, discoidal
sectors of the former oblique (fig. 4C). Wing with six
costals, the first four with distal ends merging into a
marginal dark patch; intercalaray vein present
between the radial and median; media vein branched,
cubitus with five branches (fig. 4D). Front femur
Type C,; pulvilli and arolia absent, tarsal claws sym-
metrical, simple (not serrated). Abdominal terga un-
specialized. Supraanal plate transverse, very narrow,
hind margin entire (fig. 4E). Subgenital plate asym-
metrical, with a small cylindrical style on the left side,
(Hanitsch stated, ‘styles not observed’), right side
with a pair of curved spinelike processes directed to-
wards the left (figs. 4B, E).

Colour. — Head shining black; antennae pale yel-

lowish with darker terminal segments. Pronotum
shiny black medially, dark reddish brown laterally,
and with a pair of orangish maculae on either side of
the anterior margin (fig. 4A). Tegmina uniformly
dark brown (fig. 4C). Wings hyaline, brown, with a
dark patch along the anterior costal region (fig. 4D).
Abdominal terga light brown, sterna dark brown,
subgenital plate with an orangish area distad. Legs
with coxae and femora dark brown, tibiae and tarsi
yellowish. Cerci with brown basal segments, remain-
ing cercomeres yellowish.

Female: Unknown.

Measurements (mm). — Length, 5.7; pronotum
length x width, 1.7 x 2.2; tegmen length, 7.3; interoc-
ular space, 0.8.

Ctenoneura birmanica Princis

(figs. SA-D)

Ctenoneura birmanica Princis, (1953) 1954: 206, fig. 5.
Holotype, d, Kambaiti, N. E. Burma, 7000 ft,

Rovu: Revision of Ctenoneura

25.v.1934, R. Malaise (NRSS) [examined]; Princis 1963:
102.

Ctenoneura major (nec Hanitsch, 1925); Princis 1950: 203,
204 (8), 1963: 102.

Material examined. — Holotype, and paratypes. Burma.
NRSS: 1 d, same locality and collector as holotype, 2000 m,
28.v.1934. ZIS: 1d, same data as holotype; 1 d , same local-
ity and collector as holotype. 2000 m, 29.v.1934.

Redescription. — Male: Head hidden or slightly ex-
posed; interocular space greater than distance
between ocellar spots and antennal sockets.
Pronotum subelliptical (fig. 5A). Tegmina and wings
fully developed extending well beyond end of abdo-
men, discoidal sectors of former oblique. Hind wing
with the distal ends of the proximal costal veins reach-
ing only to the dense spot near the costal margin;
intercalary vein present, media vein with two branch-
es (one forked), cubitus vein with six branches (one

forked) (fig. 5D). Front femur Type C;; pulvilli ab-

Fig. 6. Crenoneura brunnea Hanitsch, male holotype. — A. pronotum; B. subgenital plate (ventral); C, D. left tegmen and

hind wing; E. supraanal and subgenital plates (dorsal).

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

sent, tarsal claws symmetrical, simple, arolia minute.
Supraanal plate transverse, hind margin convexly
rounded, symmetrical, entire (fig. 5C). Subgenital
plate asymmetrical, divided by a medial excavation,
one lobe on the left side bearing a cylindrical style,
two overlapping lobes on the right side, the dorsal
one upturned (in the holotype only it partly covers
the right side of the supraanal plate), the ventral lobe
with minute spicules near its distal margin (figs. 5B,
©)

Colour. — Antennae dark brown with pale junc-
tions between individual segments. Head reddish
brown. Pronotal disk reddish brown, lateral areas
dark-hyaline (fig. 5A). Tegmina brownish, hyaline.
Hind wings weakly infuscated. Abdomen dark
brown, supraanal plate black. Legs brown. Cerci
brown, proximal cercomeres darker on both surfaces.

Female: Unknown.

Measurements (mm). — Body length, 7.7-8.1; pro-
notum length x width, 2.1-2.2 x 3.0; tegmen length,
10.0-10.4; interocular space, 0.8-0.9.

Ctenoneura misera Bey-Bienko

(fig. SE)

Ctenoneura misera Bey-Bienko, 1969: 832, fig. 1 (Russian).
Holotype d, S. China, Yunnan, Tapa-shan Mountains,
near Pin P’ien, 1300 m, 23.vi.1956, Huen K’e-yen; in
ZINR. [not examined]; 1970: 528 (English translation).

Description (after Bey-Bienko). — Male: Small.
Head with flat vertex, almost pendent (overhanging),
forming an obtuse angle with the frons; interocular
space slightly more than double the distance between
antennal sockets. Pronotum one and one half times as
wide as long, with broad transparent lateral parts,
hind margin with a narrower hyaline border. Hind
wings with oblique branches of the radius distinct,
vein between radius and media arched backwards;
cubitus vein with seven branches. Subgenital plate as
in fig. 5E [style in the left posterior corner]. Cerci
long, thickened towards the bases, terminal segment
not spine-like.

Colour. — Rusty yellow. Head chestnut brown, an-
tennae brownish yellow. Hind wing just visibly infus-
cate, yellowish along costal margin.

Female: Unknown.

Measurements (mm). — Body length, 6.8; prono-
tum length x width, 2.0 x 3.0; tegmen length, 8.0;
overall length, 10.0.

Comments. — According to Bey-Bienko, this spe-
cies is similar to yunnanea from China and kemneri
from Java, but clearly distinguished by the colour of
head and pronotum, small body, and structural de-
tails of the subgenital plate. The subgenital plate (fig.
SE) of misera more closely resembles that of birmani-

92

ca (fig. 5B) than it does yunnanea (fig. 7F) or kemne-
ri (fig. 7C).

Ctenoneura brunnea Hanitsch

(fig. 6)

Ctenoneura brunnea Hanitsch, 1929a: 266, 292, fig. 5;
1932b: 52, 81. Holotype, d [Hanitsch did not indicate
the sex], Gunung Singgalong (Sumatra, Westkust), 1800
m, 1925, E. Jacobson; Type Orth. 362, in HDEO [exam-
ined]; Bruijning 1948: 41, 148: Princis (1953) 1954:
208, fig. 8 (d) Princis 1963: 102.

Additional material. — Sumatra. RMNH: N. Sumatra,
Bivouac One, Mt. Bandahara, 3°43’N 97°41’E, ca. 810 m,
1 6, 25.vi. —5.vii.1972, J. Krikken.

Redescription. — Male (Hanitsch did not deter-
mine the sex of the type): Head covered, interocular
space less than distance between antennal sockets.
Pronotum subcircular (fig. GA). Tegmina and wings
fully developed extending beyond end of abdomen,
veins of the former distinct, not raised, discoidal sec-
tors oblique (fig. 6C). Hind wing with a forked media
vein, intercalary vein present, cubitus with six
branches (fig. 6D). Front femur Type C,; pulvilli ab-
sent, tarsal claws symmetrical, simple, arolia subobso-
lete. Abdominal terga unspecialized. Supraanal plate
strongly transverse, shallowly trigonal (fig. GE).
Subgenital plate asymmetrical, hind margin deeply,
widely excavated, with a small cylindrical style on the
left side along the inner margin of the excavation (fig.
6B).

Colour. — Head shiny dark reddish brown; anten-
nae dark brownish black. Pronotal disk dark reddish
brown, lateral borders broad, hyaline, partially yel-
lowish (fig. 6A). Tegmina shining brown. Wings
brownish, darker along the distal half of the costal
margin and apically (fig. 6D). Abdomen yellowish to
reddish brown, lateral borders darker. Cerci dark
brown to black. Coxae, femora, and tibiae reddish
brown, tarsi yellowish.

Female: Unknown.

Measurements (mm). — Length, 5.6-6.5; prono-
tum length x width, 1.6-1.8 x 2.1-2.3; tegmen length,
7.1-8.5.; interocular space, 0.5-0.7.

Ctenoneura kemneri Princis
(figs. 7A-E)
Ctenoneura kemneri Princis, 1967: 146, fig. 1. Holotype d,

Tjitjoroeng, Java, 23.1.1921, N. A. Kemner (ZILS) [exam-
ined] ; Princis 1971: 1138.

Redescription. — Male: Head hidden, interocular
space wrinkled, slightly less than the distance between
the antennal sockets. Pronotum strongly transverse,

Fig. 7. Ctenoneura spp. A-E.
— C. kemneri Princis, male
holotype: A. pronotum; B.
supraanal and subgenital
plates (dorsal); C. subgenital
plate (ventral); D. subgenital
plate (rear view); E. subgeni-
tal plate (left lateral). — F. C.
yunnanea Bey-Bienko, male
holotype, subgenital plate
(ventral) (redrawn from Bey-
Bienko, 1957).

its greatest width before the middle, anterior margin
cucullate above the head (fig. 7A). Tegmina extend-
ing well beyond end of abdomen, discoidal sectors of
the former oblique. Hind wing with distinct costal
veins, not thickened distad; intercalary vein present,
media vein with one branch, cubitus vein with six or
seven branches (one of the branches may be forked).
Front femur Type C,, terminal spine very small; pul-
villi absent, tarsal claws symmetrical, simple, arolia
minute. (Princis claimed that only the fourth segment
of the basitarsus had a pulvillus. However this ‘pulvil-
lus’ is spined and is no different from the pulvillar re-
gion on the other tarsomeres. Pulvilli normally are
fleshy and lack spines and I consider the basitarsus in
this species as lacking a pulvillus). Abdominal terga
unspecialized; supraanal plate short, transverse, hind
margin convexly rounded (fig. 7B). Subgenital plate
divided into two dissimilar lobes, the left one bearing
a cyclindrical style in the posterolateral corner (figs.
7B-D).

Colour. — Head dark brown with lighter mouth-
parts; antennae brown. Pronotum with yellow edging
(wider laterally than on the anterior and posterior

Rorn: Revision of Ctenoneura

margins, disk bright red brown, the lateral parts hya-
line brown. Tegmina uniformly reddish brown, veins
yellowish. Hind wings darkly infuscated, part of co-
stal vein area with a yellowish patch. Abdomen
brown. Cerci brown, darker ventrally. Coxae, femora,
and tibiae reddish brown, tarsi yellowish.

Female: Unknown. _

Measuremnts (mm). Body length, 6.5; pronotum
length x width, 2.0 x 3.2; tegmen length, 8.2; interoc-
ular space, 0.6.

Comments. — Princis stated that kemneri to some
degree resembled major, chiefly because its subgenital
plate is of the same type. However their lobes differ in
shape. Actually Princis’s drawing of the subgenital
plate of kemneri (Princis, 1967: fig. 1) resembles that
of yunnanea (fig. 7F), more closely than it does that of
major (fig. IE). My camera lucida drawing of the sub-
genital plate of kemneri differs from Princis’s but
slight differences in the angle from which the speci-
men is drawn can make a marked difference in the
shape of the structure. The shapes of the subgenital
plate lobes of kemneri (fig. 7C) show some resem-
blance to those of birmanica (fig. 5B).

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

1mm

0.5 mm

0.5 mm

Fig. 8. Ctenoneura fulva Hanitsch, male holotype. — A. pronotum; B. subgenital plate (ventral); C. supraanal and subgenital
plates (rear view); D. supraanal and subgenital plates (dorsal); E, F. left tegmen and hind wing.

Ctenoneura yunnanea Bey-Bienko

(fig. 7F)

Ctenoneura yunnanea Bey-Bienko, 1957: 896, 902, 912, fig.
1. Holotype d, Mt. Santaishan, Yunnan, China (EASC)

[not examined] (Russian, English summary); Princis

1963:102.

Description (from Bey-Bienko). — Male: Prono-
tum transverse. Tegmina and wings with typical ve-
nation; wings with distinct oblique branches of the
radius, intercalary vein between radius and media
present, cubitus vein with seven branches. Supraanal
plate very short. Subgenital plate with a deep round
emargination, right margin thickened, style long,
thin, placed on the upper left margin of the plate (fig.
7F). Cerci moniliform.

Colour. — Brownish yellow. Vertex distinctly bor-
dered above antennal sockets. Pronotal disk brownish
black, lateral parts yellow. Cerci yellow.

94

Female: Unknown.

Measurements (mm). — Body length, 7.0; prono-
tum length, 2.0; tegmen length, 8.5.

Comments. — Bey-Bienko stated that this species
strongly resembles hanitschi. However, its subgenital
plate is similar to that of brunnea and kemneri.

Ctenoneura fulva Hanitsch
(fig. 8)

Ctenoneura fulva Hanitsch, 1925: 78, 101, figs. 13-14 (d &
2). Lectotype (here designated) d , Mt. Murud, Sarawak,
6000 ft., Dr. E. Mjöberg, Sarawak Mus. dd 1925; Type
Orth. 300'/,, in HDEO [examined]. [Hanitsch listed six
specimens in his original description, three d d from Mt.
Murud, and one 6 and two 2 ® from Mt. Dulit; there
are only two paralectotypes in the HDEO, and neither one
is fulva; see comments, below]; Princis 1933: 303, 329;
Bruijning 1948: 41, 148; Princis (1953) 1954: 207, fig. 6
(8); Princis 1963: 101.

Redescription. — Male: Head covered by prono-
tum, interocular space greater than distance between
antennal sockets. Pronotum subelliptical (fig. 8A).
Tegmina and wings extending well beyond end of
body, the former with proximal costal veins arising
from a common trunk, the distal ones singly, discoid-
al sectors oblique (fig. 8E). Hind wing with interca-
lary vein, media vein with two branches, cubitus with
six simple branches (fig. 8F). Front femur Type C;
pulvilli and arolia absent, tarsal claws symmetrical,
simple. Abdominal terga unspecialized. Supraanal
plate short transverse, hind margin shallowly convex
(figs. 8C, D). Subgenital plate asymmetrical, hind
margin with a pair of weakly dissimilar lobes, and a
small cylindrical style on the left side adpressed to the
surface, its apex directed anteriorly. The style is hid-
den in ventral view but can be seen in end view. The
tips of two apically acute structures, probably genital
phallomeres, protrude laterally, and dorsally from be-
neath the supraanal plate (figs. B-D).

Colour. — Head dark reddish brown, shiny, anten-

Rotu: Revision of Ctenoneura

nae dark brown. Pronotal disk reddish brown, sur-
rounding area broad, hyaline (fig. 8A). Tegmina
brownish yellow, humeral and costal area lighter (fig.
8E). Wings pale hyaline with a yellowish tinge, ante-
rior portion with a dark area along the anterior mar-
gin where the ends of the proximal costals merge (fig.
8F). Abdominal terga brown, terminal segments
darker, supraanal plate with a pale medial zone.
Abdominal sterna light brown, margins darker. Legs
brown.

Female: Not seen.

Measurements (mm). — Length, 6.6; pronotum
length x width, 1.9 x 2.7; tegmen length, 9.5; interoc-
ular space, 0.7.

Comments. — Two of Hanitsch’s paralectotypes are
not fulva. One from Mt. Murud (Type Orth. 300'/,)
is a new species, murudensis, described below. The
other paralectotype, from Mt. Dulit (Type Orth.
300°/,), is possibly new but I am not describing it be-
cause its subgenital plate is distorted and it is difficult
to see the shape of the hind margin.

Fig. 9. Ctenoneura major Hanitsch, male holotype. A. pronotum; B, C. left tegmen and hind wing; a = intercalary vein. D.
supraanal and subgenital plates (dorsal); E. subgenital plate (ventral); F. subgenital plate (right lateral).

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Ty DSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Ctenoneura major Hanitsch

(fig. 9)

Ctenoneura major Hanitsch, 1925: 78, 102 (d & 9).
Holotype d, Mt. Murud, Sarawak, 6500 ft., x.1922-
i.1923, Dr. E. Mjôberg, Sarawak Mus., dd 1925; Type
Orth. 301, in HDEO [Hanitsch listed a d and 2 from Mt.
Murud, but according to the number 301, there is only
one type specimen in HDEO] [examined]; Hanitsch 1927:
9, 26, 42 (in part?); Bruijning 1948: 41, 148; Princis
(1953) 1954: 207, fig. 7 (8); Princis 1963: 101.

Redescription (Hanitsch described both d & 9 in
the same paragraph. His description can not be used
to distinguish between the sexes). — Male: Head
slightly exposed, interocular space less than the dis-
tance between ocellar spots and antennal sockets.
Pronotum suborbicular (fig. 9A). Tegmina and wings
extending beyond end of abdomen, the former with
distinct, but not thickened veins, discoidal sectors
oblique (fig. 9B). Hind wing with eight costals, the
distal ends of the first three merging into a thicken-
ing, intercalary vein present, media with two branch-
es, cubitus vein with eight rami (fig. 9C). Front femur
Type C;; pulvilli absent, tarsal claws simple, symmet-
rical, arolia minute. Abdominal terga unspecialized.
Supraanal plate transverse, very narrow, hind margin
convex, entire (fig. 9D). Subgenital plate asymmetri-
cal, a small cylindrical style on the left side near the
posterior corner, right side produced, the process di-
rected posteriorly (figs. 9D-F).

Colour. — Head reddish and dark brown, shining,
ocelli pale, antennae dark brown. Pronotal disk dark

reddish brown (fig. 9A). Tegmina brown (fig. 9B).
Wings brown, thickened costal vein region darker
(fig. IC). Abdomen brownish yellow, hind margin of
subgenital plate in part yellowish. Cerci fuscous
brown. Legs dull brownish yellow.
Female. Not seen.

Measurements (mm). — Length, 9.7; pronotum
length x width, 2.2 x 3.0; tegmen length, 11.1; inter-
ocular space, 0.7.

Ctenoneura hanitschi Princis

(fig. 10)

Ctenoneura hanitschi Princis, (1953) 1954: 204, fig. 1 (6d.
Holotype, d, Sibajak, Sumatra, 1600 m, Mjöberg;
(NRSS) [examined]; 1963: 102.

Ctenoneura major (nec Hanitsch, 1925); Hanitsch 1929b: 3,
18 (in part, d).

Redescription. — Male: Head hidden, interocular
space greater than the distance between antennal
sockets. Pronotum subrectangular (fig. 10A).
Tegmina and wings fully developed extending well
beyond end of abdomen, the former with oblique dis-
coidal veins. Hind wing with an intercalary vein.
Front femur Type C,; pulvilli absent, tarsal claws
symmetrical, simple, arolia minute. Abdominal terga
unspecialized. Supraanal plate transverse, very short,
hind margin shallowly convex, entire (fig. 10C).
Subgenital plate asymmetrical, with a cylindrical style
on the left side; the right side has a process that is

Fig. 10. Ctenoneura hanitschi Princis, male holotype. — A. pronotum; B. subgenital plate (ventral); C. supraanal and subgen-
ital plates (dorsal); D. supraanal and subgenital plates (rear view).

96

A | ne

RotH: Revision of Ctenoneura

TOR Ne

Fig. 11. Crenoneura mjoebergi Princis, male holotype. — A. pronotum; B. supraanal and subgenital plates (rear view); C. su-
praanal and subgenital plates (dorsal); D. subgenital plate (ventral).

curved dorsad towards the left (figs. B-D).

Colour. — Head dark reddish brown; antennae
dark brown, monochromatic. Pronotal disk with a
sublozenge-shaped reddish brown macula narrowly
margined with yellow, broad lateral zones yellowish-
hyaline (fig. 10A). Tegmina brownish. Hind wing
weakly infuscated. Abdomen brown. Legs dark to yel-
lowish brown.

Female: Unknown.

Measurements (mm). — Body length, 7.3; prono-
tum length x width, 2.2 x 2.5; tegmen length, 9.4;
interocular space, 0.6.

Ctenoneura mjoebergi Princis

(fig. 11)

Ctenoneura mjoebergi Princis, (1953) 1954: 206, fig. 4 (d).
Holotype, d, O. Borneo, Pajau River, [Kalimantan],
Mjöberg; in (NRSS) [examined].

Ctenoneura fulva (nec Hanitsch, 1925). — Hanitsch 1933:
235 (in part; Mt. Tibang and Pajau River material);
Princis 1963: 102.

Material examined. — Holotype, and paratype:
Kalimantan. 1d, same data as holotype (NRSS). Sarawak.
Mt. Tibang, 1400 m, 1 d, Mjöberg (NRSS).

Redescription. — Male: Head concealed, interocu-
lar space greater than distance between antennal sock-
ets. Pronotum subelliptical (fig. 11A). Tegmina and

wings fully developed extending well beyond end of
abdomen, the former with oblique discoidal sectors.
Hind wing with an intercalary vein, media vein with
a forked branch, cubitus with five branches (one
forked). Front femur Type C,; pulvilli absent, tarsal
claws simple, symmetrical, arolia minute. Supraanal
plate symmetrical, transverse, shallowly trigonal, hind
margin entire (fig. 11C). Subgenital plate asymmetri-
cal with several processes, one of which on the left
side is elongated and bears a single style (figs. 11B-D).

Colour. Head reddish brown; antennae with seg-
ments on proximal half yellow (except for the first
two brown antennomeres), distal half brownish.
Pronotal disk reddish brown with a pair of dark spots
on anterior margin, lateral zones hyaline (fig. 114A).
Tegmina brownish, subcostal vein yellow. Hind wing
lightly infuscated. Abdomen brown, the sterna light-
er medially and laterally. Legs and cerci brown.

Female: Unknown.

Measurements (mm). — Length, 5.5-5.8; prono-
tum length x width, 1.5-1.7 x 2.1-2.3; tegmen length,
6.6-6.8; interocular space, 0.6.

Ctenoneura tuberculata Princis

(fig. 12)

Ctenoneura tuberculata Princis, (1953) 1954: 205, fig. 2
(3). Holotype, d, Sumatra, Sibajak, Mjoeberg (NRSS)
[examined]; Princis 1963: 101.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

C _ 05mm

B 0.5 mm

Fig. 12. Ctenoneura tuberculata Princis, male from Mt. Bandahara. A. subgenital plate (left lateral); B. subgenital plate (ven-

tral); C. subgenital plate (right lateral).

Ctenoneura major (nec Hanitsch, 1925); Hanitsch 1929b: 3,
18 (à, in part); Princis 1963: 101.

Additional material. — Sumatra. N. Sumatra, Bivouac
One, Mt. Bandahara, 3°43’N 97°41’E, ca. 810 m, 1d,
25.vi. —5.vii.1972, J. Krikken (RMNH).

Redescription. — Male: Head hidden, interocular
space greater than the distance between antennal
sockets. Pronotum subelliptical. Tegmina and wings
fully developed extending beyond end of abdomen,
the former with oblique discoidal sectors. Hind wing
with intercalary vein, media vein with one branch,
cubitus vein with six branches. Front femur Type C;
pulvilli absent, tarsal claws symmetrical, simple, aro-
lia minute. Abdominal terga unspecialized. Supraanal
plate shallowly trigonal, apex rounded and medially
incised. Subgenital plate strongly asymmetrical, hind
margin deeply divided forming two asymmetrical
lobes, the one on the left bearing a small cylindrical
style below its apex on the inner margin of the exci-
sion; the lobe on the right deeply excised; a large
round swelling is located along the margin distad on
the left side (figs. 12A-C).

Colour. — Head reddish brown; antennae dark
brown. Pronotal disk reddish brown the surrounding
area hyaline and partly yellowish. Tegmina with hu-
meral and costal vein areas yellowish, remainder
brown. Hind wing with yellowish tinge. Abdomen
brown, subgenital plate reddish brown (holotype), or
partly yellowish. Legs light brown. Cerci light brown
dorsally, darker brown ventrally.

Female: Unknown. |

Measurements (mm). — Body length, 6.0; prono-
tum length x width, 1.7-1.8 x 2.3-2.5; tegmen length,
7.2-9.0; interocular space, 0.5-0.6.

98

Ctenoneura spinastyla sp. n.

(fig. 13)

Type material. — Holotype, d, Pahang, F.M.S.,
‘Cameron’s Highlands’, No. 4 Camp, 4800 ft, 14.-
viii.1923, H.M., Pendlebury (incorrectly labelled by

Hanitsch, Ctenoneura major Hanitsch); in HDEO.

Description. — Male: Head hidden, interocular
space greater than the distance between ocellar spots
and antennal sockets. Pronotum suboval. Tegmina
and wings extending well beyond end of abdomen,
discoidal sectors of the former oblique. Hind wing
with distinct costal veins, intercalary vein present,
media vein forked beyond the middle, each branch
again dividing (left wing, fig. 13E), or with two
branches (one of them forked, right wing); cubitus
vein with five or six branches (two of them branching
once or twice) (fig. 13E). Front femur Type C,; pul-
villi absent, tarsal claws symmetrical, simple, arolia
minute. Abdominal terga unspecialized. Supraanal
plate strongly transverse, very short, hind margin al-
most truncate (fig. 13A). Subgenital plate asymmetri-
cal with a pair of large dissimilar processes, the left
one with a bulbous swelling near the apex, both lobes
setose apically, those on the right much longer; on the
left margin near the base of the plate is a long, slender,
spine-like style (figs. 13A-D).

Colour. — Head reddish brown; antennal segments
black (most of the antennae are missing). Pronotal
disk reddish brown, becoming yellowish in the poste-
rior region, lateral zones yellowish-hyaline. Tegmina
with humeral zone yellowish, remainder brownish-
hyaline, mediastine vein very dark. Abdomen, cerci,
and legs brown.

Female: Unknown.

Measurements (mm). — Length, 8.2; pronotum

Fig. 13. Ctenoneura spinastyla
sp. n., male holotype. — A. su-
praanal and subgenital plates
(dorsal); B. subgenital plate
(right lateral); C. subgenital
plate (ventral); D. subgenital
plate (left lateral); E. left hind

wing.

length x width, 2.3 x 2.9; tegmen length, 11.1; inter-
ocular space, 0.8.

Etymology. — The specific name refers to the slen-
der, greatly elongated left style.

Comments. — The two setose processes, and the
spine-like style on the subgenital plate are unique and
readily identifies this species.

Ctenoneura uncata sp. n.

(fig. 14)

Type material. — Holotype, 4, Malaysia, S.W. Sabah, nr.
Long Pa Sia (East), c. 1000 m, Mal. trap 5, 1-13.iv.1987, C.
v. Achterberg; in RMNH . — Paratypes: Sabah. RMNH: The
following were collected by C. v. Achterberg: S.W. Sabah,
nr. Long Pa Sia (West), c. 1050 m, Mal. trap 3, 1 d, 1-
14.iv.1987; same data as holotype, 1 d, 25.xi. —7.xii.1987;
nr. Long Pa Sia (West), 1010 m, Mal. trap 1b, 1 d, Mal.
trap 2, 1020 m, 2 dd, 25.xi. -8.xii.1987. The following
were collected by J. Huisman: Sg. Malabit, Rd. Long Sia-L
Semado, 115°41’E 4°21’N, 1175 m, 1 d, 8.xii.1986; Long
Pa Sia, confl. S. Maga and S. Pa Sia, 4°26’30”N 115°40’E,
high moist forest, sandsoil, 1350 m, 2 d d, 16-19.x.1986.
One d retained in MCZ.

Description. — Male: Head hidden, interocular
space greater than the distance between the ocellar
spots and antennal sockets. Pronotum suboval (fig.
14A). Tegmina and wings fully developed extending

RorH: Revision of Ctenoneura

beyond end of abdomen. Hind wing with intercalary
vein, media vein with one branch, cubitus vein with
five or six branches (some may be forked). Front fe-
mur Type C,; pulvilli absent, tarsal claws symmetri-
cal, simple, arolia minute. Abdominal terga unspe-
cialized. Supraanal plate transverse, subtrigonal (fig.
14E). Subgenital plate strongly asymmetrical, the
right side with a broad, elongated lobe that curves to-
wards the left where there is a broad, apically curved,
hook-like process; style absent (figs. 14B-E).

Colour. — Head dark reddish brown; basal four an-
tennal segments brown, succeeded by pale (yellowish)
antennomeres to about the middle, those on the distal
half brown, with as many as six terminal (or pretermi-
nal) white antennomeres (some lacked, or had only
two or three white segments, but these may have been
damaged specimens). Pronotal disk with a broad, dark
reddish brown macula that extends to both the anteri-
or and posterior margins, lateral areas usually hyaline
(fig. 14A), sometimes partly opaque yellowish.
Tegmina with humeral and proximal half of the costal
vein region yellowish-hyaline, the remainder dark hya-
line-brown. Hind wing with yellowish tinge in the co-
stal vein region. Abdomen brown, supraanal plate
with a white area, subgenital plate may be a lighter
brown and sometimes partly yellow.

Female: Unknown.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 14. Ctenoneura uncata sp. n., male paratype from near Long Pa Sia. — A. pronotum; B. subgenital plate (ventral); C. sub-
genital plate (left lateral); D. subgenital plate (rear view); E. supraanal and subgenital plates (dorsal).

Measurements (mm). — Length, 5.0-5.8; prono-
tum length x width, 1.6-1.7 x 2.1-2.3; tegmen length,
7.0-7.7; interocular space, 0.6.

Etymology. — The specific name refers to the hook-
shaped structure on the left side of the subgenital
plate.

Ctenoneura poringa Sp. n.

(figs. 15A-C)

Type material. — Holotype, d, Malaysia, Sabah, Poring
hot spring, Sg. Kepungit, E. of park boundary, c. 116°42°E
6°03’N, 480 m, ML, 26.xi.1986, J. Huisman; in RMNH.

Description. — Male: Head hidden under prono-
tum, interocular space greater than distance between
ocellar spots and antennal sockets. Pronotum sub-
oval. Tegmina and wings fully developed extending
beyond end of abdomen, the former with oblique dis-
coidal sectors. Hind wing intercalary vein present,
media vein with one branch, cubitus vein with six ra-

100

mi (one may be branched). Front femur Type C;;
pulvilli absent, tarsal claws symmetrical, simple, aro-
lia minute. Abdominal terga unspecialized. Supraanal
plate transverse. Subgenital plate asymmetrical, hind
margin deeply, widely excavated, with a large whitish,
membranous area bearing a cylindrical style at its ba-
sal end (figs. 15A-C).

Colour. — Head dark brown; two basal antennal
segments dark brown succeeded by about 25 yellowish
antennomeres, then three or four brown and three or
four white segments, terminating in one or two brown
ones (some terminal segments may have been miss-
ing). Pronotal disk dark reddish brown, anterior and
posterior areas darker, lateral regions hyaline.
Tegmina with humeral region yellow-hyaline, remain-
der brown-hyaline. Hind wing infuscated, darker
along the costal vein area. Abdominal terga brown, su-
praanal plate with a white medial zone. Abdominal
sterna light brown, subgenital plate darker. Cerci with
four basal segments brown, distal three segments yel-
lowish. Coxae and femora brown, tibiae and tarsi pale.

Rorn: Revision of Ctenoneura

0.25 mm

Fig. 15. Ctenoneura spp. A-C. C. poringa sp. n., male holotype. A. supraanal and subgenital plates (dorsal); B. subgenital
plate (ventral); C. subgenital plate (rear view). - D-G. C. /uma Sp. n., male holotype. D. supraanal and subgenital plates
(dorsal); E. subgenital plate (left lateral); F. subgenital plate (right lateral); G. subgenital plate (ventral).

Female: Unknown. Ctenoneura luma sp. n.

Measurements (mm). — Length, 5.0; pronotum (figs. 15D-G)
length x width, 1.6 x 2.0; tegmen length, 6.3; interoc- Type material. - Holotype, 4, Malay Penin., Perak,
ular space, 0.5. Larut Hills, 4000-4500 ft., ii. —iii.1905, R. Shelford (incor-

Etymology. — The species is named for Poring hot rectly labelled C. fulva, by Hanitsch); in HDEO. — Paratype.
spring. Perak. HDEO: Perak, F.M.S., Jor Camp, 2000 ft, 16,

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

0.5 mm

Fig. 16. Ctenoneura scutica sp. n., male holotype. A. supraanal and subgenital plates (dorsal); B. subgenital plate (ventral); C.

subgenital plate (rear view).

6.ix.1922, 2d, at light, 27.viii.1922, E. Sumiind (labelled
C. pumosa, mss name, by Hanitsch).

Description. — Male: Head barely exposed, interoc-
ular space greater than the distance between ocellar
spots and antennal sockets. Pronotum suboval.
Tegmina and wings fully developed, the former with
oblique discoidal sectors. Hind wing with distinct co-
stal veins, the proximal ones thickened distad, interca-
lary vein present, media vein bifurcate (one of the ra-
mi also branched), cubitus vein with six branches.
Front femur Type C,; pulvilli absent, tarsal claws sym-
metrical, simple, arolia minute. abdominal terga un-
specialized. Supraanal plate trigonal, entire (fig. 15D).
Subgenital plate asymmetrical, deeply excavated medi-
ally forming two dissimilar lobes, with a slender style
on the left side (the style was accidentally broken off
and lost after the drawings were made); on the right
side, a slender, thorn-like structure appears to arise
from the margin of the plate but actually lies pressed
tightly against the dorsal surface, and its apex pro-

trudes beyond the the right lobe margin (figs. 15E-G).

Colour. Head reddish brown. Pronotal disk red--

dish brown, the macula narrowly margined with yel-
low, lateral portions hyaline. Tegmina and wings pale
hyaline. Abdominal terga light brown, supraanal

102

plate with a white macula medially near the hind
margin, subgenital plate darker except for a pale zone
around the medial excavation. Cerci pale. Legs light
brown.

Female: Unknown.

Measurements (mm). — Length, 5.5; pronotum
length x width, 1.5 x 1.8; tegmen length, 6.3; interoc-
ular space, 0.6.

Etymology. — The specific name refers to the thorn-
like structure that arises on the dorsal surface of the
subgenital plate and protrudes beyond the right lobe.

Ctenoneura scutica sp. n.

(fig. 16)

Type material. — Holotype d, Malaysia, Sabah, one half
way Rd. Melligan, Long Pa Sia, 115°42°E 4°35’N, 1200 m,
ML, 15.xii.1986, J. Huisman; in RMNH.

Description. — Male: Head hidden, interocular
space greater than the distance between ocellar spots
or antennal sockets. Pronotum suboval. Tegmina and
wings extending well beyond end of abdomen, the
former with oblique discoidal sectors. Hind wing
with intercalary vein, media vein with one bifurcated
branch, cubitus with five branches. Front femur Type

C,; pulvilli absent, tarsal claws symmetrical, simple,
arolia minute. Abdominal terga unspecialized.
Supraanal plate symmetrical, transverse, short, hind
margin entire. Subgenital plate asymmetrical, the left
side concavely excavated bearing a small cylindrical
style in a white membranous area; near the mid region
of the plate is a deep, narrow, oblique (towards the
left) excision with a long, stout, tapering whip-like
process that curves upwards and anteriorly between
the smooth walls of the excision (figs. 16A-C).
Colour. — Head dark reddish brown; antennae
brown, monochromatic. Pronotal disk reddish
brown, darker near the anterior and posterior mar-
gins. Tegmina with humeral and proximal part of the
costal vein area yellowish-hyaline, remainder slightly
darker. Hind wing with a yellowish tinge, costal vein
region darker. Abdominal terga brown, lateral edges
and terminal segments darker, supraanal plate with a

RotH: Revision of Ctenoneura

white medial zone. Abdominal sterna light brown,
lateral edges darker, distal part of subgenital plate yel-
lowish. Cerci brown, terminal segment lighter. Legs
light brown.

Female: Unknown.

Measurements (mm). — Length, 5.6; pronotum
length x width, 1.8 x 2.2; tegmen length, 7.6; interoc-
ular space 0.6.

Etymology. — The specific name refers to the whip-
like process between the narrow incision on the sub-
genital plate.

Ctenoneura parascutica sp. n.

(fig. 17)

Type material. — Holotype, d, Malaysia, Sabah, base of
S. Nologana, 1010 m, Crocker range, 115°55’E 5°10°N,
UV, 10-11.x. 1986, J. Huisman (RMNH).

Fig. 17. Ctenoneura parascutica sp. n., male holotype. A. pronotum; B. Supraanal and subgenital plates (rear view); C. sub-
genital plate (right lateral); D. subgenital plate (left lateral); E. supraanal and subgenital plates (dorsal); F. left hind wing.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

0.25 mm

Fig. 18. Crenoneura kinabaluana sp. n., male paratype from Kiau, Mt. Kinabalu. A. supraanal and subgenital plates (dorsal);
B. subgenital plate (rear view); C. subgenital plate (left lateral); D. subgenital plate (right lateral).

Description. — Male: Head exposed, interocular
space greater than the distance between ocellar spots
and antennal sockets. Pronotum subelliptical (fig.
17A). Tegmina and wings fully developed extending
beyond end of abdomen, the former with oblique dis-
coidal sectors. Hind wing with the proximal three or
four costal veins clubbed, intercalary vein present,
media vein with one branch, cubitus vein with five
branches (the right wing has one of the branches
forked) (fig. 17F). Front femur Type C;; pulvilli ab-
sent, tarsal claws symmetrical, simple, arolia minute.
Abdominal terga unspecialized. Supraanal plate
transverse, hind margin entire (figs. 17B, E).
Subgenital plate asymmetrical, hind margin excised,
with a large dark, spine-like process that curves dor-
sad, its apex hidden under the supraanal plate; the
margins of the excision on either side of the spine-like
process are irregular, and on the left side the region is
unsclerotized; a small cylindrical style is on the left
side (figs. 17B-E).

Colour. — Head reddish brown; antennae brown
(distal halves missing). Pronotal disk reddish brown,
anterior and posterior border regions darker, a small
yellowish area anteriorly on each side along the edges
of the macula, lateral regions hyaline (fig. 17A).
Tegmina with humeral and proximal costal vein re-
gions yellowish-hyaline, remainder darker. Hind
wing with a yellowish tinge in the costal vein region
(fig. 17F). Abdominal terga brown, supraanal plate
with a white macula posteromedially (fig. 17E).
Abdominal sterna brown, subgenital plate darker.

104

Cerci brown, segments on distal half lighter. Legs
pale.

Female: Unknown.

Measurements (mm). — Length, 5.2; pronotum
length x width, 1.7 x 2.3; tegmen length, 7.3; interoc-
ular space, 0.6.

Comments. — I place parascutica close to scutica be-
cause both have a large, darkly sclerotized, curved,
spine-like process on the hind margin of the subgeni-
tal plate.

Ctenoneura kinabaluana sp. n.

(fig. 18)

Material. — Holotype, &, B.N. Borneo [Sabah], Mt.
Kinabalu, Lumu Luma, 3500 ft., 16.iv.1929 [labelled C.
fulva, by Hanitsch] (HDEO). — Paratypes. Sabah. Same local-
ity and date as holotype, 3500 ft, 16; Mt. Kinabalu,
Kamboranga, 7000 ft, 2d, 30.iii.1929 (HDEO); Malaysia,
Sabah, Mt. Kinabalu Kg. Kiau, S. Kadamaian, 116°31’E
6°02’N, 720 m, UV, 1d 26.viii.1986, J. Huisman (RMNH).

Description. — Male: Head hidden, interocular
space greater than the distance between ocellar spots
and antennal sockets. Pronotum suboval. Tegmina
and wings fully developed extending well beyond end
of abdomen, discoidal sectors of the former oblique.
Hind wing with distinct costal veins, intercalary vein
present, media vein with one branch (the branch may
be bifurcate), cubitus vein with six branches (some
may be forked; also one of the wings has two veins
fused a short distance from their bases and then sep-

arate). Front femur Type C,; pulvilli absent, tarsal
claws symmetrical, simple, arolia minute. Abdominal
terga unspecialized. Supraanal plate transverse, hind
margin convex, entire (fig. 18A). Subgenital plate
asymmetrical, with a short, median process that ex-
tends postero-obliquely, or dorsad, its distal end cury-
ing anteriorly towards the supraanal plate; a cylindri-
cal style is located on the left side and the margin to
the right of the process is prolonged (figs. 18A-D).
The length of the median process varies and is longest
in the specimen from Kiau.

Colour. — Head reddish brown; antennae brown.
Pronotal disk reddish brown, lateral zones hyaline.
Tegmina with humeral and costal vein zones yellow-
hyaline, remainder brown-hyaline. Hind wing infus-
cated, darkest in the costal vein zone. Abdominal ter-

Roth: Revision of Ctenoneura

ga brown, supraanal plate with a white macula ante-
romedially. Abdominal sterna brown, distal margin
of subgenital plate yellowish. Cerci and legs brown.
The specimen from Kiau is smaller and darker than
the others: head dark brown, basal three antenno-
meres brown, remainder yellowish (distal segments of
the antennae missing); supraanal plate yellow lateral-
ly, and with a white area posteromedially (fig. 18A).
Cerci with basal segments brown, distal cercomeres
yellowish. Coxae, femora, and tibiae brown, tarsi
lighter.

Female: Unknown.

Measurements (mm) (male from Kiau in parenthe-
ses). — Length, 6.5-6.6 (5.3); pronotum length x
width, 1.8-1.9 (1.5 x 2.1); tegmen length, 9.0 (6.7);
interocular space, 0.6-0.7 (0.6).

0.25 mm

Fig. 19. Ctenoneura triprocessa sp. n., male holotype. A. subgenital plate (ventral); B. subgenital plate (dorsal); C. subgenital
plate (rear view); D. subgenital plate (right lateral); E. subgenital plate (left lateral); F. left hind wing.

105

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Ctenoneura triprocessa Sp. n.

(fig. 19)

Type material. — Holotype d, Malaysia, SE. Sabah,
Danum Valley Field C., c. 150 m, W.O., Mal. trap 5,
22.xi.-4.xii.1987, C. v. Achterberg & D. Kennedy (RMNH)
[The type is fragmented, head and thorax, tegmina and
wings, and terminal abdominal segments are mounted on
cards].

Description. — Male: Head hidden, interocular
space greater than the distance between ocellar spots
and antennal sockets. Pronotum suboval. Tegmina
and wings fully developed extending beyond end of
abdomen, discoidal sectors of former oblique. Hind
wing with a intercalary vein, media vein with one
branch, cubitus vein with three branches (one of
which may or may not be branched near its base) (fig.
19F). Front femora missing; other legs lack pulvilli,
tarsal claws symmetrical, simple, arolia minute.
Abdominal terga unspecialized. Supraanal plate
transverse, hind margin convexly rounded, entire.
Subgenital plate asymmetrical, trigonal Ge, 194),
hind margin with three long processes that curve dor-
sad; arising on the dorsal surface of the plate near the
left process is a long filament that protrudes between
the left and middle processes; style absent (figs. 19B-
E).

Colour. — Head dark reddish brown; antennae
with pale yellowish segments (about half the segments
are missing). Pronotal disk reddish brown, slightly
lighter on the anterior and posterior regions, lateral
areas clear hyaline. Tegmina with humeral and proxi-
mal half of the costal vein area yellowish-hyaline, re-
mainder light brown-hyaline. Hind wing with yel-
lowish tinge mostly in the costal vein zone (fig. 19F).
Abdomen brown, distal part of supraanal plate white,
subgenital plate yellowish brown.

Female: Unknown.

Measurements (mm). — Pronotum length x width,
1.5 x 1.8; tegmen length, 5.7; interocular space, 0.5.

Etymology. — The specific name refers to the three
prominent processes or prolongations on the hind
margin of the subgenital plate.

Ctenoneura murudensis Sp. n.

(fig. 20)

Ctenoneura fulva, Hanitsch (in part, 6 paralectotype) 1925:
101.

Type material. — Holotype, d paralectotype (Type Orth.
300°/,) of Ctenoneura fulva Hanitsch, Mt. Murud
[Sarawak], 6500 ft. (HDEO).

Description. — Male: Head hidden, interocular

Fig. 20. Crenoneura murudensis sp. n., male holotype. A. supraanal and subgenital plates (rear view); B. supraanal and sub-
genital plates (dorsal); C. subgenital plate (ventral); D. subgenital plate (right lateral); E. subgenital plate (left lateral).

106

space greater than the distance between ocellar spots
and antennal sockets. Pronotum suboval. Tegmina
and wings fully developed extending beyond end of
abdomen, the former with oblique discoidal sectors.
Hind wing with distinct costal veins, the proximal
ones clubbed distally, intercalary vein present, media
vein forked about at the middle, cubitus with eight
unbranched rami. Front femur Type C,; pulvilli ab-
sent, claws simple, symmetrical, arolia small. Abdom-
inal terga unspecialized. Supraanal plate transverse,
deflexed, hind margin convex, entire (figs. 20A-B).
Subgenital plate asymmetrical with a small, dark, cy-
lindrical style located in a membranous area on the
left side; a darkly sclerotized lobe with a U-shaped ex-
cavation on its apical margin arises medially, and on
the right side is a small acute prolongation (figs. 20A-
B)

Colour. — Head light reddish brown; antennae
dark brown, monochromatic. Pronotal disk light red-
dish brown, surrounding areas hyaline. Tegmina with
humeral and proximal half of the costal vein zone yel-
lowish-hyaline, remainder darker, hyaline. Wings
pale, with a yellowish tinge, darker in the clubbed co-
stal vein region. Abdominal terga uniformly light
brown, supraanal plate without a medial white spot.
Abdominal sterna yellowish-brown, distal part of
subgenital plate yellow. Cerci light brown dorsad,
darker ventrally. Legs light brown.

Female: Unknown.

Measurements (mm). — Length, 6.7; pronotum
length x width, 1.8 x 2.3; tegmen length, 8.5; interoc-
ular space, 0.7.

Comments. — Hanitsch incorrectly considered this
as fulva, but the subgenital plate of that species (fig.
8B) is distinctly different from murudensis (fig. 20C).

Ctenoneura crassistyla sp. n.
(figs. 21A-D)
Type material. — Holotype, d, Pahang, F. M. S.,

Cameron’s Highlands, Tanak...., 4800 ft., 22.v.1931, H.
M. Pendlebury (HDEO).

Description. — Male: Head hidden, interocular
space greater than the distance between ocellar spots
and antennal sockets; antennae missing. Pronotum
suboval. Tegmina and wings fully developed extend-
ing beyond end of abdomen, the former with oblique
discoidal sectors. Hind wings with distinct costal
veins, media vein bifurcate, cubitus vein with six
branches. Front femur Type C,; pulvilli absent, tarsal
claws symmetrical, simple, arolia minute. Abdominal
terga unspecialized, supraanal plate strongly trans-
verse, very short, hind margin almost truncate, entire
(fig. 21A). Subgenital plate asymmetrical, hind mar-
gin deeply, unevenly excavated, with a large, stout

Rorn: Revision of Ctenoneura

style on the basal corner of the left side of the excava-
tion; the apex of the style is split and a small piece of
‘fleshy’ tissue protrudes between the ends; right side
with distal region concavely depressed (figs. 21B-D).

Colour. — Head reddish brown. Pronotum with
narrow yellow edging, disk reddish brown, lateral re-
gions hyaline. Tegmina hyaline yellow, darker on the
anal field region. Hind wings pale with yellowish
mostly in the costal vein region and apical part of the
anterior field. Abdominal terga light brown, supraa-
nal plate yellowish with a medial white area near the
hind margin (fig. 21A). Abdominal sterna brown,
subgenital plate darker. Cerci pale dorsally. Legs light
brown.

Female: Unknown.

Measurements (mm). — Length, 6.4; pronotum
length x width, 1.9 x 2.5; tegmen length, 8.3; interoc-
ular space, 0.7.

Etymology. — The specific name refers to the large,
stout, left style.

Ctenoneura sp.

Material. — South Vietnam. HDEO: 1d, S. Annam,
Longbian Peaks, 5500-7500 ft, iv.-v.1918, C. B. Kloss (la-
belled Ctenoneura major Hanitsch, by Hanitsch).

Comments. — This species has a subgenital plate
similar to that of crassistyla, with a large, stout, style
on the basal corner of the left side (the style is not
split at the apex). The media vein of the hind wing
has two branches, and the cubitus vein has seven
branches, one or two of which may be branched. The
tarsal claws are long, simple, symmetrical, arolia ab-
sent. The supraanal plate is transverse, hind margin
weakly convex. This is the only species recorded from
Vietnam.

Measurements (mm). — Length, 7.8; pronotum
length x width, 2.3 x 3.1; tegmen length, 10.2; inter-
ocular space, 0.8.

Ctenoneura simulans Bey-Bienko

(figs. 21 E-F)

Ctenoneura simulans Bey-Bienko, 1969: 833, figs. 2,3, (d)
(Russian). Holotype 3, S. China, Yunnan: Hsiamonyang
vicinity, 900-1100 m, 6.v.1957, Hung Kuang-ti (ZINR)
[not examined]; 1970: 528 (English translation).

Description (from Bey-Bienko). — Male: Small.
Vertex passing roundly into frons, interocular space
two and one half times wider than distance between
antennal sockets. Pronotum somewhat less than one
and one half times as wide as long. Hind wings with
cross branches of the radius vein distinct, intercalary
vein between the radius and media, almost straight;

107

TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 21. Ctenoneura spp. — A-D, C. crassistyla sp. n., male holotype. A, supraanal and subgenital plates (dorsal); B. subgenital
plate (ventral); C. subgenital plate (left lateral); D. subgenital plate (right lateral). — E, F. C. simulans Bey-Bienko; E. sub-
genital plate (ventral); F. subgenital plate (rear view) (redrawn from Bey-Bienko, 1969).

cubitus vein with six branches, the first one from the
wing base bifurcate. Subgenital plate when viewed
from below as in fig. 21E; when viewed from behind
it has an irregular appendage on the right (fig. 21F).
Cerci moderately long, fourth to sixth segments oval,
tip of apical segment blunt, rounded.

Colour. — Brownish yellow. Head dark chestnut.
Pronotum chestnut brown with broad, hyaline lateral
parts. Wings just perceptably infuscated.

Female. Unknown.

Measurements (mm). — Body length, 8.0; prono-
tum length x width, 2.2 x 3.0; tegmen length, 9.0;
overall length, 11.5.

108

Comments. — According to Bey-Bienko, outward-
ly this species resembles misera, but is clearly distin-
guished from all known species in the genus by the
structure of the subgenital plate. Bey-Bienko does not
mention whether or not there is a style (which could
be small on the dorsal surface of the plate, and not
visible in ventral or rear views).

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. I am most grate-
ful to Dr. Horst Bohn, Zoologisches Institut der
Universitat Miinchen, for taking the time from his
own taxonomic studies to review my manuscript.

REFERENCES

Bey-Bienko, G. Ya., 1957. Blattoidea. I. The results of the
Chinese-Soviet zoologico-botanical expeditions to
southwestern China 1955-1956 . — Entomologicheskoe
Obozrenie, Moskva 36: 895-915. (Russian; English sum-
mary).

Bey-Bienko, G. Ya., 1969. New genera and species of cock-
roaches (Blattoptera) from tropical and subtropical Asia.
— Entomologicheskoe Obozrenie, Moskva 48: 831-862.
(Russian). (English translation of this article:
Entomological Review, Washington 48 (1970): 528-
548).

Bruijning, C. F. A., 1948. Studies on Malayan Blattidae. —
Zoologische Mededelingen Leiden, 29: 1-174.

Hanitsch, R., 1925. On a collection of Blattidae from
northern Sarawak, chiefly Mt. Murud and Mt. Dulit. —
Sarawak Museum Journal 3 (1): 75-106.

Hanitsch, R., 1927. On a collection of Blattidae from south-
ern Annam. — Journal of the Siam Society, Natural
History Supplement 7(1): 7-48.

Hanitsch, R., 1928. Spolia Mentawiensia: Blattidae. —
Bulletin Raffles Museum 1: 1-44.

Hanitsch, R., 1929a. Fauna Sumatrensis. Blattidae. —
Tijdschrift voor Entomologie 72: 263-302.

Rotu: Revision of Ctenoneura

Hanitsch, R., 1929b. Dr. E. Mjöberg’s zoological collec-
tions from Sumatra. 11. Blattidae. — Arkiv for Zoologi
21A (2): 1-20.

Hanitsch, R., 1932a. On a collection of blattids from the
east coast of Sumatra. — Miscellanea Zoologica
Sumatrana 62: 1-8.

Hanitsch, R., 1932b. Beccari and Modigliani’s collection of
Sumatran Blattidae in the Museo Civico, Genoa. —
Annali del Museo civico di Storia naturale di Genova 56:
48-92.

Hanitsch, R., 1933. On a collection of Bornean and other
oriental Blattidae from the Stockholm Museum. —
Entomologisk Tidskrift 54: 230-245.

Princis, K., 1950. Entomological results from the Swedish
expedition 1934 to Burma and British India. — Arkiv for
Zoologi 1: 203-222.

Princis, K., 1954. Kleine Beitrige zur Kenntnis der
Blattarien und ihrer Verbreitung. VI. — Entomologisk
Tidskrift 74 (1953): 203-213.

Princis, K., 1963. Blattariae: Subordo: Polyphagoidea, Fam.
Polyphagidae. /n Beier (ed.): Orthopterorum Catalogus
4: 77-172. ’s-Gravenhage.

Princis, K., 1967. Kleine Beitrige zur Kenntnis der
Blattarien und ihrer Verbreitung. X. — Opuscula
Entomologica 32: 141-151.

Princis, K., 1971. Blattariae: Subordo Epilamproidea, Fam.:
Ectobiidae. /n Beier (ed.): Orthopterorum Catalogus. 14:
1039-1224. ’s-Gravenhage.

Received: 12 November 1992
Revised version accepted: 19 February 1993

109

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GEORGE W. BECCALONI MCZ

NERC Centre for Population Biology, Imperial College, UK L IB RA R Y

JAN 1 4 1994
A NEW SPECIES OF EXTATOSOMA GRAY
(PHASMATODEA: PHASMATIDAE) FROM PAPUA
NEW GUINEA, WITH REMARKS ON THE SPECIES IN
ime GENUS

HARVARD
UNIVERSITY

Beccaloni, G. W., 1993. A new species of Extatosoma Gray (Phasmatodea: Phasmatidae) from
Papua New Guinea, with remarks on the species in the genus. — Tijdschrift voor Entomologie
136: 113-123, figs. 1-16, tabs. 1-2. [ISSN 0040-7496]. Published 10 December 1993.
Extatosoma carlbergi sp. n. (Phasmatidae), is described from Papua New Guinea. The biology,
ecology and distribution of this and the three other species of Extatosoma Gray is discussed and
a hypothesis is advanced to explain the discrete altitudinal ranges of the two New Guinean spe-
cies. The phylogenetic relationships of the species are considered and a key to the adult females
is presented.

G. W. Beccaloni, Department of Entomology, The Natural History Museum, Cromwell Road,
London SW7 5BD, UK.

Key words. — Phasmatodea, Phasmida, Extatosoma, New Guinea, Australia, biology, ecology,

distribution, phylogeny.

The genus Extatosoma is widely known due to fa-
miliarity with an Australian species, £. tiaratum
(MacLeay 1826). This is one of the stick-insects most
commonly cultured by amateur enthusiasts (Bragg
1991) and it is also a popular laboratory animal on
which many physiological, behavioural and morpho-
logical studies have been conducted (for references see
Carlberg 1983, 1987a).

Two other species are currently recognised as be-
longing to the genus : E. bufonium Westwood, 1874
from Australia and £. popa Stal, 1875 from New
Guinea. Another nominal species, E. elongatum
Froggatt, 1922, was synonomised with E. bufonium
by Vickery (1983). The holotype of E. bufonium (in
the Hope Entomological Collection, Oxford, UK) is
a penultimate (?) instar female nymph, not a male as
stated by Vickery (1983), and Froggatt (1922) based
E. elongatum on the adult female of this species. A
third, as yet undescribed, species may exist in the vi-
cinity of Brisbane, Queensland, Australia (P. D.
Brock, in prep.). The adult female of this taxon (fig.
in Wilson 1991) is superficially similar to E. tiaratum
in appearance, while the egg resembles that of £. bu-
fonium.

Both £. popa and E. bufonium are poorly known
and only the former species has been cultured in cap-
tivity (Harman 1985). The males of these two species
have not been figured or formally described, although
Harman briefly noted that the male of £. popa can be

distinguished from that of E. tiaratum by its clear
wings and a white dot behind its head. From person-
al observation of a series of preserved adult male £.
popa cultured by Harman and deposited in the BMNH,
it is evident that the ‘white dot’ referred to is in fact a
pale v-shaped marking on the mesonotum, similar to
that possessed by the adult female of this species.

The existence of a second New Guinean species of
Extatosoma was first suggested in print by Harman
(1985) who stated that there is ‘....another species [of
Extatosoma] which occurs in the mountainous regions
of Papua New Guinea. I have seen a number of spec-
imens in the collection at the Wau Ecology Institute
Museum’. During July and August of 1990 the au-
thor led an Imperial College (University of London)
expedition to Papua New Guinea (PNG). For the
first four weeks we were based at the Wau Ecology
Institute (WEIC) and the author took the opportunity
to investigate Harman’s claim. Three adult females of
an undescribed Extatosoma were present in the WEIC
collection, together with a note stating that the taxon
was new.

A reward was offered to two Papuan youths to col-
lect living specimens of this species and one day later
an adult female (fig. 1) was received. This individual
was placed in a cylindrical net cage on cut foodplant
standing in water and was kept alive for twelve days
for observation.

Despite further requests and a search of the locality

113

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 1. Extatosoma carlbergi sp. n. — Holotype © resting on
twig of Calliandra surinamensis Benth.

from which the specimen had been collected, no
other individuals were obtained. Several local
Papuans who were interviewed stated that they were
familiar with this species but that it was only rarely
seen (more rarely than E. popa).

MATERIALS AND METHODS

The terminology used to describe eggs is after
Clark (1976a), with the following additions and mo-
difications : ‘total egg length’ is the length of the egg
capsule plus the length of the operculum ; ‘length of
operculum’ is the maximum length of the operculum
from its base to its summit ; ‘diameter of operculum’
is the maximum diameter across the base of the oper-
culum ; ‘micropylar arm’ is a new term given to the
lateral extensions of the micropylar plate located on
either side of the micropylar cup region ; ‘length of
micropylar arm’ is the length of one micropylar arm
measured from the centre of the micropylar cup to
the apex of the arm; “distance across micropylar arms’
is the maximum distance between the tips of the mi-
cropylar arms ; ‘height of micropylar mound’ is the
height with which the micropylar plate is raised abo-
ve the surface of the egg capsule and is measured mid-
way along the plate’s length; ‘chorionic membrane’ is
a new term given to the membrane or layer which co-
vers the entire outer surface of the egg. This membra-
ne expands and detaches from the egg surface if the
egg is immersed in alkali (e.g. sodium hydroxide solu-
tion). The membrane is thinner and weaker around

114

the base of the operculum and the portion of mem-
brane covering the operculum remains attached to it
when it detaches on hatching of the egg. The eggs of
all four Extatosoma are covered by a chorionic mem-
brane, but it appears to be absent from the eggs of
other stick-insect species (Clark 1976b); ‘capitulum’,
in light of the recent discovery that this structure has
a function analogous to the elaiosome of certain plant
seeds, and acts as a handle which enables ants to carry
stick-insect eggs back to their nests (Compton &
Ware 1991, Hughes & Westoby 1992), the author
proposes to restrict Clark’s (1976a) definition to
structures attached to and protruding from the sur-
face of the operculum.

The coloration of £. carlbergi is described from
slides of the living holotype taken on ‘Kodachrome
64’ slide film under natural lighting conditions.
Specimens of E. carlbergi and E. popa collected by the
author were killed by freezing and preserved by injec-
tion of embalming fluid (formula in Cogan & Smith
1974). Eggs (fully developed) were removed from the

NS

i

si
ik
#

4
3
1
ad
TA
a

Fig. 2. Extatosoma carlbergi sp. n. — Holotype 9, dorsal hab-

itus.

20 mm

Figs. 3-4. Egg
of Extatosoma
carlbergi sp. n.
taken from ovi-
positor of holo-

type. — 3, dorsal
view; 4, lateral
view.

abdomen of the £. carlbergi holotype by first relaxing
the specimen (in a sealed plastic box with damp tissue
paper in the base) and then by softening a section of
intersegmental membrane between tergites and ster-
nites with 10% aqueous ammonia solution, so that an
incision could easily be made and the eggs removed
with forceps. The chorionic membrane was removed
from eggs for study, by placing the eggs for c. 5 min-
utes in a 5% aqueous solution of sodium hydroxide at
room temperature. Drawings were produced using a
drawing tube on a Wild M5 stereomicroscope. All
measurements are given in millimetres. Measure-
ments of eggs (accurate to 0.01 mm) were made using
an ocular micrometer, except for micropylar plate
length, which because of the plate’s curvature, was
first drawn using a drawing tube and then measured
using inelastic fishing line and a ruler. All other meas-
urements (accurate to 1 mm) were taken from speci-
mens using a pair of dividers and a ruler.

Abbreviations of depositories

BMNH: The Natural History Museum, London,
England; BPBM: Bishop Museum, Hawai'i; TPNG:
Central Reference Insect Collection, Department of
Agriculture and Livestock, Port Moresby, Papua New
Guinea; FICB: National Forest Insect Collection,
Forest Research Institute, Lae, Papua New Guinea;
NHMN: Nottingham Natural History Museum,
Nottingham, England; UPNG: University of Papua
New Guinea, Port Moresby, Papua New Guinea;
WEIC: Wau Ecology Institute, Wau, Papua New
Guinea.

BECCALONI: A new species of Extatosoma

‘TAXONOMY

Extatosoma carlbergi sp. n.

(figs. 1-5, 8, 12-14)

Type material. — Holotype 9: Papua New Guinea,
Morobe Province, Wau, golf course, c. 1150 m, ex
Calliandra surinamensis Benth., 14.vii.1990, G. W.
Beccaloni (BMNH). — Paratypes: PAPUA NEW GUINEA: 29,
Southern Highlands Province, Tari, 1600 m, ex Casuarina,
4.x.1963, R. Straatman (BPBM); 19 (fragmented), Chimbu
Province, Karimui, 4.vi.1961, Gressitt (BPBM); 1 2, Morobe
Province, Wantoat, feeds on Casuarina, 1.xii.1957, R. W.
Paine (BMNH).

Additional material examined. — NEW GUINEA: 19, no
other data (NHMN); PAPUA NEW GUINEA: 22, Southern
Highlands Province, Pangia, defoliating Casuarina oligodon,
1.vi.1971, J. Lowien (FICB); 19, Southern Highlands
Province, Pangia, found on Casuarina trees, 30.iv.1971, J.
Wallis (TPNG); 22, no data (WEIC); 12 (nymph, penulti-
mate ? instar), Morobe Province, Wau, 1200 m, 14.x.1966,
G. A. Samuelson (BPBM); 1 2, Morobe Province, Wau, 1100
m, collected on Casuarina sp., 10.xii.1974, A. D. Hart
(WEIC); 1 2, Morobe Province, Wau, Wau Ecology Institute
Office, 1150 m, 27.xi.1986, J. Somp (UPNG).

Description

Based on adult female holotype and four adult fe-
male paratypes. Measurements of selected structures
given in table 1. Male unknown.

Female (adult) . — Head typical for the genus; hy-
pognathous with conical occiput. Occipital medial
spines large and robust; four pairs anterior to crest
and two pairs posterior to crest. A large lamellate
compound spine present on either side of occipital

115

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 5-6. Lateral views of ab-
domens of adult Q
Extatosoma spp. showing spi-
nation on tergites 3-10. — 5,
Extatosoma carlbergi sp. n.
(holotype); 6, Extatosoma tia-
ratum.

crest. Eyes small and globular, situated anterolateral-
ly. Ocelli indicated by slight swellings covered by in-
tegument. Antennae simple, with 25 segments (n=2).
A large spine present between eyes and antennal sock-
ets.

Pronotum elongate-rectangular, slightly rounded
posteriorly. Spines around margin variable in size and
number; a pair of large medial lamellate spines usual-
ly present on posterior third of segment.

Mesonotum with a well developed pair of pre-me-
dian spines and a large pair of compound lamellate
spines situated between tegminal bases. Lateral bor-
ders with 3-6 large spines, the two directly above the
tegminal bases often confluent basally. Tegmina re-
duced, lobose, extending to posterior margin of meta-
notum.

Metanotum with a pair of well developed lamellae
above posterior margin. Wings rudimentary, extend-
ing into anterior third of first tergite.

Tergites one to nine bearing a pair of medial com-
pound lamellate spines on the posterior third, with a
well developed simple spine posterior and to either
side of these. The lamellate spines on tergites five and
six especially well developed (fig. 5) ; those on tergites
seven to nine are sometimes simple. The posterolater-
al margins of tergites two to nine expanded laterally

116

and strongly serrated or spined. The anterolateral
margin of the tenth tergite is similarly produced.
Lateral expansions of tergites five to seven are c. twice
the size of those on the other tergites. Tenth tergite
usually with a pair of medial spines on posterior half
and the hind margin has large posteriorly directed
spines (fig. 5).

Supra-anal plate with large pair of posteriorly di-
rected medial spines. Cercus 2.6 mm in length, bi-
lobed, setose (fig. 14). Both structures concealed be-
neath tenth tergite.

Operculum keel-shaped, the anterior third ventral-
ly expanded with a serrated edge (fig. 5). Sternites two
to seven with many long to short irregularly arranged
robust spines. A pair of double weakly confluent me-
dial spines may be present posteriorly. Spination of
sternites four to six reduced in comparison with that
of sternites two to three. Meso- and metathoracic ba-
sisterna with few irregular short robust spines.
Ventral margins of meso- and metathoracic episterna
with long to short laterally or ventrally directed
spines.

Fore femur with strongly serrated dorsal and later-
al expansions. Fore tibia with dorsal expansion arcu-
ate and ventral expansion denticulate (fig. 8).
Proximal tarsomere of fore leg expanded into lobe

BECCALONI: A new species of Extatosoma

Table 1. Measurements of selected structures of type specimens and female (? penultimate instar) nymph of £. carlbergi. All
measurements in mm (accurate to 1 mm). “excluding occipital spines; “n = 1.

Structures measured Holotype Paratypes Nymph
(Lengths) (n=4)
x+SD Range

Body 128 IEN) 116 - 140 70
Antenna 27 - — 11
Head ' 19 195=#13 18-21 11
Pronotum 9 10.5 + 1.3 Se 2 5
Mesonotum 15 16.0 + 0.8 15-17 10
Metanotum 12 11.5 + 0.6 lil = 12 5
Fore femur 25 DD SEA DAT 15
Median femur 20 20.3 + 1.0 19-21 11
Hind femur 30 28.8 + 1.0 28 - 30 15
Fore tibia 23 23.5 + 0.6 23 - 24 13
Median tibia 21 Mil) ze 12 VO = 22 12
Hind tibia 32 DAB ae 17 3135 16
Operculum 26 24.3 + 0.5 VA = WS) 7

dorsally towards distal end. Dorsal surfaces of median
and hind femora (figs. 12-13) with serrated expansion
distally. Outer ventral margin serrated and expanded
ventrally. Inner ventral margin spined and expanded
distally. Dorsal expansions of hind tibia usually
strongly arcuate (fig. 12), rarely weakly so (fig. 13);
median tibia likewise. Inner and outer ventral mar-
gins of median and hind tibiae with large ventrally di-
rected spines. Legs and body densely setose.

Coloration in life: Mesonotum with conspicuous
v-shaped marking which is paler than rest of segment
(this marking is usually white or cream in preserved
specimens). Arms of ‘v’ extend from anterolateral cor-
ners of segment, with base of ‘v below middle of seg-
ment. Area between arms usually black or dark
brown. Dorsal and ventral surfaces of body (includ-
ing spines, tegmina and wings) pale green with dark-
er green, white and black mottling; mirrored on each
side of body. Intersegmental membranes cream to
light brown. Legs with similar coloration to rest of
body; dorsal surfaces with a higher incidence of larger
black and dark green marks often arranged as trans-
verse stripes; ventral surfaces (less so of fore legs) apos-
ematically patterned with thick irregular black, ochre-
ous and white transverse bands. Median and hind
tibiae often with black predominating.

Nymph (female, penultimate ? instar). —
Measurements given in table 1. Morphology similar
to that of adult, except that wings not developed
(small, c. 1.5 mm long wing buds present) and oper-
culum is relatively shorter; not extending beyond pos-
terior margin of ninth tergite and not covering the su-
pra-anal plate or cerci. Coloration in life presumed
similar to that of adult, as coloration is similar to that

of the preserved adult specimens examined (i.e. with
light to dark brown replacing pale green ground
colour).

Egg (figs. 3-4). — Measurements given in table 2.
Operculum conical. Chorionic membrane transpar-
ent, not thickened and modified into a capitulum on
the operculum (see below). Capsule smooth and
glossy, marked with mottled pattern of black and grey
on cream. Micropylar plate cream, slightly raised
above surface of capsule on micropylar mound, with
lateral arm on either side of the micropylar cup region.

Etymology
This species is named in honour of Dr. Ulf

Table 2. Means, sample standard deviations and ranges of
dimensions of 17 eggs from E. carlbergi holotype (1 egg tak-
en from ovipositor, 16 extracted from abdomen). All meas-
urements in mm (accurate to 0.01 mm).

Egg dimensions x + SD Range
Total egg length 6.54+0.12 6.40 - 6.85
Capsule length 5.170072 5062531
Capsule height 4.49 +0.03 4.42 - 4.56
Capsule width (maximum) SDE 3771025284
Length of operculum 1.37220.12 NAR = 1154
Diameter of operculum

(maximum) 2.32+0.07 2.20 - 2.43
Micropylar plate length 6.09+0.28 5.70 - 6.73
Micropylar plate width

(at middle) 0.67 +0.03 0.61 - 0.70
Length of micropylar arm 1.74+0.06 1.60 - 1.79
Distance across micropylar

arms 3.00+0.07 2.88 - 3.07
Height of micropylar mound 0.15+0.03 0.13 - 0.19

iy

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

L
L

Syn

118

Carlberg, in recognition of his work on and passion
for Extatosoma.

Biology and behaviour

In common with the other members of the genus,
E. carlbergi is arboreal and when not feeding hangs
motionless upside down under thin branches of the
host tree (fig. 1). The holotype was collected on
Calliandra surinamensis (Leguminosae), an introduc-
tion from South America commonly grown in PNG
gardens (Verdcourt 1979). In captivity the specimen
accepted Leucaena sp. (Leguminosae) and guava,
Psidium sp. (Myrtaceae). Both originate from the
Neotropics and the former is cultivated in Wau as a
shade tree for coffee, while the latter is grown for its
fruit. J. Somp (pers. comm.) informed the author that
he has collected E. carlbergi on Sterculia sp. (Sterculia-
ceae), and Casuarina sp. (Casuarinaceae) is the only
foodplant recorded on the data labels of this taxon
examined during the course of this study.

Both £. tiaratum and E. popa are also polyphagous.
The latter has been collected on Casuarina, Diospyros
ferrea (Ebenaceae), C. surinamensis and Hibiscus sp.
(Malvaceae) (taken from data labels listed in
Appendix) and it will also feed on Leucaena and
Psidium in captivity (pers. observation). For a list of
the foodplants of E. tiaratum see Carlberg (1987b).
The only foodplant known for E. bufonium is
Japanese plum, ? Prunus cerasifera (Rosaceae) (from
data label of adult female in Queensland Museum,
Brisbane, Australia; collected at Mt. Tamborine, near
Brisbane, December 1924).

When disturbed the £. carlbergi holotype was ob-
served to exhibit a defensive behaviour similar to that
shown by adult females of £. tiaratum (Carlberg,
1981) and E. popa (pers. observation). The abdomen
is curled over the body, presenting the dense spina-
tion on the sternites towards the potential predator.
The hind legs are raised and extended to reveal the
aposematic pattern of black, white and ochreous
bands on the ventral surfaces of the femur and tibia.
This threat posture is maintained until any possible
danger has passed. Should a predator approach too
close, the insect will attempt to grasp it between the
femur and tibia of the hind legs; retracting and ex-
tending the legs with a rapid, synchronous scissor-like
action. The ventral margins of these appendages are
furnished with sharp downwardly directed spines,
which are capable of inflicting injury to human skin if
caught between them.

The egg of E. carlbergi (figs. 3-4), which resembles

BECCALONI: A new species of Extatosoma

a seed, is held in the ovipositor for several hours and
is then catapulted to the ground with a vigorous flick
of the abdomen. Eggs of E. popa (A. J. E. Harman,
pers. comm.) and £. tiaratum (Carlberg, 1984) are
deposited in a similar manner.

ECOLOGY AND DISTRIBUTION OF THE EXTATOSOMA
SPECIES

The known distributions of the two New Guinean
Extatosoma species is shown in fig. 16. The locality re-
cords plotted on this map were taken from data labels
of specimens examined by the author or from the li-
terature. Records for E. popa are listed in the
Appendix while those for £. carlbergi are listed under
the species description. Dubious records or possible
misidentifications have been omitted.

From fig. 16 it can be seen that £. popa has a wider
recorded geographical distribution than £. carlbergi,
ranging from Jayapura in Irian Jaya to Mt. Obree in
the Owen Stanley Range, PNG. Altitude appears to
be the most important factor in determining the dis-
tributions of these two species. Giinther (1929)
placed E. popa in an altitude band of 600-1500 m
above sea level. However, the highest that this species
has been recorded is c. 1200 m at Budemu, PNG. In
contrast, all known specimens of Æ. carlbergi have
been collected in the range 1100 m (Wau, PNG) to
1600 m (Tari, PNG). Wau (1100-1200 m) and
Wantoat, PNG, (1200 m), where the species are sym-
patric, are located within the c. 100 m altitudinal
overlap of the two species ranges.

The altitudinal ranges of E. popa and E. carlbergi
match the distributions of two discrete vegetation
belts, lower montane rain forest and mid-montane
rain forest respectively, which differ from each other
both in terms of species composition and in the struc-
ture and physiognomy of the vegetation. Lower mon-
tane rain forest has a mean canopy height of 30-40 m
and is distinctly drier in aspect than the mid-mon-
tane, with few species of ferns and mosses. In con-
trast, the mid-montane rain forest has a mean canopy
height of 20-30 m and its main characteristic is the
abundance and high species diversity of terrestrial and
epiphytic ferns, mosses and lichens (Johns 1982).

It is proposed that the gross differences in colour
pattern and morphology seen between E. popaand E.
carlbergi (nymphs and adult females only; adult males
of Extatosoma are macropterous and differ from
nymphs and adult females in both body form and be-
haviour) are the result of these species evolving cryp-

Figs. 7-15. Extatosoma spp. adult © appendages. — 7-8, lateral views of left fore tibia and proximal tarsomere: 7, tiaratum, 8,
carlbergi sp. n. (holotype). 9-13, lateral views of left hind femur, tibia and proximal tarsomere: 9, tiaratum; 10, popa; 11, bu-
fonium; 12, carlbergi sp. n. (holotype); 13, carlbergi sp. n. (paratype from Tari, PNG). 14 - 15, ventral views of left cercus: 14,

carlbergi sp. n. (holotype); 15, tiaratum.

119

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 16.
Sketch map
of Papua New
Guinea and
eastern Irian
Jaya, showing

© E. carlbergi OS

a À. popa ==

known distri-
butions of E.
popa (13
records) and
E. carlbergi
(11 records).

PAS AURA:
NEW
GAU RENDER

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sis adapted to different habitat types. Once evolved
this would tend to restrict the species to the habitat in
which it is best concealed from predators.

The body coloration of £. popa is primarily brown
and when the insect is hanging under a branch at rest
the foliose expansions of the legs (fig. 10) and abdo-
men give it a resemblance to a bunch of dried leaves.
In the lower montane rain forests, where this species
is found, dry brown leaves are presumably more com-
mon than in the wetter mid-montane forests occu-
pied by £. carlbergi. In these latter forests, the bark of
trees and other plants is covered by dense growths of
lichens, and mosses are three times as abundant as in
the lower montane (Johns 1982). Thus the dark
green, white and black mottling of £. carlbergi, to-
gether with the arcuate flanges of the tibiae (figs. 8,
12) and small lateral expansions of the tergites (fig. 2),
are probably adaptations for resembling foliose epi-
phytic lichens, such as the abundant Usnea, or leaves
covered with epiphyllous growth.

The crypsis of the Australian Extatosoma mirrors
that of the New Guinean species, with one, £. tiara-
tum, a leaf mimic (Brock 1992) and the other, £. bu-
fonium, a probable lichen mimic. Both taxa are re-
stricted to the coastal forests of eastern Queensland
and New South Wales (NSW), although the former
species has doubtfully been recorded from Lord
Howe Island (Redtenbacher 1908, Giinther 1931).
E. tiaratum is distributed from Daintree in north-east

120

Queensland (data from a final instar female nymph in
BMNH) to Kiama, near Sydney, NSW (Froggatt 1922)
and appears to range further inland than Æ bufonium
(Beccaloni, unpublished). It is possible that literature
records of ‘E. tiaratum from Brisbane southward rep-
resent the undescribed taxon referred to earlier. E.
bufonium ranges from Moreton Bay, near Brisbane,
south-east Queensland (data from a female nymph in
BMNH) to Camden, near Sydney, NSW (Froggatt,
loc. cit.). This species may also occur in north-east
Queensland, as a female nymph probably belonging
to this taxon was collected in Atherton in 1989 (P. D.
Brock, pers. comm.). A patchwork of different forest
types (ranging from rain forest to dry eucalypt wood-
land) is found in eastern Queensland and NSW and
without further detailed study it is not possible to de-
termine in which habitats the Australian Extatosoma
live. However, considering the types of crypsis exhib-
ited by these species, it is hypothesised that E. bufoni-
um will be found to inhabit a habitat type with a
higher representation of lichens and other epiphytes
than that in which £. tiaratum occurs.

COMPARATIVE MORPHOLOGY AND PHYLOGENETIC
RELATIONSHIPS

The characters discussed below are taken from
adult females and eggs, as only males of E. popa and
E. tiaratum are known and they do not exhibit suffi-

cient characters to make comparisons worthwhile.

Both £. popa and E. tiaratum range from light to
dark brown in colour, although the latter species is
known to have a green form which is not uncommon.
The dorsal foliose expansions of the femur and tibia
of the legs (figs. 7, 9-10) are broad and well developed
in both species, as are the lateral expansions of the
fifth to seventh tergites (as in fig. 6). The lateral ex-
pansions of tergites two to four and eight to nine are
reduced (£. popa) or obsolete (E. tzaratum).

The coloration of E. carlbergi and E. bufonium is
again similar: both are green, with white and black
mottling in varying proportions. The dorsal expan-
sions of the tibiae (figs. 8, 11-13) are arcuate in both
taxa and the lateral expansions of the fifth to seventh
tergites (as in figs. 2, 5) are much reduced relative to
those of E. popa and E. tiaratum. Lateral expansions
of tergites two to four and eight to nine are relatively
well developed.

The superficial similarities in coloration and body
form (listed above) between £. popa and E. tiaratum
and between £. carlbergi and E. bufonium, are prob-
ably due to convergence upon analogous cryptic mor-
phologies and thus do not reflect the phylogenetic re-
lationships of the group. Characters not obviously
related to crypsis (see below) are incongruent with
those characters involved in crypsis and suggest a
grouping reflecting the biogeographical division of
the taxa (New Guinea / Australia).

The New Guinean taxa share a number of features
not possessed by the Australian taxa. Both are larger
than the Australian species and the body spination is
relatively longer and more robust. Both possess a dis-
tinctive pale v-shaped marking on the mesonotum,
and the pair of medial lamellate spines on the posteri-
or third of the meso- and metanotum are large and
well developed. The paired medial lamellae of the
fifth and sixth terga each occupy about one-third of
the length of the segment and their basal width is less
than their height (as in fig. 5). The posterior margin
of the tenth tergite has large posteriorly directed
spines (as in fig. 5) and the apex of the cercus is bi-
lobed (as in fig. 14). The ventral expansion of the fore
tibia has a denticulate margin (as in fig. 8) and the in-
ner ventral margin of the median and hind femur is
expanded ventrally towards its distal end (figs. 10, 12-
13). The egg of E. popa (fig. 100 in Sellick, 1980) is
virtually identical in appearance and dimensions to
that of E. carlbergi (figs. 3-4).

The Australian taxa share the following characters:
the pale mesonotal v-shaped marking (of E. popa and
E. carlbergi) is absent in the adults (although present
in newly hatched nymphs of £. tiaratum) and the pair
of medial spines on the posterior third of the meso-
and metanotum are poorly developed and, if double,
are only weakly confluent basally. The paired medial

BECCALONI: A new species of Extatosoma

lamellae of the fifth and sixth tergites each occupy
about one half of the segment and their basal width is
greater than their height (as in fig. 6). The posterior
margin of the tenth tergite has small posteriorly di-
rected spines (as in fig. 6), which may be subobsolete,
and the cercus is tapered towards its apex (as in fig.
15). The margin of the ventral expansion of the fore
tibia is entire (as in fig. 7) and the inner ventral mar-
gin of the median and hind femur is only weakly (or
not) expanded ventrally towards its distal end (figs. 9,
11). The eggs of E. tiaratum (fig. 8 in Clark, 1976a)
and E. bufonium (pers. observation) share a number
of characters not possessed by eggs of the New
Guinean taxa. Eggs of both species lack micropylar
arms and the opercula are only c. half the length (cor-
rected for differences in egg size) of E. popa | E. carl-
bergi eggs. In addition, the chorionic membrane on
the opercula of eggs of the Australian species is thick-
ened and modified into an opaque hollow dome or
capitulum. The chorionic membrane on the opercula
of eggs of the New Guinean species is not modified
into a capitulum, and instead is transparent and lies
in contact with the surface of the operculum. (Note
that the capitulum of £. tiaratum is relatively much
larger than that of £. bufonium. Also note that the egg
of E. bufoniumis c. half the size of that of £. tiaratum,
the capitulum is dark brown in colour and the egg
capsule has a pitted and irregular surface. The capitu-
lum of £. tiaratum is cream in colour and the egg cap-
sule is smooth and glossy like the capsules of the New
Guinean taxa).

It has not been possible to determine the polarity of
the above characters, as appropriate outgroups to
Extatosoma have not been identified. The other gene-
ra (e.g. Podacanthus Gray, Tropidoderus Gray,
Didymuria Kirby) placed with Extatosoma in the tribe
Tropidoderini, differ greatly from Extatosoma in both
adult and egg morphology preventing useful compar-
ison. However, the non-polar characters described
above imply that either the two New Guinean species
are sister taxa and/or the Australian species are sister
taxa. This ambiguity can only be resolved through
polarity determination.

Key to the adult females of Extatosoma

1. Mesonotum with conspicuous pale v-shaped
marking; ventral expansion of fore tibia with den-
tieulatermazsın (asin fig 6). D

— Mesonotum lacking a pale v-shaped marking;
ventral expansion of fore tibia with entire margin
(ASUS) ot ann, 3

2. Lateral expansions of tergites 5-7 small, non-
overlapping (fig. 2); dorsal expansions of median
and hind femur arcuate (figs. 12-13); body color-
ation pale green, with white and black mottling

121

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

pied seen che) tas MOD ba A aed ia carlbergi Beccaloni
— Lateral expansions of tergites 5-7 large and over-
lapping; dorsal expansions of median and hind
femur broad, not arcuate (fig. 10); body colora-
tion light to dark brown ....................... popa Stal
3. Tergites 2-4 and 8-10 bearing lateral expansions;
dorsal expansions of median and hind femur ar-
cuate (fig. 11); body coloration green, with dark-
er green, white and black mottling......................
ART MA RIST EEN Ci bufonium Westwood
— Tergites 2-4 and 8-10 without lateral expansions;
dorsal expansions of median and hind femur
broad, not arcuate (fig. 9); body coloration
brommtongmeen MINO Ein otter

ACKNOWLEDGEMENTS

This paper is based on the author’s final year BSc (Hons)
project, written while an undergraduate in Zoology at
Imperial College (University of London). The author would
like to thank Mrs J. Marshall and Mr J. Reynolds for assi-
stance, advice and for enabling him to conduct much of his
work in the BMNH. He is grateful to the following individu-
als for either sending him specimens or enabling him to exa-
mine material included in this paper: Mr S. Fellenberg
(Sydney, Australia); Dr M. Hopkins (UPNG); Prof. R.
Kumar (Department of Agriculture and Livestock, Port
Moresby, PNG); Dr G. McGavin (Hope Dept. of
Entomology, Oxford, UK); Dr G. B. Monteith
(Queensland Museum, Brisbane, Australia); Dr M. S.
Moulds (Australian Museum, Sydney, Australia); Dr H.
Roberts (Forest Research Institute, Lae, PNG); Dr H. A.
Rose (University of Sydney, Australia); Mr J. Somp (WEIC);
and Mr J. S. Strazanac (BPBM).

The author’s thanks also go to Mr P. D. Brock and Mr A.
J. E. Harman (UK) for providing useful information, to Dr
U. Carlberg (Sweden) for his enthusiasm and help, and to
Jerry and Willie of Wau (PNG) for collecting the holotype
of E. carlbergi.

The author is also very grateful to Mr P. E. Bragg (UK),
Dr V. K. Brown (Imperial College, UK), Mr R. I. Vane-
Wright (BMNH) and Dr M. Wilkinson (BMNH) for reading
and commenting on the manuscript and he is indebted to all
the organisations and individuals who approved and spon-
sored the ‘Imperial College Expedition to Papua New
Guinea 1990’.

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Brock, P. D., 1992. Rearing and studying stick and leaf in-
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Carlberg, U., 1981. Defensive behaviour in adult female
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Carlberg, U., 1983. Bibliography of Phasmida (Insecta) I.
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Froggatt, W. W., 1922. Description of a new phasma be-
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Linnean Society of New South Wales 47: 344-345.

Giinther, K., 1929. Die Phasmoiden der deutschen Kaiserin
Augusta-Fluss-Expedition 1912-13. — Mitteilung aus
dem Zoologisches Museum in Berlin 14: 597-746.

Günther, K., 1931. Beiträge zur Systematik und Geschichte
der Phasmoidenfauna Ozeaniens. — Mitteilung aus dem
Zoologisches Museum in Berlin 17: 753-835.

Gurney, A. B., 1947. Notes on some remarkable
Australasian walkingsticks, including a synopsis of the ge-
nus Extatosoma (Orthoptera: Phasmatidae). — Annals of
the Entomological Society of America 40: 373-396.

Harman, A., 1985. The genus Extatosoma Gray. — Phasmid
Study Group Newsletter 25: 5.

Hughes, L. & M. Westoby, 1992. Capitula on stick insect
eggs and elaiosomes on seeds: convergent adaptations for
burial by ants. — Functional Ecology 6: 642-648.

Johns, R. J., 1982. Plant zonation. p. 309-330. — In: J. L.
Gressitt (Ed.), Biogeography and Ecology of New
Guinea. Junk, The Hague.

Redtenbacher, J., 1908. p. 380-381. — Die Insektenfamilie
der Phasmiden 3. Verlag Von Wilhelm Engelmann,
Leipzig.

Sellick, J. T. C., 1980. A study of the eggs of the insect or-
der Phasmida with particular reference to the taxonomic
value of egg structure in this group. — Unpublished Ph.
D. thesis, University of London, England.

Verdcourt, B., 1979. — A manual of New Guinea Legumes.
Kristen Press Inc., Madang, PNG, 645 pp.

Vickery, V. R., 1983. Catalogue of Australian stick insects
(Phasmida, Phasmatodea, Phasmatoptera, or
Cheleutoptera). — CSIRO Australia, Division of
Entomology Technical Paper 20: 1-19.

Wilson, S. K., 1991. Throwing your babies at the enemy. —
GEO Australasia’s Geographical Magazine 12: 106-113.

Received: 25 May 1993
Accepted: 30 August 1993

APPENDIX

The following records of Extatosoma popa were used to
plot fig. 16. The data have been taken from material exam-
ined by the author, unless stated otherwise. IRIAN JAYA:
16, Hollandia, 3°10’S 140°E, 300-600 m, i.1937-8, W.
Stüber (BMNH). PAPUA NEW GUINEA: Madang Prov.: 19,
Finisterre Mts., Damanti, 1065 m, 2-11.x.1964, M. E.
Bacchus (BMNH); 19, Finisterre Mts., Budemu, c. 1200 m,
15-24.x.1964, M. E. Bacchus (BMNH); Morobe Prov.: 22,
Wantoat, 1200 m, Casuarina, x.1957, R. W. Paine (BMNH);

BECCALONI: A new species of Extatosoma

29, Wau, golf course, 1150 m, ex Calliandra surinamensis
Benth., 13.vii.1990, G. W. Beccaloni (BMNH); 19
(nymph), Wau, c. 1200 m, ex Hibiscus, 1.vii.1990, G. W.
Beccaloni (BMNH); 1 ® (nymph), Wau, 1150 m, 24.vi.1974,
Gewise Otaweto (WEIC); 16, Wau, 1180 m, 18.x.1974, A.
D. Hart (WEIC); 16, Wau, 1150 m, coll’d on Diospyros fer-
rea, 6.x11.1975 (WEIC); 12, Bulolo, v.1990 (BMNH);
Northern Prov.: 12, Mt. Obree, 1.x.1921, R. Neill (BMNH).

The following records were taken from Gurney (1947):
Papua NEW GUINEA: Morobe Prov.: Sattelberg, vic.
Finschafen; Northern Prov.: Buna.

125

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

(FIORENZA.

XX INTERNATIONAL CONGRESS
OF ENTOMOLOGY

Florence (Italy)
August 25 - 31, 1996

Palazzo dei Congressi

First Circular

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— Opening Plenary Lecture

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BE UE Gt GE BE de GE GE LE GE AE LE GE EE LE AE Gt WE GE e at dè

124

DK) D

CONGRESS PRESIDENT
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see page 288

THOMAS W. DONNELLY

Binghamton, N. Y., U. S. A.

TWO NEW GENERA OF ISOSTICTID DAMSELFLIES
FROM NEW BRITAIN, BOUGAINVILLE, AND THE
SOLOMON ISLANDS (ODONATA: ZYGOPTERA)

Donnelly, T. W., 1993. Two new genera of isostictid damselflies from New Britain,
Bougainville, and the Solomon Islands (Odonata: Zygoptera). — Tijdschrift voor Entomologie
136: 125-132, figs. 1-23, tab. 1. [ISSN 0040-7496]. Published 10 December 1993.

Titanosticta and Cnemisticta are new genera of the Isostictidae described from the eastern
Papuan - Melanesian region. New species Titanosticta macrogaster (New Britain), Cnemisticta
angustilobata (New Britain), and C. latilobata (Bougainville, Gizo, Malaita, and Guadalcanal)
are also described. The two genera bear no particularly close relationship with any of the
Papuan (Selysioneura, Tanymecosticta), New Caledonia (Jsosticta), or Australian (seven genera)

taxa.

Dr. Thomas W. Donnelly, 2091 Partridge Lane, Binghamton NY 13903 USA.

Key words. — Odonata, New Guinea, Solomon Islands, new genera, new species.

The damselfly family Isostictidae is reported from
the Australian subcontinent, the Papuan region (ex-
tending westward to Tanimbar and Halmahera), and
the island of New Caledonia, where the type genus
occurs. In Australia, Watson et al. (1991) recognize
seven genera. Tanymecosticta is widespread on New
Guinea, has a single species in New Britain and an-
other on Woodlark on the east, and occurs on Misool
and Tanimbar on the west. Selysioneura is widespread
on New Guinea, and occurs on Misool, Halmahera,
and Morotai on the west, and Woodlark,
D’Entrecasteaux, and the Louisiade Archipelago on
the east. I herein describe two new Papuan genera:
Titanosticta and Cnemisticta, which occur in New
Britain and the Solomon Islands. The new genera
overlap only slightly in range with either of the two
Papuan genera: thus, they fill a geographic gap in the
range of the family.

The Isostictidae had long been considered a sub-
family of the Protoneuridae, but has recently been
separated into a separate family, largely on the basis of
larval characters (Lieftinck 1975). Adults resemble
the protoneurids, and differ from most otherwise
similar Oriental Coenagrionoid families by their nar-
row wings, with the anal portion highly reduced.
They differ from the Protoneuridae by their absence
of bright colours (protoneurid adults are almost in-
variably marked with bright colours, especially in the
males), their generally longer CuP, and larvae (as far
as known) with saccoid, rather than flat, caudal la-
mellae with subapical constriction. Some genera of

Oriental protoneurids have a very short anal vein; this
vein is always totally lacking in isostictids. Another
character of most genera of the isostictids is a promi-
nently and elaborately developed hind lobe of the
prothorax; Selysioneura and one species of Cnemisticta
are Conspicuous exceptions.

Fraser (1955) discussed this group (which he con-
sidered a subfamily of the Protoneuridae) and sug-
gested a relationship with the platycnemidids. In
common with the Protoneuridae, but conspicuously
different from the Platycnemididae, the Isostictidae
have short legs with short leg spines. However
Cnemisticta has a broader wing which is abruptly nar-
rowed distal to the pterostigma and a relatively long
CuP, both of which are reminiscent of the
Platycnemididae. The larvae of the Platycnemididae
have highly variable caudal lamellae and do not seem
to form a single coherent group.

In the absence of larval forms for most genera it is
not profitable at this time to suggest relationships
within the family. The two new genera do not seem
to have an especially close affinity with any other
Papuan or Australian genus. Cremisticta is notably
distinct within the family because of its broader wing
with abrupt terminal narrowing, and long CuP.

Abbreviations for collections in which type materi-
al is located: UZMC — Universitetets Zoologiske
Museum, Copenhagen; RMNH — Rijksmuseum van
Natuurlijke Historie, Leiden; TD — Collection of T.
Donnelly, Binghamton.

125

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-2. Venation of hind
wings. — 1, Cnemisticta an-
gustilobata, gen. n., sp. n. 2,
Titanosticta macrogaster, gen.
n., sp. n.

Titanosticta gen. n.

(figs. 2, 3-7)

This is the largest damselfly in the family
Isostictidae and one of the longest Zygoptera in the
Oriental region. It is distinguished by the short CuP,
which is two cells long (rarely one). Also, Ac is just
proximal to Ax1 in the fore wing, and slightly distal
in the hind wing; the arculus is distal to the Ax2 of
both wings; IR2 arises at the 7-8th Px of the fore wing
and at 6 or 7th (rarely 5th) in the hind wing; R4+5
arises distal to the subnodus. The pterostigma is
rhombic and longer than broad. The wing is nar-
rowed gradually distal to the pterostigma. The inferi-
or appendage of the sole species is much larger than
the superior. In common with other genera of the
family, the legs are short with short spines, and the
body colours are dull.

Type species: T. macrogaster sp. n.; the only known
locality is New Britain.

Etymology: ‘Titan’, from mythology, signifying
very large size, and ‘-sticta’, the predominant suffix
for genera in this family.

In addition to its very large size, Titanosticta is dis-
tinguished from all other isostictid genera by the fol-
lowing combination of characters: inferior appendage
much larger than the superior; CuP two cells long,
vein R4+5 arising distal to subnodus, very small tarsal
claws, and Ac close to Axl in the fore wing and slight-
ly distal to Axl in the hind wing.

Titanosticta macrogaster sp.n.

(figs. 2, 3-7)

Type material. — Holotype d : Papua New Guinea. New
Britain. Yalom, 1000 m, 20 May 1962. Noona Dan Exped.
(UZMC). Allotype 9: Papua New Guinea. New Britain.
Metelen River, 20 mi. S.E. of Ruango, July 1970, V.
Jindrich (RMNH). Paratypes: 2 d same locality as holotype,
19-20 May 1970 (RMNH, TD); 18 Papua New Guinea. New
Britain. Metelen River, 20 mi. S.E. of Ruango, July, 1970,

126

V. Jindrich (rp).

Additional material. — An additional 2 in the Donnelly
collection, with the same data as Metelen River paratype, is
not designated a paratype because it lacks both a head and
the terminal segments of the abdomen.

Male holotype. — This is an obscurely marked spe-
cies varying from pale gray brown to dark, shining
brown.

Head: Labium pale with rounded triangular central
excavation about 2/5 the length of the segment. Face
mainly pale; labrum shining black with thin apical
yellow rim and pale central, basal spot; postclypeus
pale with prominent black anterior and lateral mar-
gins; frons pale with small black spots anterior and
posterior to antennae and irregular black central
stripe enclosing front of lateral ocelli and median
ocellus; pedicel about 1.5 times the length of the
scape, both pale.

Prothorax: Pale, small paired central black spots on
hind lobe and very small black marks on remainder
pronotum; hind lobe sharply recumbent, broadly T-
shaped, with a basal constriction.

Pterothorax: Mesostigmal laminae small, slightly
tapered apically, with rounded tips; pterothorax gray-
ish green; mesepisternum with stripe on dorsal carina
occupying 1/6 of sclerite; very thin dark line on ante-
humeral suture; 2nd lateral suture with small dark
spot on caudal sixth.

Legs: Yellow with contrasting dark spines: 4 on
outer row of hind femur, 5 on hind tibia.

Wings: Venation described in generic diagnosis;
venation brown, rhombic, pale red brown.

Abdomen: Pale, darker at tip; 1 and 2 greenish
gray; 3 to 8 yellow, darkening apically; 9 and 10 shin-
ing dark brown. Superior appendage dark, shorter
than 10, constricted apically to terminate in rounded,
cylindrical tip, with low, rounded basal-ventral prom-
inence. Inferior appendage 1.5 times length of super-
ior, forcipate, laterally flattened, expanded abruptly
in caudal half, excavate on dorsal-apical corner to

DonneLLy: /sostictid damselflies

Table 1. Characters of genera of Isostictidae. — Explanation: CuP: Number of cells length; both wings considered. Inferior
appendage: of male, whether large or smaller than superior appendage. Hind lobe of prothorax, male: whether prominent
(flared, with horns, or with other processes) or low. Ac or Axl: which crossvein is in more distal position in hind wing. Tarsal

claws: size compared with other coenagrionoids. R4+5: origin of vein with respect to nodus.

Genus Range length of CuP, male hind lobe tarsal R4+5 hw: Ac or Axl
cells inferior prothorax claws to nodus distal
appendages male
Isosticta New Caledonia 1-6 equal prominent normal distal, Axl; some spp.
proximal, nearly aligned
or aligned
Selysioneura New Guinea, 0-1 smaller low very small distal Ax (nearly
Misool, aligned in
Halmahera, some spp)
Morotai,
Woodlark,
D’Entrecasteau,
Louisiade Arch.
Tanymecosticta New Guinea, 1 slightly prominent small distal Ax
New Britain, smaller
Woodlark,
Misool,
Tanimbar
Titanosticta New Britain 2 (1) larger prominent very small distal Ax
Cnemisticta New Britain, (7)8-10(11) larger prominent normal proximal Ac
Bougainville, or low
Solomon I.
Rhadinosticta Australia 1 larger prominent normal proximal Axl
Oristicta Australia 1 smaller prominent very small distal Ac
Neosticta Australia 5-6 smaller prominent normal proximal Ac (close)
Labidiosticta Australia 0-2 smaller prominent very small distal Axl
Eurysticta Australia 122 smaller prominent normal proximal Axl
Lithosticta Australia 5-7 equal prominent normal proximal Axl
Austrosticta Australia 4-6 larger prominent normal proximal Ac (close)

form two rounded tips. Penis with terminal segment

Y-shaped.

Female allotype. — Similar to male, but face more
pale: labrum yellow with obscure paired lateral spots;
anteclypeus mottled brown; postclypeus pale with
brown anterior and lateral margins, this dark colour
extended posteriorly in center and at lateral extremes;
top of head gray-green, black as follows: short trans-
verse dashes anterior to median ocellus; jagged trans-
verse stripe enveloping median ocellus extending pos-
terior to lateral ocelli.

Thorax: Hind lobe of prothorax quadrate, scarcely
narrowed at base, the central portion of hind rim ex-
cavate in the center with a medial, rounded knob.
Colour of side of thorax greenish yellow. Basal seg-
ments of abdomen as in male; 5 terminal abdominal
segments lacking.

Dimensions. — Holotype male: abdomen 60.5
mm, hind wing 32 mm. Allotype female, hind wings
dissimilar, 29.5 and 31 mm.

The dimensional range of the paratype series is as
follows: two males have abdomens 46 and 63 mm,

and hind wings 32-32.5 mm. The 2nd female has the
hind wing 30 mm.

Variation within paratype series. — The three males
have notably different abdomen lengths. The
Metelen River male has the abdomen (46 mm) more
than a centimeter shorter than the Yalom males (60.5
and 63 mm), but the hind wing of this specimen (29
mm) is only slightly shorter than the other two (32
mm). There are only trivial differences among the
three males in colour pattern. One male has a broad-
er black stripe on mesepisternum (4 rather than % the
width of the sclerite). The females, which are both re-
grettably incomplete, appear essentially identical.

Etymology. — The name, a noun in apposition, re-
fers to the very long abdomen.

Remarks. — The difference in abdomen length of
the males (which have very similar hind wing lengths)
violates the normally firm linear allometric relation-
ship (in this case the ratio between abdomen and hind
wing) among Zygoptera. The constancy of this ratio
can be attributed to the necessity of maintaining a
uniform dimensional ratio to enable the insect to fly
vigorously. There are similar disparities in this di-

127

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 3-7. Titanosticta macrogaster, gen. n., sp. n. — 3, male hind lobe of prothorax, dorsal view. 4, female hind lobe of pro-
thorax, dorsal view. 5, male abdominal appendages, dorsal - lateral - apical view. 6, male abdominal appendages, lateral view.

7, penis, ventral and lateral view.

mensional ratio among New World Pseudostigmat-
idae, which have extremely long abdomens and which
fly much less vigorously than other Zygoptera. The
violation of a constant ratio in the few specimens of
Titanosticta raises the question as to whether the
flight of this genus (which has not been recorded)
might resemble the weak flight of the pseudostigmat-
ids.

Cnemisticta gen. n.

(figs. 3, 8-23)

This genus contains two species of large isostictids.
It is immediately recognized by the length of CuP,
which in the type species C. angustilobata is 8 to 10
cells long in the fore wing) and 8 (rarely 7) to 10
(rarely 11) in the hind wing. IR2 arises at the 7-8th
Px in the fore wing (rarely 9th) and 5 to 6th in the
hind wing. In C. latilobata IR2 arises at the 5-6th
(rarely 7th) Px in the fore wing and 4-5th Px in the
hind wing; CuP is 8-10 cells long in the fore wing and
7-9 in the hind wing. In both species vein Ac arises
just distal to the Ax1 of the fore wing and %o the dis-
tance between the Ist and Ax2 in the hind wing; the
arculus is distal to the Ax2 in both wings; R4+5 arises
proximal to the subnodus in both wings. The pteros-
tigma is rhombic and as wide as long. The tarsal claws
are of normal length and the inferior abdominal ap-

128

pendage of the male is much larger than the superior.
The two species range from New Britain to Bougain-
ville, Gizo, Malaita, and Guadalcanal.

Type species. — C. angustilobata sp. n., from New
Britain.

Etymology. — ‘Cnem-’ from the slight resemblance
to some platycnemidids.

This genus is distinguished from all other isostic-
tids by its much longer CuP and abrupt narrowing of
the wing beyond the pterostigma. The two species
vary from each other in the extent of development of
the hind lobe of the prothorax. The species C. latilob-
ata has a very prominent hind lobe, similar to that of
T. macrogaster, and to most of the other members of
the family. The species C. angustilobata has a highly
reduced hind lobe, in common with the most wide-
spread Papuan isostictid, Selysioneura.

Cnemisticta angustilobata sp. n.
(figs. 1, 8-15)

Type material. — Holotype d : Papua New Guinea. New
Britain: Yalom, 1000 m, 18 May 1962, Noona Dan Exped.

_(UZMC). Allotype 9: same data as holotype, except coll. 19

May 1962 (UZMC). Paratypes: 3d 19 same data as holotype
except coll. 18-19 May 1962 (RMNH); Papua New Guinea.
New Britain. Metelen River, 20 mi. S.E. of Ruango, July
1970, V. Jindrich (rp); 1 & Papua New Guinea. New

Britain: Gazelle Pen., upper Warango, 250-600 m, 28-30
Nov 1962, J. Sedlacek (RMNH).

Additional material. — 4 additional d collected by
Sedlacek (RMNH) along with last named paratype are not
designated as paratypes because they lack the terminal seg-
ments of the abdomen.

Description

Male holotype. — This is a dark, obscurely marked
damselfly.

Head: labium pale with rounded excavation in ap-
ical 4; frons rounded; face and top of head black; la-
brum black with pale, obscurely defined central
stripe; genae pale, anteclypeus obscurely pale; pedicel
slightly longer than scape, both dark; rear of head
pale.

Prothorax: Pale laterally, mottled dark on dorsum,
with large tringular paired dark spots on median
lobes; apodyne extended posteriorly as commissure
nearly to hind lobe; hind lobe small and low, round-
ed triangular with central part slightly thickened.

Pterothorax: Mesostigmal laminae simple, parallel
sided, rounded, slightly elevated lateral tip; mesepi-
sternum black, mesepimeron dark with pale on mesal
% (adjacent to antehumeral suture) and cephalic %;
mesinfraepisternum pale on outside, central half
dark; the dark colour of the mesepisternum essential-
ly extensive to metepisternum with only thin pale
lines on the two lateral sutures; metepisternum pale
with black spot on caudal %; metepipimeron and
metastermum pale.

Legs: Pale with thin dark dorsal lines on femora;
spines brown, not contrasting with legs, 5 on hind fe-
mur and hind tibia.

Wings: Venation as for genus; IR2 arising at 7 (fore
wing) and 5 (hind wing); CuP 9 cells long in fore
wing and hind wing; pterostigma rounded, rhombic,
as long as wide, pale reddish brown.

Abdomen: Pale, dark on dorsum of 1 and 2, ob-
scurely dark on dorsum of 3 to 7, with apical % dis-
tinctly darker on 3 to 6; 7 uniform; 8 with obscurely
dark dorsal mark in caudal #; 10 with obscure paired
dark spots on caudal 4. Superior appendage shorter
than 10, blunt, rounded, slightly expanded and
branched in dorsal view, with mesal branch very
short; with ventrally directed spine. Inferior append-
age more than twice as long than superior, forcipate,
laterally flattened, with caudal half expanded into
prominent rounded keel on dorsal and apical por-
tions. Penis with terminal segment Y-shaped.

Allotype female. — Similar to male. Face pale; post-
clypeus pale with dark lateral margins; top of head
pale, about half dark, with an irregular dark mark
centered on the ocelli, small dark spots anterior to an-
tennae, and short transverse dark stripes posterior to
lateral ocelli. Thorax: Hind lobe of prothorax similar

DonneELLY: Isostictid damselflies

to male but slightly larger, low triangular, with a
stepped appearance. Colour of thorax: mesepister-
num with pale stripe above antehumeral suture % the
width and in the cephalic % of the sclerite; mesepime-
ron with short central elongate dark mark;
Metepisternum with small caudal dark mark.
Abdomen: Dorsum of 8 to 10 pale; ovipositor dark,
extending distinctly beyond 10 (cerci).

Dimensions. — Holotype male: abdomen 44 mm,
hind wing 29 mm. Allotype female: abdomen 45.5
mm. The dimensional range of the paratype series is
as follows: 5 paratype males have the abdomens 43.5
to 47.5 mm; 8 males have the hind wing 26 to 29
mm. A paratype female has the abdomen 45.5 mm
and the hind wing 32.

Variation within paratype series. — The Gazelle
Peninsula and Metelen River males are paler than the
Yalom series; the dark colour of the side of the thorax
ranges to a short central dark mark on mesepimeron
and metepisternum almost totally pale.

Etymology. — The name refers to the vary narrow
hind lobe of the prothorax.

Cnemisticta latilobata sp. n.
(figs. 16-23)

Type material. — Holotype d : Solomon Islands. Malaita
I., Kware’a River and tributaries, 8 km E of Dala, 100m, 19
Feb 1987, T. Donnelly (RMNH); Allotype 2 Solomon
Islands. Guadalcanal I., Honiara, Rove Creek at Reservoir,
13-14 Feb 1987, T. Donnelly (RMNH). — Paratypes: 1d
same data as holotype (TD), 1d, Papua New Guinea,
Bougainville I., South Nasiooi, Agriculture Field Station, 19
May 1975, Howard R. Wimmer (RMNH).

Additional material. — 16 Solomon Islands. Gizo I., low
bush area in valley, 7,8 May 1975, H. Wimmer (specimen
lacks terminal segments of abdomen) (RMNH), 1d Papua
New Guinea. Bougainville I., Boku Bong, 5 June 1956, J. L.
Gressitt (RMNH) (highly teneral specimen).

Description

Holotype male. — A pale, obscurely mottled, dam-
selfly.

Head: Labium white, with a quadrate apical exca-
vation % of its length; frons rounded; face black, la-
brum with thin yellow apical rim; genae brown with
yellow mesal margin; pale pedicel slightly longer than
dark scape; top of head black bordered at rear by yel-
low; obscure pale spots lateral to lateral ocelli; rear of
head yellow.

Prothorax: pale, fore lobe with dark rim; medial
lobes low, apodyne continued into a commissure
nearly to hind lobe; hind lobe large, T-shaped, sharp-
ly recumbent, basally constricted, and sharply notch
in center of hind rim.

129

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 8 - 15. Cnemisticta angustilobata, gen. n., sp. n. — 8, male hind lobe of prothorax, dorsal view. 9, male hind lobe of pro-
thorax and mesostigmal lamina, dorsal- lateral view. 10, female hind lobe of prothorax, dorsal view. 11, female hind lobe of
prothorax and mesostigmal lamina, dorsal- lateral view. 12, male abdominal appendages, dorsal - lateral - apical view. 13,
male abdominal appendages, lateral view. 14, ovipositor. 15, penis, ventral and lateral views.

Pterothorax: Mesostigmal laminae parallel sided,
flat, with rounded tips; colour of pterothorax pale;
mesepisternum with dark stripe on dorsal carina oc-
cupying % of sclerite; antehumeral suture with thin
black line; 2nd lateral suture with tiny black spot on
caudal %. Legs: Yellow, with 4 contrasting black
spines (outer row) on hind femur, 5 on hind tibia.
Wings: Venation as in generic diagnosis; IR2 arising
at 6 (fore wing) and 5 (hind wing); CiP 8 (fore wing),
9 (hind wing) cells long.

Abdomen: Pale laterally and darker dorsally; 3 and
4 dominantly pale dorsally, slightly darker in basal
half and on apical 4; 5 obscure, darker on apical %; 6
slightly pale in basal %; 7-10 dark, with limited pale
on sides, 8 and 9 with pale longitudinal dorsal lines
on basal %, tapering caudally. Superior appendage
brown, shorter than 10, rounded and expanded api-

130

cally in dorsal view, with rounded, upturned apices.
Inferior apendage 3 times as long as superior, forci-
pate, with blunt, laterally flattened, rounded tip and
dorsal keel on expanded apical half, and with low,
rounded, conical, dorsally directed spine in basal %.
Penis with terminal segment Y-shaped.

Allotype female. — Similar to male. Head: Face
pale, paired dark spots anterior to lateral ocelli. Small
rounded pale protuberances on rear of head. Thorax:
Hind lobe of prothorax prominent, triangular, divid-
ed into three nearly equally wide lobes by deep sulci;
the central lobe emarginate, and the two lateral lobes
low and tapering laterally to points. Colour of thorax:
Mesepisternum pale, thin black line adjacent to pale
dorsal carina; thin black line along caudal half of an-
tehumeral suture; mesepimeron with an obscure cen-

DonnELLy: /sostictid damselflies

Figs. 16-23. Cnemisticta latilobata gen. n., sp. n. — 16, male hind lobe of prothorax, dorsal view. 17, male hind lobe of pro-
thorax and mesostigmal lamina, dorsal-lateral view. 18, female hind lobe of prothorax, dorsal view. 19, female hind lobe of
prothorax and mesostigmal lamina, dorsal- lateral view. 20, male abdominal appendages, dorsal-lateral-apical view. 21, male
abdominal appendages, lateral view. 22, ovipositor. 23, penis, ventral and lateral views.

tral dark mark deflected laterally in cephalic portion.
Abdomen: 9 and 10 with prominent pale basal-later-
al spots; ovipositor pale, extends to end of 10 (cerci).

Dimensions. — Holotype male: abdomen 39 mm,
hind wing 26 mm. Allotype female: abdomen 35.5
mm, hind wing 26.5 mm. The dimensional range of
the paratype series is as follows: 2 paratype males have
abdomens 37 and 38.5 mm; 3 paratype males have
the hind wing 24 to 26 mm. The 2nd female speci-
men is very teneral; the hind wing is 27 mm.

Variation within paratype series. — The male from
Bougainville has the pale colour of the top of the head
more extensive than the holotype, extending anterior-
ly nearly to the antennal bases. A very teneral female
specimen has the hind lobe of the prothorax longer

and apparently more Y-shaped than the allotype.
However the apparent difference in shape could be
due to twisting upon drying.

Etymology. — The name refers to the wide and dec-
orative hind lobe of the prothorax.

Remarks. — The two species of Cnemisticta resem-
ble each other in their peculiarly wide and apically
abruptly narrowed wings, the long CuP, and the male
abdominal appendages. The species angustilobata is
slightly larger than /atilobata. The species differ most
prominently in the hind lobe of the prothorax, which
is prominently developed in /atilobata and reduced in
angustilobata. The species latilobata is pale, but with a
darker abdomen, and angustilobata is dark, but with a
paler abdomen. The allotype female of latilobata has

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

a rounded prominence on the rear of the head; this is
lacking in angustilobata. The female of angustilobata
has a much longer ovipositor than /atilobata. The ve-
nation of the two species (above) differs more than is
appropriate for the slight dimensional differences
between the two species.

ACKNOWLEDGEMENTS

I am indebted to the RMNH, Leiden, for the loan of
specimens which M. A. Lieftinck had been studying
at the time of his death. I am further very grateful to
Vladimir Jindrich, at that time a graduate student in
our department, for specimens of two of the new spe-
cies collected during a geological trip to New Britain.
Comments from Drs. Jan van Tol and Anthony
Watson on a draft of the manuscript have been very
helpful and are gratefully acknowledged; I am further

132

grateful to Dr. Watson for the loan of specimens of
four Australian genera which enabled me to complete
the table of generic characters.

REFERENCES

Fraser, F. C., 1955. A new isostictine dragonfly from
Australia with some remarks on the subfamily Isostictinae
nov. (Odon., Protoneuridae). — Entomologist’s Monthly
Magazine 91: 227-230.

Lieftinck, M. A., 1975. The dragonflies (Odonata) of New
Caledonia and the Loyalty Islands. Part 1. Imagines. —
Cahiers O. R. S. T. O. M., série Hydrobiologie 9(3):
127-166.

Watson, J. A. L., G. Theischinger, and H. M. Abbey, 1991.
The Australian dragonflies. — CSIRO, Canberra and
Melbourne, 278 p.

Received: 14 April 1993
Accepted: 14 June 1993

MATTI HAMALAINEN

Department of Applied Zoology, University of Helsinki

DESCRIPTION OF NEUROBASIS DAVIESI SP. N. FROM
PALAWAN, WITH TAXONOMIC NOTES ON OTHER
SRECIES OP DEE NCHINENSISGROUP (ODONATA,
EAEOPTERYGIDAE)

Hämäläinen, M. 1993. Description of Neurobasis daviesi sp.n. from Palawan, with taxonomic
notes on other species of the N. chinensis group (Odonata, Calopterygidae). — Tijdschrift voor
Entomologie 136: 133-136, figs 1-4. [ISSN 0040-7496]. Published 10 December 1993.
Neurobasis daviesi sp. n. (holotype male, Brooke’s Point, S. Palawan) is described in both sexes.
The new species is closest to the Bornean N. longipes Hagen, 1887, which is considered to be
specifically distinct from N. chinensis (Linnaeus, 1758). A key for both sexes to the three recog-
nized species of the N. chinensis group is presented.

M. Hämäläinen, P. O. Box 53, FIN-02151 Espoo, Finland.

Key words. — Neurobasis, Palawan; new species; key.

As well pointed out by Lieftinck (1949, p. 13-15),
the definition of the limits of each species in the ge-
nus Neurobasis Selys, 1853 has always been difficult
and subject to considerable personal interpretation.
This applies well also to the present assessment of the
taxonomic status of a new Neurobasis taxon from
Palawan, work on which necessitated a reconsidera-
tion of Lieftinck’s (1940, 1954) view of recognizing
three subspecies in Neurobasis chinensis (Linnaeus,
1758), viz. chinensis, florida Hagen and longipes
Hagen.

In my opinion it is difficult to regard /ongipes and
florida to have an equal status as ‘subspecies of
chinensis. Whereas the Javanese florida comes quite
near to the continental forms, the Bornean longipes
differs markedly from them and represents a distinct
good species.

Neurobasis chinensis (Linnaeus, 1758)

(figs. 1-2).

N. chinensis is a widely distributed insect in South
and South-East Asia. It is known to extend from NE
Pakistan in the west to South China Sea in the east. In
the western part of its range it spreads from NE
Pakistan, Himachal Pradesh, Uttar Pradesh and
Nepal in the north through whole India to Ceylon. In
the eastern part its area extends from Fujien and
Yunnan provinces in China southwards to the
Malaya Peninsula, Sumatra (& Simalur and Nias)
and Java.

In males the breadth of the wings and the relative
size of the metallic area in the hind wing and in fe-
males the presence or absence as well as the size of
pseudopterostigma and the size of the whitish nodal
spots are subject to considerable variation within the
vast range of the species. Documentation of this vari-
ability and revealing its nature is worth of a thorough
study.

Results of this study would also be useful in decid-
ing on the taxonomic status of the Javanese popula-
tions, which indeed differ most from the topotypical
Chinese populations. Lieftinck (1940, 1954) consid-
ered the Javanese populations to represent a distinct
subspecies - florida Hagen, 1854. He also pointed out
the very confusing usage of the name florida in the
early literature and concluded that the original de-
scription of florida applies to the Javanese specimens
only. However, the correct citation to the author of

florida should be Hagen in Walker, 1853.

Neurobasis longipes Hagen, 1887 stat. n.
(fig. 3).

This species is universally distributed in Borneo
and it occurs in lowland streams in forested areas.
The following brief comparison with chinensis ex-
plains my decision to consider longipes as a distinct
species. In longipes
— the size is smaller and the whole appearance is ‘-
slimmer’
— legs are proportionally longer in both sexes; espe-

133

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-4. Wings of males (not in scale). - 1, N. chinensis, North Thailand; 2, N. chinensis (ssp. florida), West Java; 3, N. lon-
gipes, Borneo, E. Sabah; 4, N. daviesi sp. n., Palawan (paratype).

cially the anterior femora are very long, in male as
long and in female longer than the length of the 3rd
abdominal segment (in chinensis much shorter)

— in the male hind wing the opaque colouring is
much more reduced at base (Fig. 3); in subcostal field
starting at the level of ca. 10th antenodal and below at
the level of apical end of quadrangle

— in male the shape of hind wings is different (Fig. 3),
the broadest point is situated usually beyond the mid-
dle of its length; fore wing base with only 2 complete
rows of cells between the lowest sector of IA and the
wing border

— female is differently coloured; metallic green colour
on thorax has a strong coppery tint instead of emerald
metallic green in chinensis. In abdomen the shining
metallic green colour on dorsum is restricted to seg-
ments 1 and 2, the rest being dark brownish; in chi-

134

nensis the dorsum of segments 3 and 4 is similarly
shining metallic green as on segment 2; the metallic
colour getting darker green more apicad

— whitish nodal spots and pseudopterostigma are
lacking in all wings of the female.

The species was originally described by Hagen
(1887) as ‘Abart’ of chinensis on the basis of a single
male specimen from Mindai. Hagen’s contribution
remained unnoticed by other authors, like Kirby
(1890), de Selys Longchamps (1897) and Förster
(1897), who treated the genus in detail. Lieftinck
(1940), who had studied a series of specimens from
different parts of Borneo, reintroduced the name /on-
gipes to apply to all Bornean specimens and presented
a complete list of early references on this taxon.

Neurobasis daviesi sp. n.

(fig. 4).

Type material. — Holotype d: Philippines, Palawan,
Brooke’s Point, (325 m), l.viii. 1980, Leg. R. B. Rodriquez;
in Museum Leiden (RMNH). — Paratypes: 2d, locality and
date as in the holotype (RMNH); 3d, Philippines, S.
Palawan, Brooke’s Point, 1982, Leg. R. B. Rodriquez
(RMNH); 198, 49, Philippines, Palawan, Matalangao River
[between Roxas and Port Barton], 21.xii.1975, [collector
data lost, probably R. B. Rodriquez] (coll. D. A. L. Davies,
Cambridge, U.K. ; 6d, 29 in coll. M. Hämäläinen).

Description

Instead of a detailed description, I prefer to high-
light the characters by comparison of the new species
with N. longipes and N. chinensis. N. daviesi is a some-
what larger insect than chinensis and very distinctly
larger than longipes.

Male. — Colouring of head resembles that of chi-
nensis. However, the yellow colour on labrum is re-
stricted to form two widely separated spots. Thorax
brilliant metallic green as in the related species. Sides
of thorax showing, however, a slight coppery hue.
Legs long; anterior femora proportionally longer than
in chinensis, but shorter than in longipes; shorter than
the length of the 3rd abdominal segment. Abdomen
metallic green with pale markings on sides of seg-
ments 1-2 as in the other species.

Wings. The shape of hind wing is closer to that in
longipes than in chinensis (Fig. 4). Opaque metallic
area in hind wing extends more basad than in /on-
gipes, but base of hind wing distinctly less coloured
than in chinensis. Hind wing border at base as broad-
ly hyaline as in the Javanese specimens of chinensis
(ssp. florida), but the line with the opaque patch much
less distict. As in the other species the hind wing re-
flects brilliant metallic green or blue colour according
to angle of view; the apical third with violaceous re-
flections. The intensity of the metallic reflections is
less pronounced than in chinensis, thus the outer line
of the metallic patch looks less sharp.

Measurements. Fore wing 35.5-37 mm, hind wing

33.5-35 mm; abdomen 48-49 mm.

Female. — Rather robust insect. Thorax less shining
metallic green than in chinensis. The colouring of ab-
domen resembles that of longipes, metallic green
colouring being restricted to dorsum of segment 1
and the basal 4/5th of segment 2. Rest of the abdo-
men brown, darker brown on dorsum of segments 6-
10. Dorsal carina narrowly yellow broadening to a
typical band on segments 8-10.

Legs very long, proportionally longer than in chi-
nensis. Anterior femora as long as the 3rd abdominal
segment. Hind femora extending (in enveloped speci-
mens) beyond half of the 3rd abdominal segment as
in longipes.

HAMALAINEN: Neurobasis daviesi sp. n.

Wings hyaline with yellowish tint; as in longipes,
the hind wings only slightly more heavily tinted than
the fore wings. As in longipes, no trace of pseudopte-
rostigma or whitish nodal spots in fore and hind
wings.

Measurements. Fore wing 38.5-39 mm, hind wing
40-41 mm, abdomen 47.5-48.5 mm.

Etymology. — The new species is named after Dr
D. A. L. Davies (Cambridge, U.K.) to appreciate his
diligent efforts to search for dragonfly rarities in dif-
ferent corners of the globe.

Discussion. — In spite of the striking size difference,
daviesi seems to be nearer to longipes than to chinensis.
This is apparent especially from the female characters:
long legs, absence of nodal spots and pseudopterostig-
ma in all wings and brownish colouring of abdomen.
However, the colouring of thorax is slightly different.
Males show more ‘intermediate’ characters between
chinensis and longipes.

I think that it is better to to treat N. daviesi as a dis-
tinct species, rather than a subspecies of /ongipes. Both
taxa have well defined separate ranges in different is-
lands, although separated only by the narrow Balabac
Strait.

It is remarkable that M. daviesi is missing from the
large dragonfly collection amassed by Roland A.
Miiller and his coworkers from many localities (even
from Matalangao River) in different parts of Palawan
in 1985-1992. N. daviesi is either a very rare species
or (most likely) its flying period is confined to the ad-
vanced rainy season. The bulk of Miiller’s specimens
have been collected at the beginning of the rainy sea-
son in May-June or during the dry season in March.

Key to the species of Neurobasis chinensis group

Males

1. Hind wing broadly ‘hyaline at the base; opaque
patch starts at the level of the apical end of
quadrangle at the central part of the wing. In fore
wing only two complete rows of cells between the
lowest sector of IA and the wing border. Anterior
femora as long as the length of the 3rd abdominal
SESCNENRL: Has ee tone ear senses wale longipes

— Hyaline area in hind wing base less extensive. In
fore wing at least 3 rows of cells between the low-
est sector of IA and the wing border. Anterior fe-
mora shorter than the length of the 3rd abdomi-
DAL SEP IMEN by us Lira ?

2. Athind wing base the whole subcostal and cubi-
tal fields opaque and reflecting strong metallic
colour (only 1-2 basal cells of cubital field hya-
mes RUE PP LEE LP M eae chinensis

— Several hyaline or subhyaline cells in subcostal

135

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Females

1. All wings with (more or less distinct) whitish no-
dal spots. Abdominal segments 3-4 predominant-
ly shining metallic green on dorsum .....chinensis

— No whitish nodal spots present on wings. Ab-
dominal segments 3-4 predominantly brownish
OMdOrsumae A Lee A Aes ne 2

2. Anterior femora clearly longer than the length of
the 3rd abdominal segment. Small species, hind
fm lees nn en longipes

— Anterior femora as long as the length of the 3rd
abdominal segment. Large species, hind wing
OVER SA sum: daviesi

ACKNOWLEDGEMENTS

I am intebted to Drs Jan van Tol (RMNH, Leiden)
and Dr D. A. L. Davies (Cambridge, U.K.) for the
loan of the necessary specimens and for valuable com-
ments on an earlier version of the manuscript. My
thanks are also due to Mr Matti Viitasaari and Mr
Reino Tyynelä for the wing photographs.

136

REFERENCES

Förster, F., 1897. Contributions a la faune odonatologique
Indo-Australe. — Annales de la Societe Entomologique de
Belgique 41: 204-211.

Hagen, H., 1887.Ueber Neurobasis und Vestalis. —
Abhandlungen der Zoologisch-Botanischen Gesellschaft
in Wien 37: 647-648.

Kirby, W. F., 1890. A synonymic catalogue of Neuroptera
Odonata or dragonflies. With an appendix of fossil spe-
cies. — Gurney & Jackson, London.

Lieftinck, M. A., 1940. Descriptions and records of South-
east asiatic Odonata (II). — Treubia 17: 337-390.

Lieftinck, M. A., 1949. The dragonflies (Odonata) of New
Guinea and neighbouring islands. Part VII. Results of the
Third Archbold expedition 1938-1939 and of the Le
Roux Expedition 1939 to Netherlands New Guinea (II.
Zygoptera). — Nova Guinea (N.S.) 5: 1-271.

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 (Suppl.) 22: i-xiii + 1-202.

Selys Longchamps, E. de, 1897. Causeries Odonatologiques
No 10. — Annales de la Societe Entomologique de
Belgique 41: 427-432.

Received 1 September 1993
Revised version accepted 14 October 1993

CHRISTOPH L. HAUSER

Museum Alexander Koenig, Bonn

AeA NO TATED GHECKLIST OF THESPECIES OF
THE PARNASSIINAE (LEPIDOPTERA: PAPILIONIDAE)

Hauser, C. L., 1993. An annotated checklist of the species of the Parnassiinae (Lepidoptera:
Papilionidae). — Tijdschrift voor Entomologie 136: 137-146. [ISSN 0040-7496]. Published 10
December 1993.

A total of 76 binominal taxa including 22 cases of questionable species status are listed for the
Parnassiinae, a subfamily comprising eight genera of predominately palaeartic swallowtail but-
terflies. The taxonomic history of all doubtful species is briefly reviewed and possible affinites
of these taxa are indicated. The majority of taxa contributing to a considerable increase in spe-
cies number compared to previous accounts represent changes in status rather than genuine dis-
coveries of new taxa. This is seen in relation to a change of attitude in taxonomic practise.

C. L. Hauser, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee
160, D-53113 Bonn, Germany.

Key words. - Papilionidae, Parnasiinnae, checklist, species number.

The Parnassiinae represent one of the three cur-
rently recognized subfamilies of the Papilionidae, the
‘Swallowtail Butterflies’. Among the swallowtails, the
‘Apollo Butterflies’ in particular have for long been
highly sought after by butterfly collectors because of
their great aesthetic appeal and considerable intraspe-
cific variation in wing pattern. This specialized inter-
est has led in the past to voluminous works devoted to
catalog and illustrate all the many named species-
group taxa and infraspecific forms of the Parnassiinae
(e.g., Austaut 1889, Verity 1905-1911, Stichel
1907a, 1907b, Rothschild 1918a, 1918b, Bryk 1934,
1935, Eisner 1974, 1976). In contrast, comparatively
little attention has hitherto been paid to species-level
taxonomy, and to morphological or ecological pecu-
larities of the group.

The subfamily is largely restricted to the palaearctic
region with three species occuring in western North
America, and completely absent from tropical zones
(Stichel 1907a, 1907b, Reinig 1937, Ackery 1975).
In an evolutionary view, the Parnassiinae have always
been regarded as a basic stock of the Papilionidae
from which all other higher taxa of the family arose
(e.g., Spuler 1892, Reuter 1897, Ford 1944, Munroe
1961). In phylogenetic terms, the Parnassiinae are
considered accordingly as the sistergroup of the
Papilioninae, which comprise all remaining swallow-
tails, with the exception of one monotypic genus
(Hancock 1983, Miller 1987, Hauser 1993).

Since the last comprehensive review of the
Parnassiinae by Ackery (1975) several species have
been described as new, mostly in the genus Parnassius
(e.g., Koiwaya 1987, Korshunov 1988, Weiss &

Michel 1989, Watanabe 1990). Furthermore, a num-
ber of formerly recognized species has been shown to
consist of two or more closely related, but reproduc-
tively well isolated species, often on the basis of accu-
mulated knowledge on their early stages and biology
(e.g., Larsen 1973, de Freina 1985, Kreuzberg 1985).
It seems therefore appropriate to provide an updated
species checklist for this group. This list is also meant
as a preliminary basis for a much needed phylogenet-
ic analysis of the Parnassiinae at the species level, with
the intention to point out all cases of presently prob-
lematic species delimitations.

CHECKLIST

The present list can only be regarded as a first step
towards a complete revision of the Parnassiinae even
at species level. For this list, a ‘splitter’ approach was
followed in many cases, in that geographically well se-
parated and morphologically distinct taxa are rather
given species status than treated as conspecific with
their geographic vicariants. None of the many availa-
ble infraspecific names appear in this list, but a num-
ber of former subspecies have been elevated here to
species rank following suggestions elsewhere in the li-
terature. Similarly, all recently described species in
the genus Parnassius are retained here in binominal
combination. However, all closely related, but strictly
allopatric species-group taxa are indicated in the pre-
sent list by indentation and omitting the generic na-
me. Under a more rigorous application of the biolo-
gical species concept, most of these cases might well
turn out as conspecific, and could be treated as such

157

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

for purely classificatory purposes.

The Parnassiinae are seen here to comprise the
eight genera listed below following recent classifica-
tions of the Papilionidae (Munroe 1961, Ackery
1975, Hancock 1983, Miller 1987), despite the fact
that the subfamily as currently defined might not rep-
resent a monophyletic group in phylogenetic terms
(Hiura 1980, Hauser 1990, 1993). All taxa are listed
fully with author and year, and their sequence follows
current ideas about taxonomic and phylogenetic rela-
tionships. Comments on individual taxa are present-
ed as a series of notes, which are numbered consecu-
tively and appear at the end of the checklist.

Subfamily Parnassiinae Swainson, 1840

Genus: Hypermnestra Ménétriés, 1848 O1
Hypermnestra helios (Nickerl, 1846)

Genus: Parnassius Latreille, 1804 02
Subgenus: Parnassius Latreille, 1804 03

Parnassius apollo (Linnaeus, 1758)

P. nomion Fischer De Waldheim, 1823

P. phoebus (Fabricius, 1793)
sacerdos Stichel, 1906 04
ruckbeili Deckert, 1909 05

P. actius (Eversmann, 1843)

P. jacquemontii Boisduval, 1836

P. tianschanicus Oberthür, 1879

P. epaphus Oberthür, 1879

P. bremeri Bremer, 1864

P. honrathi Staudinger, 1882

P. apollonius (Eversmann, 1847)

Subgenus: Driopa Korshunov, 1988 06

P. mnemosyne (Linnaeus, 1759)
P. stubbendorfii Ménétriés, 1849

hoenei Schweitzer, 1912 07
P. glacialis Butler, 1866

P. eversmanni Menetries, 1855

felderi Bremer, 1861 08
P. ariadne (Eversmann, 1843) 09
nordmanni [Ménétriés] in
Siemaschko, 1850 09, 10
clodius Ménétriés, 1855 09
P. orleans Oberthür, 1890
Subgenus: Tadumia Moore, 1902 11
P. hardwickii Gray, 1831
P. hunnyngtoni Avinoff, 1916 12
P. acco Gray, 1853
przewalskii Alpheraky, 1887 13

138

baileyi South, 1913

P. szechenyii Frivaldszky, 1886

P. cephalus Grum-Grshimailo, 1891
pythia Roth, 1932

P. maharaja Avinoft, 1916
labeyriei Weiss & Michel, 1989
nosei Watanabe, 1990

P. schultei Weiss & Michel, 1989

Subgenus: Koramius Moore, 1902

P. delphius (Eversmann, 1843)
maximinus Staudinger, 1891

P. cardinal Grum-Grshimailo, 1887

P. staudingeri Bang-Haas, 1882
stenosemus Honrath, 1890

P. stoliczkanus Felder & Felder, 1865
nandadevinensis Weiss, 1990

P. patricius Niepelt, 1911
priamus Bryk, 1913
hide Koiwaya, 1987

P. acdestis Grum-Grshimailo, 1891

Subgenus: Sachaia Korshunov, 1988

P. tenedius Eversmann, 1851

P. arcticus (Eisner, 1968)
ammosovi (Korshunov, 1988)

P. simo Gray, 1853
andrei Eisner, 1930
simonius Staudinger, 1889

P. boedromius Püngeler, 1901

Subgenus: Kailasius Moore, 1902

P. charltonius Gray, 1853

P. loxias Piingeler, 1901
autocrator Avinov, 1913

P. imperator Oberthür, 1883

P. inopinatus Kotzsch, 1940

Genus: Archon Hiibner, [1822]

Archon apollinus (Herbst, 1798)
A. apollinaris (Staudinger, 1892)

Genus: Zerynthia Ochsenheimer, 1816

Zerynthia rumina (Linnaeus, 1758)

Z. polyxena ([Denis & Schiffermüller], 1775)

Genus: Allancastria Bryk, 1934

Allancastria cerisy (Godart, [1824])
cretica (Rebel, 1904)
louristana (Le Cerf, 1908)

37
38

39

40

41

42
43
44

A. deyrollei (Oberthiir, 1869) 45
A. caucasica (Lederer, 1864) 46

Genus: Sericinus Westwood, 1851

Sericinus montela Gray, 1853

Genus: Bhutanitis Atkinson, 1873 47
Bhutanitis mansfieldi (Riley, 1939) 48
B. thaidina (Blanchard, 1871)

B. lidderdalii Atkinson, 1873

B. ludlowi Gabriel, 1942

Genus: Luehdorfia Criiger, 1878

Luehdorfia puziloi (Erschoff, 1872)
L. japonica Leech, 1889

L. chinensis Leech, 1893 49
L. longicaudata Lee, 1982 50
Notes

01.—The genus Hypermnestra has long been inclu-
ded in the subfamily (Spuler 1892, Stichel 1907a,
Bryk 1935, Ackery 1975), but recent phylogenetic in-
vestigations suggest that it might not form part of the
Parnassiinae as a monophyletic group (Hiura 1980,
Hauser 1993).

02. — The splitting of the genus Parnassius into sev-
eral genera had started by Moore (1901-1903), and
was further elaborated by Bryk (1935), and, most re-
cently, by Korshunov (1988, 1990). This is not re-
garded here as useful taxonomic practice (Sokolov
1929, Munroe 1961, Ackery 1975), particularly as
long as the genus is assumed to be monophyletic.
Some of the generic names proposed, however, are re-
tained here to designate presumably monophyletic
species-groups, which for taxonomic purposes can be
treated as subgenera (Munroe 1961; see notes 03, 06,
205056)

03. — The subgenus Parnassius or the so-called
‘apollo-group’ comprises a morphologically and eco-
logically homogeneous group of species with larval
hostplants restricted to Crassulaceae and Saxifra-
gaceae (Bryk 1935, Ackery 1975, Kreuzberg 1987a).
The proper allocation of many taxa from Central
Asia, where several species of the group coexist, is still
problematic and awaits further detailed studies. For
the present list, with the exceptions noted below, the
arrangement of species by Ackery (1975) is followed.

04. — The separation of the European taxa as a spe-
cies distinct from P. phoebus is largely based on pecu-
liar ecological requirements and morphological dif-
ferences of the larvae, in addition to considerable
geographic separation. P. sacerdos is restricted to hab-

Hauser: Parnasstinae checklist

itats along and near small streams of running water at
elevations between 1500 and 2600 m in the
European Alps, and its larvae exclusively feed on
Saxifraga aizoides (Weidemann 1986, Descimon et al.
1989, Schweizerischer Bund fiir Naturschutz 1991).
In contrast, habitat preferences are quite variable in P.
phoebus in Northern Asia and Western North
America where the species can occur from near sea
level in Eastern Siberia and steppe zone habitats in
the Altai Mountains (Kurentzov 1970, Mracek 1989)
up to the nival zone above the tree line in the Rocky
Mountains (Ferris 1976). In North America, larval
hostplants of P. phoebus are various species of Sedum,
whereas local species of Saxifraga, as well as S. aizoides
are not accepted by larvae in captivity (Scott 1986:
189, Nardelli 1991).

05. — P. ruckbeili had been originally described as a
subspecies of P. phoebus, and was treated subsequent-
ly as such by most authors (Rothschild 1918b, Bryk
1935, Eisner 1976). The taxon, however, shows also
affinities both in external appearence and male geni-
talia to P. actius (Bryk & Eisner 1935, Hering 1935),
and Eisner (1961: 180) finally placed it as a separate
species. The known geographic range of P. ruckbeili is
restricted to mountains North-East of the Tarim ba-
sin completely separated from both P. phoebus, and P.
actius, which leaves the species status doubtful. The
name is often spelt as ‘riickbeil? or ‘rueckbeilî (e.g.,
Bryk 1935, Hering 1935, Eisner 1976), both of
which represent unjustified emendations.

06. — The ‘mnemosyne-group’ was in the older liter-
ature often referred to by the (sub-)generic name
Doritis Fabricius, 1807 (Moore 1901-1903, Sokolov
1929, Bryk 1935, Munroe 1961), which, however,
has P. apollo as its type-species, and thereby is invalid
as an objective synonym of Parnassius Latreille, 1804
(Hemming 1967). The name Adoritis Koçak, 1989 is
invalid as an objective synonym of Driopa
Korshunov, 1988.

07. — P. hoenei from the Japanese Island of
Hokkaido had originally been described as a subspe-
cies of P. stubbendorfii and was subsequently treated
as such by most authors (Bryk 1935, Eisner 1974,
Ackery 1975). Recently, the taxon has been elevated
to species rank by several Japanese authors (Fukuda et
al. 1982, Kitahara 1990), but the conclusive evidence
in support of such a view is not clear.

08. — P. felderi had been described as species and
was accepted as such by most early authors (Stichel
1907a, Verity 1905-1911, Rothschild 1918b; but see
Elwes 1886). Later on, the taxon was generally placed
as a subspecies of P. eversmanni (Bryk 1922, Bang-
Haas 1927, Eisner 1961, 1974, Kurentzov 1970,
Ackery 1975), but some doubts about its proper stat-
us remained (e.g., Bang-Haas 1927: 8, Eisner 1974:
90). The allopatric distribution of P. felderi and typi-

139

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

cal forms of P. eversmanni (see Iwamoto & Inomata
1988), and the occurrence of apparent intermediate
taxa (Bryk 1935, Eisner 1961), give little support to
maintain it as a distinct species.

09. — P. ariadne, P. nordmanni, and P. clodius have
always been treated as distinct species despite their
close morphological and ecological similarity, and
their strictly allopatric distribution (Elwes 1886,
Stichel 1907a, Bryk 1935, Eisner 1974, Ackery
1975). This practice is followed here, but it should be
pointed out that the degree of morphological differ-
ence between these taxa, for example, in male and fe-
male genitalia (Hering 1932, Orr 1988), especially,
between P. nordmanniand P. clodius is not very large.
For consistency, then, comparable cases of geograph-
ically vicariant taxa should also be classified as differ-
ent species (e.g. 04).

10. — The author of P. nordmanni is sometimes
been given as ‘Nordmann 1851’ (Ackery 1975,
Hancock 1983), based on conclusions by Hemming
(1934) that a description of the taxon by Ménétriés
was never published. However, according to biblio-
graphic studies by Nekrutenko & Kerzhner (1986)
the original description of P. nordmanni by Ménétriés
was indeed published in 1850 and thus takes prece-
dence.

11.—In the subgenus 7adumia, the so-called ‘acco-
group, Stichel (1907a), Bryk (1935), and Eisner
(1976), had included P. acco, P. simo, P. tenedius, and
related forms, but not P. szechenyi, P. cephalus, P.
maharaja, and P. hardwickii, which were placed in
different (sub-)genera. The separation of the szmo-
and tenedius- groups from Tadumia on morphologi-
cal grounds (Munroe 1961, Ackery 1975) is further
supported by biological characters (Kreuzberg 1985,
1987a). A close relationship between P. acco, P. sze-
chenyii, P. hardwickii, and P. cephalus with all asso-
ciated taxa is supported by similarity in male genital-
ia, sphragis structure, and larval hostplants (Hering
1932, Kreuzberg 1987a, Weiss & Michel 1989).

12. — P. hunnyngtoni had originally been described
as a separate species on the basis of its extremely small
size and different wing pattern (Avinoff 1916), but it
was subsequently treated as conspecific with P. acco
by Bang-Haas (1927), Eisner (1976), and Ackery
(1975). In contrast to all other Parnassius, this species
flies at high altitudes in early spring (Riley 1927,
W.Eckweiler, pers. com.). Furthermore, its sympatric
occurrence with P. acco (Riley 1927, Weiss & Michel
1989) leaves little doubt as to place it as a separate
species. The species name is often found spelt as ‘han-
nyngtont or ‘hanningtonî (Bryk 1935, Eisner 1976,
Weiss 1992), both of which are unjustified emenda-
tions.

13. — P. przewalskii had been described as a separ-
ate species and was treated as such by Stichel (1907a),

140

Verity (1905-1911), Bryk (1935), and Munroe
(1961), whereas it was considered as conspecific with
P. acco by Hering (1937), Ackery (1975), Eisner
(1976), Hancock (1983), and Weiss (1992). Recent
discoveries have extended the known range of P. prze-
walskii considerably (Huang & Murayama 1991,
Schulte 1992), which now covers most of North-
Western Tibet and Qinghai. Some of the new taxa
described appear intermediate in wing pattern with P.
acco, and leave further doubts about the species status
of P. przewalskii (Weiss 1992).

14. — P. baileyi is another questionable species geo-
graphically separated but close to P. acco, which had
originally been described as a subspecies of P. acco
from South-Eastern Tibet. Subsequently Bryk
(1932a) described P. rothschildianus as a species separ-
ate from P. acco from Western Sichuan, which he lat-
er united with baileyi and another new taxon (Bryk
1932b). All these taxa were later on placed by Bryk
(1935) as subspecies of P. przewalskii, whereas Bang-
Haas (1927), Eisner (1976), and Ackery (1975) con-
sidered them all conspecific with P. acco. New reports
of ‘typical’ P. acco occurring in South-East Tibet
(Nosé 1990), in addition to reported constant differ-
ences in genitalia (Weiss 1992) suggest that possibly
two species might occur in that region, and give rea-
son to tentatively maintain P. baileyi as a species.
However, a thorough taxonomic revision of the entire
acco-group is certainly needed and should yield a
more definite conclusion.

15. — The description of P. pythia is based on a sin-
gle small female specimen from South-Western
Tibet. The taxon was placed as a subspecies of P.
cephalus by Bryk (1935), but it was maintained as a
separate species in a list by Munroe (1961). The type
locality of P. pythia was at the time of its discovery
well separated from the then known range of P. ceph-
alus, but geographically intermediate populations
have since been found, which all appear conspecific
with P. cephalus (Weiss 1992).

16. — Originally, P. maharaja had been described as
a separate species (Avinoff 1916), but it was placed as
a subspecies of P. cephalus by Bryk (1922), Eisner
(1958, 1976), and Ackery (1975) despite constant
differences in wing venation and genitalia between
the two taxa (Hering 1932, Bryk 1935). The sympat-
ric occurrence of newly discovered taxa closely related
to P. maharaja with P. cephalus in Southern and
South-Eastern Tibet (see below, 17 + 18) give further
support to maintain P. maharaja as a separate species
from P. cephalus (Bryk 1935, Munroe 1961, Weiss
1992):

17. — The recently discovered ?. labeyriei from
Southern Tibet is morphologically very similar to P.
maharaja from Ladakh (see Michel & Weiss 1989,
Watanabe 1990), and probably just represents a geo-

graphic vicariant form of the latter. However, pos-
sibly intermediate records are currently missing from
Western Tibet, which certainly is one of the areas that
have never been adequately surveyed. Additional new
discoveries indicate that this species is more widely
distributed in Western China (Weiss 1991); see be-
low, 18.

18. — P. nosei from E. Tibet is also very similar to P.
maharaja, and has already been regarded as conspecif-
ic with P. labeyriei (Weiss 1991). However, both P.
labeyriei and P. nosei reportedly occur next to each
other in South-Eastern Tibet (Nosé 1990), which is
the reason to retain the latter provisionally here as a
distinct species. Probably at least one of the two taxa
will turn out to be conspecific with P. maharaja.

19. — P. schultei has been described from only three
worn specimens from Southern Tibet (Weiss &
Michel 1989). The genitalia of the male holotype ap-
pear to differ considerably from P. cephalus, and also
from P. maharaja and P. labeyriei, which gives reason
to place it here as a distinct species.

20. — The status of most of the following species
listed under the subgenus Koramius is still problemat-
ic, and a comprehensive revision of the entire ‘delphi-
us-group’ is much needed. Previous authors have gen-
erally recognized only a single (Stichel 1907a, Verity
1905-1911, Hering 1932), two (Elwes 1886, Bang-
Haas 1915, Rothschild 1918b), three (Bang-Haas
1927, Ackery 1975, Hancock 1983), four (Bryk
1935, Collins & Morris 1985), or at most five differ-
ent species (Munroe 1961). However, detailed com-
parative investigations on the ecology and genital
morphology (Kreuzberg 1985) indicate that in sever-
al mountain ranges of Central Asia (Tian-Shan, Alai,
Pamirs) two to four distinct species of the ‘delphius-
group’ coexist. The species recognized by Kreuzberg
(1985, 1987a) are accepted for the present list (Weiss
1992), but it should be noted that an exact delimita-
tion of these species and a proper allocation of the
more than 50 taxa named formerly lumped under P.
delphius still awaits to be accomplished.

21. — P. maximinus was recognized as a distinct
species from P. delphius by Kreuzberg (1985) on the
basis of different ecological requirements and differ-
ences in larval colouration. The species seems restrict-
ed to the Western Tian-Shan range and shows no sig-
nificant overlap in its range with P. delphius (Weiss
1992), which leaves some doubt about its status as a
separate species.

22. — P. cardinal had originally been described as a
subspecies of P. delphius, and was first recognized as a
separate species by Stshetkin (1979) based on its sym-
patric occurrence with another de/phius-group taxon.
Subsequent studies by Kreuzberg (1985) confirmed
the coexistence of P. cardinal and P. staudingeri in
Tadzhikistan, where P. cardinal occurs mainly at ele-

Hauser: Parnassiinae checklist

vations between 3000 and 3200 m.

23. — P. staudingeri was reinstated as a different
species from P. delphius by Kreuzberg (1985) on the
basis of differences in male and female genitalia as
well as wing pattern, in addition to ecological differ-
ences regarding habitat and larval hostplants. The
species appears to co-exist in parts of its range with P.
cardinal, but is strictly allopatric with P. delphius and
P. stenosemus (Weiss 1992).

24. — P. stenosemus had been described as a subspe-
cies of P. delphius and it was treated as such or as a
subspecies of P. stoliczkanus by Stichel (1907a),
Verity (1905-1911), Rothschild (1918b), Bang-Haas
(1927), Eisner (1959a, 1976), and Ackery (1975).
The taxon was first given species status by Moore
(1901-1903), followed by Bryk (1935), Munroe
(1961), and Weiss (1992), on the basis of constant
differences in wing pattern and male genitalia with
the sympatrically occurring P. stoliczkanus. However,
the known range of P. stenosemus shows no overlap
with P. staudingeri, and further studies are needed to
confirm its species status.

25. — P. stoliczkanus had been described as a separ-
ate species, but was later considered a subspecies of P.
delphius by Stichel (1907a), Verity (1905-1911),
Avinov (1913), Rothschild (1918b), Eisner (1959a,
1976) and Ackery (1975). It occurs sympatrically
with P. stenosemus, and had been already given species
status by Elwes (1886), Bang-Haas (1927), Bryk
(1935), Munroe (1961), and Collins & Morris
(1985).

26. — The original description of P. nandadevinen-
sis is based on a single worn male specimen from
Northern Uttar Pradesh, India. According to illustra-
tions of the holotype and parts of its genitalia, it ap-
pears intermediate between P. acdestis and P. stolicz-
kanus, and could possibly represent an aberrant
specimen of the latter species.

27. — Since its original description P. patricius had
generally been accepted as a species separate from P.
delphius (Bryk 1935, Munroe 1961, Ackery 1975,
Hancock 1983, Weiss 1992; but see: Bang-Haas
1915), and recent morphological and ecological stud-
ies further support this view (Kreuzberg 1985).

28. — P. priamus had been originally described as a
subspecies of P. acdestis, and subsequently the taxon
was considered as either belonging to this species or
to P. delphius (Bang-Haas 1915, Bang-Haas 1927).
Marked differences in the shape of the sphragis
between P. priamus and P. acdestis led Bryk (1932b,
1935) to separate it from the latter species and to
place it as a subspecies of P. patricius. Apparent differ-
ences in male genitalia between P. priamus and P. pa-
tricius could support the view of two different species
(Weiss & Michel 1989), but both taxa occur allopat-
rically (Weiss 1992).

141

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

29. — P. hide has been described recently from the
Kunlun range, and was subsequently also found in
Western Sichuan and South-Eastern Tibet (Koiwaya
1987, Nosé 1990, Shinkai 1990). The taxon appears
not to differ much in male genitalia from P. priamus
from Southern Tibet, and probably is conspecific
with the latter or with P. patricius (see Weiss &
Michel 1989: 12).

30. — The type species of Sachaia is P. tenedius by
original designation (Korshunov 1988), and Kreiz-
bergius Korshunoy, 1990 is available as a separate ge-
neric name for P. simo and related taxa in this group.

31. — Originally, P. arcticus had been described as a
northern subspecies of P. simo, a species which does
not occur in Siberia, but Eisner (1969) later himself
corrected this misjudgement, and placed the taxon as
a subspecies of P. tenedius (Eisner 1976). In 1984,
however, another Parnassius species was discovered in
Central Siberia that on ecological and morphological
grounds is clearly separate from P. tenedius, without
much consideration, this species has been repeatedly
referred to as P. arcticus (Mracek 1989, Weiss 1991),
but proper identification must still await critical ex-
amination of the holotype (see below, 32).

32. — As judged from the original description
(Korshunov 1988), P. ammosovi appears not to be
conspecific with P. tenedius, and probably represents
the second Siberian species of that group (see above,
31); this taxon, however, would then be a junior
synonym of P. arcticus depending on the identity of
the holotype of the latter.

33. — P. andreji has been separated recently togeth-
er with another six subordinate taxa from P. simo by
Weiss (1991), based on differences in wing pattern
and male genitalia; both character sets, however, have
not yet been adequately documented. The fact that P.
andrei is not found sympatrically with P. simo, which
occurs over most of Tibet into South-West China,
makes this separation furthermore questionable.

34. — P. simoniushad been described as a subspecies
of P. simo and was treated as such by most authors
(Bryk 1935, Munroe 1961, Ackery 1975). However,
Avinov (1913) already had remarked on constant dif-
ferences in male genitalia between P. simonius and P.
simo (see also, Hering 1932), but did not separate
both taxa at the species level. In a comparative study
on the group, Kreuzberg (1985) elevated the taxon to
species rank supported by constant differences in
postabdominal morphology. As far as known, how-
ever, P. simonius and P. simo occur allopatrically and
the ranges of the two taxa show no overlap (Weiss
199):

35. — P. boedromius had originally been described
as a separate species, but was since generally regarded
as a subspecies of P. simo (Stichel 1907a, Bryk 1935,
Ackery 1975), except for Avinov (1913) followed by

142

Rothschild (1918b), who had recognized that it oc-
curs sympatrically with typical forms of P. simo.
Recent studies by Kreuzberg (1985) supported
Avinov s early conclusions through the confirmation
of sympatric occurrences of the two species, and eco-
logical and morphological differences.

36. — The type species of Kailasius is P. charltonius
by original designation, and a separate generic name,
Eukoramius Bryk, 1935, is available for P. imperator
(Hemming 1967).

37. — P. loxias had been described as a species and
has since been treated as such by most authors (Verity
1905-1911, Bryk 1935, Eisner 1959b, Munroe 1961,
Ackery 1975, Kreuzberg 1985), except for Bang-Haas
(1927) who regarded it as a subspecies of P. charlton-
ius. The recent discovery of its biology confirms its
status as a separate species from P. charltonius (Kreuz-
berg & Pljushch 1989).

38. — P. autocrator was originally described as a
subspecies of P. charltonius (Avinov 1913, Bryk
1935), but because of its unique wing pattern was lat-
er on universally accepted as a separate species (Bang-
Haas 1927, Eisner 1959b, Ackery 1975). In terms of
genitalia and ecology, however, it shows a close affin-
ity to P. loxias (Bryk 1937, Kreuzberg 1987b), from
which it is geographically well separated (Weiss
1991).

39. — A. apollinaris had always been regarded as a
subspecies of A. apollinus (Stichel 1907a, Bryk 1934,
Igarashi 1979), until de Freina (1985) proposed the
existence of two distinct but allopatric Archon species
based on differences in genitalia and larval coloura-
tion. The recent discovery of areas of sympatry of two
distinct taxa in the province of Mardin in Southern
Turkey further supports this view (Carbonell 1991).

40. — The generic name frequently used for
Zerynthia in the older literature is Thais Fabricius,
1807, which is invalid as a junior primary homonym
(Hemming 1967). The older replacement name
Parnalius Rafinesque, 1815 has been suppressed for
the principle of priority by the Plenary Powers of the
International Commission on Zoological Nomen-
clature (Opinion 1134).

41.—The genus Allancastria has been synonymized
by some authors with Zerynthia (Ackery 1975,
Igarashi 1979, Olivier 1991), but it is retained here as
a separate genus on the basis of considerable differ-
ences in genitalic structures (see: Bryk 1934,
Nekrutenko 1990).

42. — The species name has often been incorrectly
cited as ‘cerisyi Godart, 1822’ (e.g., Rothschild
1918a, Bryk 1934, Bernardi 1970, Eisner 1974)

based on a reference that was never published (see

Cowan 1970: 17). A. cerisy was long regarded as the

only species in the genus (Stichel 1907b, Bryk 1934,
Munroe 1961, Eisner 1974, Ackery 1975, Igarashi

1979), and the discovery of two sympatric Allan-
castria species in Lebanon (Larsen 1973, 1975) has fi-
nally led to the separation of several other species (see
below, 43-46). A complete revision of all Allancastria
taxa appears, therefore, much needed (Olivier 1991).

43. — The elevation of A. cretica endemic to the isle
of Crete as a different species from A. cerisy was first
proposed by Koçak (1981), and is presumably based
on differences in wing shape and colouration only. As
a number of phenotypically different populations of
A. cerisy from other Eastern Mediterranean islands
have also been named as subspecies (see Bryk 1934,
Olivier 1991), the single treatment of this taxon as a
species (e.g., Kudrna 1986: 169) must appear ques-
tionable, and cannot be justified by a few distinguish-
ing features and geographic separation alone.

44. — À. louristana had been described as a subspe-
cies of A. cerisy, and it was first treated as a separate
species by Kuhna (1977), followed by Blom & Eisner
(1979), de Freina (1979), and Collins & Morris
(1985). The distinguishing characters supporting
such a view, however, have not been clearly stated.
Areas of sympatry between A. louristana and either A.
cerisy or A. deyrollei are presently not known, and the
status of this taxon therefore remains questionable.

45. — Although A. deyrollei had originally been giv-
en species rank when first described, it was recognized
only recently as a separate species from A. cerisy based
on differences in wing pattern, genital morphology
and early stages (Bernardi 1970, Larsen 1973, 1975).
The species occurs partly sympatrically with A. cerisy
in Southern Turkey and Lebanon (Kuhna 1977, de
Freina 1979, 1986).

46. — A. caucasica had been described as a subspe-
cies of A. cerisy, and it was first regarded as a different
species by Kuhna (1977) based on characters of wing
pattern, colouration and genitalia. This view has since
been largely accepted (de Freina 1979, Collins &
Morris 1985, Nekrutenko 1990), but the exact de-
limitation of the species and the allocation of various
subspecific taxa to A. cerisy and A. caucasica, respec-
tively, is not clear.

47. — At present, the genus Bhutanitis is held to
comprise the four species listed here (Ackery 1975,
Collins & Morris 1985). Recent field studies in
South-West China, however, seem to indicate that, at
least, three different Bhutanitis species exist in that
area (Hou 1992). As so far only two species were
known to occur in the region (Bryk 1934, Ackery
1975), the existence of an additional species in this
genus is quite likely. Although several new taxa were
named in Hou’s abstract (1992), these names cannot
be accepted as available. In the older literature, the ge-
nus is often referred to by the name Armandia
Blanchard, 1871, which is invalid as junior homo-
nym (Hemming 1967).

Hauser: Parnassiinae checklist

48. — A separate genus, Yunnanopapilio Hiura,
1980, has been proposed for B. mansfieldi, but a ge-
neric separation of this species from Bhutanitis seems
not justified (Saigusa & Lee 1982), particularly when
considering its biology and early stages (Lee 1986a,
1986b).

49.— L. chinensis had been described as a subspecies
of L. japonica, and was treated as such by most au-
thors (Verity 1905, Rothschild 1918a, Ackery 1975),
except for Bryk (1934), and later on Eisner (1974)
who placed it as a subspecies of L. puziloi. The taxon
was first recognized as a species by Lee (1978) based
on marked differences in the early stages, a view
which has now become widely accepted (Collins &
Morris 1985, Nosé 1990).

50. — L. longicaudata was recently discovered in
Western China, and occurs partly sympatrically with
L. chinensis. The differences seen in adult morpholo-
gy and early stages seem to furthermore justify its stat-
us as a separate species (Nosé 1990).

CONCLUSIONS

In the present list, 76 species taxa are included in
the subfamily Parnassiinae. Although 22 of these bi-
nominal taxa are indicated as questionable species
some of which will eventually have to be synonymi-
zed, the remaining number of recognized species will
still exceed the fourty to fifty species listed in most
previous taxonomic reviews of the subfamily.
Generally, species numbers for the eight genera inclu-
ded here in the Parnassiinae range from 39 species re-
cognized by Stichel (1907a, 1907b) at the turn of the
century, to 42 (Eisner 1974, 1976), 44 (Bryk 1934,
1935, Ackery 1975), 48 (Hancock, 1983), 49
(Munroe 1961), or, at most, 53 species listed by
Collins & Morris (1985). This new increase in spe-
cies number should come somewhat unexpected, par-
ticularly, in a much collected and taxonomically well
studied group of papilionid butterflies. Most of the
taxa recently recognized.as distinct species, however,
were already known at the beginning of this century,
and only very few represent genuine discoveries (e.g.,
Parnassius schultei, P. arcticus, Luehdorfia longicauda-
ta). Thus, the majority of recently discovered ‘new’
species in the Parnassiinae came about by a change of
status rather than by the description of new taxa.

This recent increase in species number can prob-
ably be explained by a change of general concepts in
systematics, which is well reflected by the different at-
titudes of specialists dealing with the Parnassiinae at
different times. In the first half of this century, most
authors were primarily concerned with the study of
(intraspecific) variation in wing pattern exclusively
based on large collections of adult specimens (e.g.,

Austaut 1899, Verity 1905-1911, Bryk 1934, 1935).

143

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

This approach produced an immense number of in-
fraspecific taxa named for the Parnassiinae (Bryk
1922, Eisner 1974, 1976), and it naturally obscured
somewhat the question of delimitation of species. In
recent times, more and more authors tried to rely for
their taxonomic decisions on other morphological,
biological and ecological characteristics, many of
which usually exhibit larger differences at the species
level (e.g., Larsen 1975, Lee 1978, Kreuzberg 1985).
This shift away from a purely descriptive taxonomy
relying on a single set of adult characters was partly
driven by and coincided with the change from a typo-
logical towards a biological species concept in system-
atics (Mayr 1982). In the Parnassiinae, the long ap-
plied taxonomic practise to record and name all forms
of infraspecific variation in wing pattern still persists
today (e.g., Schulte 1992, Weiss 1992), but is slowly
giving way to a more biological aproach considering
all possible characteristics and pecularities of the
group. From this historical perspective it appears
therefore understandable that, despite the consider-
able taxonomic efforts already undertaken in the past,
a comparatively high number of unresolved cases at
species level in the Parnassiinae remains. The primary
purpose of this paper is to direct attention to these
cases and encourage further, more detailed studies.

ACKNOWLEDGEMENTS

Helpful comments and criticisms on a preliminary
draft of this paper were received from Dr. W.
Eckweiler, Frankfurt/Main, Prof. Dr. C. M.
Naumann, Bonn, and Ju. Ju. Stshetkin, Taganrog,
Russia. Financial support during the preparation of
this paper by the Deutsche Forschungsgemeinschaft
is gratefully acknowledged.

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Entwicklung -
— J. Neumann-Neudamm,

Received: September 1993
Accepted: 16 September 1993

WILLEM HURKMANS

Institute of Taxonomic Zoology (Zoological Museum), Amsterdam

A MONOGRAPH OF MERODON (DIPTERA:
SYRPHIDAE). PART 1

Hurkmans, W., 1993 A monograph of Merodon (Diptera: Syrphidae). Part 1. — Tijdschrift
voor Entomologie 136: 147-234, figs. 1-106. [ISSN 0040-7496]. Published 10 December 1993.
This first part of a revision of the genus Merodon deals with 61 species characterized by two syn-
apomorphies, viz. arcuate abdominal pruinose bands and abdomina with tapering second ter-
gite. Eleven monophyletic species groups are recognized, accounting for 51 species. Seven spe-
cies are placed in an undefined set while three species are discussed separately. Descriptions of
the species groups are provided. All species are described and their important characters figured.
Of the 91 nominal taxa mentioned in the checklist, 61 are species, 2 subspecies, 6 varieties and
22 synonyms, 4 of which are new. Of these, the following 28 species and one subspecies are new
to science: M. mariae, M. ottomanus, M. testaceoides, M. altinosus, M. hypochrysos, M. mar-
ginicornis, M. sophron, M. ankylogaster, M. auronitens, M. oidipous, M. persicus, M. tangeren-
sis, M. xanthipous, M. aberrans isperensis subsp. n., M. cupreus, M. lusitanicus, M. splendens.,
M. warnckei, M. kaloceros, M. vandergooti, M. bequaerti, M. elegans, M. lucasi, M. nitidi-
frons, M. satdagensis, M. schachti, M. taniniensis, M. toscanus, M. aureotibia.

Lectotypes are designated for 23 nominal taxa; three new combinations are introduced. An an-
notated list of species excluded from Merodon is given. Azpeytia shirakii nom. n. is proposed for
Merodon scutellaris Shiraki.

Willem Hurkmans, c/o Institute of Taxonomic Zoology (Zoological Museum), Department of
Entomology, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands.

Key words. — Diptera; Syrphidae; Merodon; new subdivision; new species; lectotypes; excluded
species; nomen novum.

CONTENTS Index suites lose CORAL) zend oa LOLA 235
Ie Cao teen ite 147 facere
LEE alesh SRO EA 148
Nie tera cn de Sn 148 The genus Merodon is a moderately large group of
The genus Merodon Meigen, 1803 ..................... 149 syrphidflies naturally occurring in the palaearctic and
ROretheSpEcIEStreaTe di... 153 ethiopian faunal regions. As far as known the deve-
SA Pnt Gye i nai ina 157 lopment of the larvae takes place in bulbs or rhizomes
desuncusassemblape>— ron 157 of Liliaceae and Amaryllidaceae. The last comprehen-
TEBE CROWD eee per > cdi ae lu 162 sive discussion of the genus, by Sack (1931), is outda-
TAD § > li, Sa Sek nude a 169 ted, contains several errors, and deals with only about
slzumpespionpiio- > rr 175 half the species currently known. The descriptions
[TE OAD). —— — > 185 and discussions of Merodon species published so far
OA CON SOUPE. ari a 186 largely disregard important male genital characters
DLC TOO EC DIE io Lot (i. (i. 188 considered suitable for grouping purposes and species
LIT NEO DR sen 189 recognition.
ZOE RON ne hen re ne 191 This paper deals with eleven monophyletic species
GEASS EUTOTIS) PCO) sn 193 groups defined by apomorphies and a remaining un-
ÉGALE e or oS 194 defined set of species. All species discussed in this
CROIENT 197 paper are grouped by two synapomorphies, viz. arcu-
Remamino species Lin 203 ate pruinose bands on the abdominal tergites, and an
Annotated list of species discussed ...................... 204 abdomen with tapering second tergite. Only the
Annotated list of species excluded from males are used for grouping purposes, since their gen-
broder boni levi lana 207 italia yield enough data for subdivision. The female
Neknowledpementst-.:- orione 208 genitalia are rather uniform; moreover the ornamen-
References: teste mens en 208 tation of the legs and specialized pubescence areas are

147

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

usually much less developed in the females than in the
males.

MATERIAL

Material was studied from the following institutes
and private collections (with acronyms used in this
paper): BMNH, British Museum (Natural History),
London, England; MNHN, Muséum National d’
Histoire Naturelle, Paris, France; LSF, Museo
Zoologico della Specola, Florence, Italy; MCSN,
Museo Civico della Storia Naturale, Milan, Italy; BSA,
Benediktinerstift Admont, Admont, Austria; NHMW,
Naturhistorisches Museum Wien, Vienna, Austria;
ZMHB, Museum für Naturkunde, Von Humboldt
Universität, Berlin, Germany; DEIC, Institut für
Pflanzenschützforschung, Eberswalde, Germany;
HLMD, Hessisches Landesmuseum, Darmstadt,
Germany; MENS, Museum für Naturkunde, Stuttgart,
Germany; LAU, Musée Zoologique, Lausanne,
Switzerland; KBIN, Koninklijk Belgisch Instituut voor
Natuurwetenschappen, Brussels, Belgium; MRAC,
Museum voor Midden Afrika, Tervuren, Belgium;
ZMUC, Zoological Museum, Copenhagen University,
Copenhagen, Denmark; sAL, Zoological Institute of
Salamanca University, Salamanca, Spain; IEE,
Instituto Espanol de Entomologia, Madrid, Spain;
ZMAS, Zoological Institute of the Russian Academy of
Sciences, St. Petersburg, formerly Leningrad, Russia;
TMA, Természettudomänyi Museum Allättara,
Budapest, Hungary; BTM, Bakonyi Természettudo-
mänyi Museum, Zirc, Hungary; LHW, Entomological
Institute, Agriculture University, Wageningen, the
Netherlands; RMNH, Rijksmuseum van Natuurlijke
Historie, Leiden, the Netherlands; JLR, J. A. W.
Lucas, Rotterdam, the Netherlands (the most impor-
tant single source of material; material from several
other, mainly private collections was made available
by Mr Lucas as well); ZMAN, Institute of Taxonomic
Zoology, Zoological Museum, University of Amster-
dam, Amsterdam, the Netherlands; PC, S. J. Para-
monov collection. Many types of species described by
Paramonov were formerly preserved in his private
collection. After Liepa (1969) this collection is consi-
dered to have been lost.

METHODS

Preparation of specimens

In order to study the male genitalia, specimens pri-
or to examination were relaxed using dilute ethanol
vapour at room temperature for 6 to 48 hours depen-
ding on specimen size. After relaxation the genitalia
were extracted with an insect pin with hooked tip.
Isolation of the genitalia often proved necessary; in
such cases the genitalia were stored in microvials con-

148

taining a 60%-40% mix of glycerol and ethanol. The
genitalia were examined in the dry state since boiling
or prolonged submersion in KOH led to maceration
and damage to the pubescence. The pubescence
found in many species could hardly be seen properly
in wet state. We were confronted with the same pro-
blem with genitalia already isolated and stored in
Canada balsam. Genitalia already stored in microvials
(in ethanol/glycerol mixture) were repeatedly washed
with 70% ethanol and left to dry prior to examina-
tion.

In Merodon, the female genitalia have been of little
use for taxonomic purposes since they are hardly scle-
rotized. If dried they shrivel completely, and if stored
in a liquid medium the various pubescences are diffi-
cult to distinguish.

Methods used in grouping

The species have been grouped on the basis of apo-
morphic characters. Characters were considered apo-
morphic after a preliminary survey of the correspon-
ding characters in other species groups and in the
genus Eumerus, the presumed sister group of
Merodon, on the ground of similarities in the genita-
lia and the banding pattern on the abdomen of some
species of Eumerus formerly placed in the genera
Megatrigon Johnson and Amphoterus Bezzi (see also
Hull 1949: 395). On the basis of this comparison a
slightly oblique to parallel, straight banding pattern,
comparatively simple male genitalia and a comparati-
vely stout abdomen are therefore considered to be
plesiomorphic.

Terminology

The terms used to describe the various structures
are illustrated in figs. 1-4 (head), 5 (thorax), 6 (wing),
7 (leg) and 8-17 (male genitalia). The terms are main-
ly derived from McAlpine (1981), but some terms
from Speight (1987), Van der Goot (1981), Hull
(1949), Sack (1931) and Metcalf (1921) have been
used for elements occurring specifically in Syrphidae.
A few terms are introduced here to describe structures
in the male genitalia; they are illustrated in figs. 8-17.
The vertex angle and ocellar angle, as used here in de-
scribing the head, are illustrated in fig. 3.

The ratio obtained by dividing the compound eye
touchline length by the vertex height, measured from
its angle to the border with the occiput, is cited as the
tl-v ratio. This ratio is often a useful distinguishing
character. The antennal ratio, illustrated in fig. 2, is
also frequently used. Sternites and tergites are count-
ed in Roman numerals and abbreviated S and T, re-
spectively.

Drawings and maps
All drawings are original; they were made with the

aid of various stereo microscopes with drawing tubes
under 10-80 X magnification; all scale lines represent
1 mm.

Distribution maps are provided for all species. On
the distribution maps only those records are repre-
sented which are based on examined material or on
reliable references. References from literature for spe-
cies easily confused with other species have not been
included. Where many records for a species occur,
single records are represented by small symbols. The
large symbols represent five records in the same area.

THE GENUS MERODON MEIGEN, 1803

Merodon Meigen, 1803: 274. — Type-species Syrphus clavipes
Fabricius, 1781: 427 [designation of Westwood 1840:
137].

Lampetia Meigen, 1800: 34 [Suppressed by the I. C. Z. N.,
Opinion 678]. — Type-species Syrphus clavipes Fabricius,
1781: 427 [designation of Coquillett 1910: 557].

Exmerodon Becker, 1912: 604 (as subgenus of Merodon). —
Type-species Merodon (Exmerodon) fulcratus Becker,
1912: 604 [original designation].

Taxonomic and nomenclatural history

From 1781 to c. 1900

In 1800, Meigen established the genus Lampetia to
accommodate Syrphus clavipes Fabricius, 1781
(Meigen 1800). In 1803 he introduced the new name
Merodon for this genus, mentioning that Syrphus cur-
vipes [error for clavipes| belongs in Merodon (Meigen
1803). The work of Meigen (1800) has been placed
on the Official Index of Rejected and Invalid Works
in 1963 (I. C. Z. N. Opinion 678); since then only
the Meigen (1803) names should have been used.
Several species described by Fabricius and Rossi prior
to the description of Merodon were later incorporated
in this genus by e.g. Fabricius (1805).

At the start of the nineteenth century species now
included in Merodon were described in several genera
such as Eristalis Latreille, 1804, and Milesza Latreille,
1804. As, in the first half of the nineteenth century,
the concept of the various syrphid genera crystallized,
many new species were rightly described in Merodon.
During this period Fabricius (1805), Meigen (1822-
1838), Macquart (1834, 1842) and Rondani (1843-
1868) described and redescribed many species.
Several synonyms were created through interpreta-
tion of colour forms as species. Walker (1849-1860)
assigned many species to this genus, all but one incor-
rectly. Loew (1862a-b, 1869) described several new
species and also provided redescriptions for several
known species. Strobl (1893-1909) described several
new, and redescribed many known taxa. The authors
mentioned so far almost never discussed the differ-
ences between species, except when establishing a

Hurkmans: Monograph of Merodon

new taxon; they were not concerned with subdividing
the genus or with discussing its position within the

Syrphidae.

The twentieth century

The first author providing a comprehensive generic
description for Merodon was Verrall (1901) who loca-
ted this genus in the subfamily Eristalinae. Villeneuve
(1909-1934) published several new species and recog-
nized a number of synonyms within Merodon. Becker
(1912) subdivided Merodon into two subgenera, the
nominal subgenus Merodon and the new subgenus
Exmerodon Becker, 1912. The latter subgenus at that
time comprised only the type-species Merodon fulcra-
tus Becker, 1912. Stackelberg (in Stackelberg &
Richter 1968) included his new species dichopticus in
the same subgenus, established to accommodate spe-
cies with dichoptic eyes in the males. In the only mo-
nograph on Merodon so far, by Sack (1913a),
Merodon was not assigned to any subfamily although
it was compared with several genera in the Eristalinae.
Other publications by Sack (1913-1938) contain a
wealth of information on Merodon, many descrip-
tions of new species and a subdivision of the genus
based on external similarities. Major contributions to
the knowledge of Merodon were made by Paramonov
(1924-1937). He divided the genus into species
groups, not all of which were published. These
groups, like those of Sack (1913a), were based on ex-
ternal similarities. Moreover Paramonov critically re-
viewed papers by Becker (1912) and Sack (1931), re-
jecting some species and expressing his doubts about
others.

Other contributions were made by Gil Collado
(1930), Curran (1939) and, more recently, Séguy
(1961), who surveyed the species known from west-
ern Europe. Van der Goot (1981) in his work on the
north-west European Syrphidae provided a key for
the species occurring in that area.

The works mentioned so far did not make use of
genital characters to identify the species. In Merodon
many problems existed as to the separation of species,
which easily could have been solved by superficial ex-
amination of the genitalia. The first papers in which
genitalia of Merodon are discussed and figured were
by Glumac (1958) and Gaunitz (1969). Regrettably
the drawings provided are insufficient, but they af-
ford an insight into the great variety within the genus.

There is a considerable amount of literature on one
single species, the narcissus bulb fly Merodon equestris
(Fabricius, 1794), which is a pest in flower bulb cul-
tivation. This species probably received more atten-
tion in terms of literature produced, than all other
species together. This literature is mainly concerned
with economical aspects, genetics and life cycle. Some
of this literature is referred to under that species. No

149

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

other species of Merodon are of known economical
importance.

Diagnostic characters

The genus Merodon is characterized by the follo-
wing characters: (1) Radius 4+5 curves deeply into
cell r5 (e.g. fig. 19 d); (2) the apical part of Media is
strongly recurrent; its subapical portion is slightly si-
nuate (e.g. figs. 6, 19 d); (3) dm-cu is weakly,
smoothly curved, not sinuate (e.g. figs. 6, 19 d); (4)
femora 3 have a concave lower face and apically bear
a single projection at the distal end (e.g. figs. 20 a
through e).

These features are found in both sexes; the size and
shape of the projection on femora 3 is rather variable,
but this projection is always present although it is re-
duced in some species. |

Species of Merodon have sometimes been assigned
to other genera. More often, species of Mallota
Meigen, 1822, and Eumerus Meigen, 1822, are
wrongly assigned to Merodon. The genus Azpeytia
Walker, 1865, includes species that can be mistaken
for Merodon.

In Azpeytia the femora 3, which are less stout than
in Merodon, bear two projections or series of spinules
distally; the scutellum is much wider than in
Merodon; R4+5 curves only weakly into cell r5; dm-
cu is distinctly sinuate to kinked; moreover the oral
margin is less projecting than in the majority of
Merodon species (figs. 18 a, 19 b, 20 k, ]).

In Eumerus the femora 3 have a convex lower face
and bear two projections or spinule series as in
Azpeytia, the 3rd antennal article is relatively larger
and shorter in Eumerus than in most Merodon species;
R4+5 does not or hardly curve into cell r5 while dm-
cu is strongly kinked in Eumerus, in several Eumerus
small appending veinlets of the dm-cu are found;
these are not found in Merodon (figs. 18 c, 19 a, 20 f,
g, h, j).

In Mallota the apical part of vein M is not recurrent
as in Merodon; moreover Mallota species have a dis-
tinct facial knob, not found in Merodon (figs. 18 d, 19
c, 20 m, n).

Platynochaetus Wiedemann, 1830, is rather similar
in appearance to Merodon; the males may be distin-
guished by their curious antennae, while both sexes
show a facial knob; the position of the bands on the
abdomen is different (anteriorly in Platynochaetus, in
mid-region in Merodon). However, the wing venation
and the shape of femora 3 are quite similar to that in
Merodon (figs. 18 e, 19 e, 20 0, p).

The larvae of Merodon may be separated from
those of other genera by using the key of Heiss
(1938), who figured the larva of Merodon equestris. As

the larvae of most species are still unknown, identifi-

150

cation may be problematic. According to Maldonado
Capriles & Berrios (1977) the puparium of Merodon
can be confused with that of Copestylum species.

Description of the imagines

(fig. 23 d)

In all respects species of Merodon conform to the
criteria established for the Syrphidae by many au-
thors, e.g. Verrall (1901), Lundbeck (1916), Sack
(1931), Hull (1949) and Van der Goot (1981).

Habitus: Stoutly built to moderately slender, rarely
very slender syrphid flies. Body length 5-25 mm,
wing span 6-35 mm. Head, thorax and abdomen
about equally wide. Legs rather stout, the trochanters
3 and femora 3 in many species swollen and orna-
mented in the males. In many species pruinose bands
of microtrichiae are present on the thorax and abdo-
men. The thoracic surface is black or dark brown, in
several species metallic lustrous; the abdomen is black
to brown, often with yellow to red lateral spots which
may extend over all the abdomen. Wings mostly clear
although infuscate wings rarely occur. A typical repre-
sentative of the genus, M. distinctus Palma d is
shown in fig. 23 d. Many species mentioned in this
paper are superficially similar, especially those of the
alagoezicus, avidus, crassifemoris, nigritarsis and van-
dergooti groups.

Head: In the majority of species the males are hol-
optic while intermediate and dichoptic conditions oc-
cur in males of several species; females always dichop-
tic. Compound eyes dark brown to reddish, usually
bearing moderately dense whitish pubescence.
Antennae shorter than head except in a few species
with lengthened third article; third article usually
longest and widest, often conspicuous; arista inserted
rather basally on dorsal margin. Coloration of anten-
nae most variable. Face lacking knob; oral margin
clear, often conspicuous, seldom weak; genae absent;
facial surface little sculptured, dark, often metallic
lustrous, often yellow to white pruinose and pubes-
cent. Frons concolorous, often denser pubescent than
face; in females lateral strips on frons often pruinose,
leaving free a lustrous midstripe; species with an all
lustrous or all pruinose frons occur. The pruinose
frontal strips do not occur in dichoptic males. In
males with incompletely touching eyes the narrow
frontal strip is bald. Holoptic males show consider-
able variety in length of the touchline of the eyes. The
vertex is dark, in some species metallic lustrous, and
often conspicuously dark pubescent in the ocellar re-
gion. Occiput dark, pubescence concolorous, often
white to yellow pruinose down sides (figs. 3-4, 18 b).

Thorax: Slightly elongate, humeral calli well
marked, dorsal suture clear laterally only; colour dark,
often metallic lustrous on dorsum, scutellum, kate-

pisternum, anepisternum and anepimeron. Pube-
scence often conspicuous, sometimes strongly
coloured; length and density most variable; species
with very even, and with uneven pubescence occur.
Pruinose bands, if present, are longitudinal, often co-
alescent anteriorly; in some cases weak crossbands oc-
cur along the dorsal suture. The scutellum is without
specialized structures or specialized pubescence (fig.
5).
Wings: The wing surface is clear to partially or
strongly infuscate, in most species slightly, evenly
troubled. All species show the vena spuria characteris-
tic of the Syrphidae. Stigmal crossvein oblique, stig-
ma absent; cell rl open anteriorly; R4+5 strongly
curving into r5, dm-cu proximally sinuous, distally
rather strongly recurrent; r-m reaches M1+2 at ap-
proximately the middle of the anterior border of cell
dm; no appending veinlets occur on R4+5 or dm-cu;
the curves in CuAl and dm-cu are smooth. The
colour of the veins ranges from dark yellow to black
and is rather constant within species. The alula is in-
conspicuous; the squama and antisquama (upper and
lower calypters) are pale and bear a fringe of pale, yel-
low, even pubescence. The halteres usually are well
clear of the thoracic pubescence; the stalk is yellow,
the knob usually pale, sometimes dark (fig. 6, 23 d).
Legs: Legs 1 and 2 moderately robust. Leg 3 very
robust, with specialized structures: a single, usually
triangular projection on the femora 3 at the distal end
in all Merodon. Various ridges and spines occur on the
trochanters 3, femora 3, tibiae 3 and metatarsi 3 in
the males, seldom so in the females, of many species.
In several species the femora, the metatarsi, or both,
may be swollen or modified strongly in the males,
weakly in females. The colour is most variable, species
with all orange and with all black legs occur. In many
species the dark legs bear short, recumbent, golden
pubescence giving a false impression of yellow leg
colour. Specialized pubescences occur on the tro-
chanters 3 where a tuft of stout hairs is found in some
species, often inserted on a projection; on the tibiae 3
where a tibial floccus (a bunch of hairs) occurs in
some species, in most cases inserted on a spur; and on
the metatarsi 3 where stout short hairs occupy part of
the lower face in some species (figs. 40 c, 42 d).
Abdomen: Outline variable, usually comparatively
stout in the females, with some notable exceptions,
more slender in the males. The abdominal apex in the
males is posteriorly rounded with the genitalia often
clearly visible, in females it is posteriorly rounded to
more or less acute. In males of some species T IV is
conspicuously swollen to hold the large genitalia.
Colour in most species dark throughout with yellow
lateral spots on T II; in other species all dark, or
mainly to all red, yellow or orange. Pubescence often
concolorous with surface, most variable in density,

Hurkmans: Monograph of Merodon

evenness, length and colour; pruinose bands often
present, in many cases combined with yellow to red
background. These bands are quite variable in width
and curvature throughout Merodon and are lacking
altogether in several species. In the males S IV is usu-
ally slightly modified posteriorly; in some species
large paired appendages or a strong keel occur (figs.
36 b, 39 c, 40 b, 41 d, 42 e, 43 e, 44 b).

Male genitalia: The male genitalia are often intri-
cate and show enormous variation. They consist of
two main parts, the aedeagus and the surstyle. Both
parts are basally connected, by epandrium and basale
respectively, to the inside of the genital cap. The ae-
deagus is relatively the least variable, consisting of a
usually tapering shaft, which apically bears paired chi-
tinous plates, encasing the sperm duct. These plates
vary in shape, size and degree of sclerotization and
can be recumbent, erect or intermediate. The outer
face of the shaft may bear paired spines, lobes or
strongly sclerotized plates, or may have a subbasal ex-
cavation. The inner face may show pubescence. The
curvature of the aedeagus varies between species; both
slender and stout aedeagi occur, while the apical shaft
part (defined in fig. 10 a) may be weakly to strongly
lengthened. The surstyle which envelops the cercus is
much more variable. In its most simple form it con-
sists of a paired, more or less S-shaped sclerotized
plate, the apical part of which is identified as the pos-
terior surstyle lobe which is always present. In most
species additional lobes of the surstyle occur; often
there is an anterior surstyle lobe which may either be
free and projecting, or coalescent to various degrees
with the posterior surstyle lobe; in some cases a mid-
dle surstyle lobe is present as well. The anterior and
posterior surstyle lobes may bear accessory lobes and /
or specialized pubescences. The cercus is variable in
shape and in the length and density of the pubes-
cence. The genital cap (S X of Metcalf 1921, S VIII of
McAlpine 1981, T VIII of Speight 1987) encasing
the genitalia proper does not show much variability,
although in one species, Merodon caudatus Sack,
1913, this cap is strongly elongate and bears an apical
floccus conspicuous to the naked eye (figs. 8-17).

Female genitalia: Simple, consisting of a telescope
tube terminally fringed with pubescence (see figs. 52-
55 in Speight 1987). The structure of female genital-
ia is quite uniform throughout Merodon and of little
significance in establishing species groups or in spe-
cies recognition.

Sexual dimorphism: Rather pronounced in most
species. In species with dichoptic males, eyes of fe-
males wider apart. Metallic lustre in females usually
less strong. Wings in females often less infuscate. In
females pubescence on head, thorax and abdomen
shorter and duller, but in many species the pruinose
bands on thorax and abdomen clearer. In species with

191

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

males showing interalar pubescence bands these are
sometimes lacking in females.

Biology

Merodon is widely distributed over the Palaearctic
and Ethiopian regions and contains about 150 spe-
cies. In general, very little is known of the biology, in
spite of several species being quite common. Merodon
species may account for a considerable part of the
Syrphidae in the fauna of a region or country. There
are over 50 species of Merodon in Turkey, where ma-
ny species are very common. In Israel about 20% of
the Syrphid species belong to Merodon (Kaplan &
Thompson 1981: 198).

The known larvae all develop in underground
bulbs or rhizomes of monocotyledons. It is assumed
that most if not all species share this way of life.
Larvae were reared of Merodon amaryllidis Villeneuve
de Janti, 1934 (Villeneuve de Janti 1934), Merodon
bombiformis Hull, 1944 (Stuckenberg 1956) and
Merodon equestris (Fabricius, 1794) (e.g. Lindner
1949).

In the temperate zone the larvae survive the winter
underground. Various species occur in regions having
severe winters, e.g. the eastern Turkish mountains at
3000 m altitude, where species of Merodon are com-
mon. The larvae are sheltered by the snow cover. On
the other hand, Merodon occurs in tropical and sub-
tropical regions as well. In these regions the larvae
survive unfavourable climatic conditions, mainly
drought, in their sheltered underground habitat. In
relatively dry surroundings the various species seem
to prefer relatively wet places (Hurkmans 1988).
Some are mainly found at higher altitudes (e.g.
Hurkmans 1988, Marcos Garcia 1989) where lower
temperatures cause less evaporation, and therefore
less extreme drought. In view of these adaptations
Merodon should be considered xero-tolerant rather
than xerophilic. In the northern and moister parts of
the range, Merodon occurs in relatively drier habitats.
This probably reflects that dry areas warm up quicker
in spring and therefore allow a longer season. This is
important given the restricted flight period. In view
of the overall distribution, Merodon can be considered
thermophilous, but this holds true only for the flight
period. Cold season or drought conditions seem less
important in view of the sheltered conditions during
larval development.

The various species of Merodon appear to have a
preference for flowers of Umbelliferae (e.g. Suster
1959, Hurkmans 1985, 1988).

Territorial behaviour was recorded in several
Merodon species, e.g. by Fitzpatrick & Wellington
(1983) and Hurkmans (1985, 1988). Defence of ter-

ritories is more or less aggressive according to territo-

152

rial space available, presence of females, population
density and weather. In some species the males emit
piping sounds presumed to play a role in territorial
defence and mating (Conn 1978; Treiber 1987;
Hurkmans 1988).

Copulation is either airborne or sedentary; both
modes may occur in the same species (Conn 1978;
Hurkmans 1988) and may be linked with morpho-
logical differences (Conn 1978). Merodon equestris is
capable of prolonged flight in copula. Duration of
copulation is variable, from few seconds (Kabos
1939) to several minutes. In Merodon rufus copula-
tion is accompanied by a loud piping sound (Treiber
1987).

During oviposition several modes appear to occur:
Kabos (1939) recorded that in Merodon equestris eggs
were not attached to plants but pressed into the soil,
while Hurkmans (1988) found that in Merodon loewi
(Van der Goot, 1964) the females pressed their ab-
dominal tip to the stems of Ornithogalum plants as if
ovipositing; regrettably no larvae could be reared
from the plants dug out.

Apparently the number of generations per year var-
ies throughout the range. Northern records are main-
ly from July, while Mediterranean records of a single
species in one region may extend from May through
September. Marcos Garcia (1985) recorded that
Merodon avidus (Rossi, 1790), and Merodon aeneus
Meigen, 1822 are bivoltine in western-central Spain.
Split occurrences of many species reflect only avail-
able collection data and do not indicate that these
species are bivoltine.

Several species of Merodon have a dense and
colourful pubescence and mimic bumblebees by their
size and laborious flight, close to the soil. Brown
(1951) noted that in Merodon equestris the colour
forms most numerous in his samples mimicked the
most common bumblebee. A cluster of small, strong-
ly lustrous species resemble halictid bees in size, ap-
pearance and behaviour.

Division into species groups

The divisions of Merodon by Becker (1912), Sack
(1913a) and Paramonov (1926b) are based on extern-
al similarities. The genitalia were not considered
while some problematical species were not included
in the species groups presented.

The subgenus Exmerodon Becker, 1912, estab-
lished to accommodate the type-species [by original
designation], Merodon (Exmerodon) fulcratus Becker,
1912, is based on the dichoptic condition in males of
fulcratus. Sack (1913a, 1931) and Paramonov (1924-
1937) did not follow Becker’s subdivision. The sub-
division by Becker is not followed here for two rea-
sons: first, dichoptic males are also found in other

species not closely related to fulcratus and second, the
fulcratus group does not occupy an isolated position.

A first attempt to present a subdivision based on
synapomorphous characters, including those of the
male genitalia, is given here for the species having ar-
cuate pruinose bands and abdomina with tapering
second tergite. These two characters are judged to be
apomorphic. The corresponding plesiomorphic con-
ditions are: straight, oblique pruinose bands and non-
tapering abdomina.

Key to the species treated

Note: Several species had to be left out of the key.
Their genitalia could not be checked, since type-ma-
terial could not be studied. These are: Merodon bes-
sarabicus Paramonov, 1924 (the types are presumably
lost); Merodon dobrogensis Bradescu, 1982; Merodon
kiritshenkoi Stackelberg, 1960; Merodon tener Sack,
1913 (the only types left are 2, the d types are pre-
sumably lost).

Males: Key to species groups and species

A. No anterior surstyle lobe present on genitalia, or:
the surstyle consists of one mass in which the an-
terior part may be differentiated (e.g. figs.); or:
the outline of the surstyle ridge is fluent, semicir-
cular or broadly rounded; or. anterior surstyle lo-
be weak, posterior lobe very long (African spe-
cies); or: anterior surstyle lobe medially recurved,
totally coalescent with surstyle mass; or: three sur-
style lobes of equal size, separated by two very
deep sulci; or : abdomen relatively stout, T II not
tapering, pruinose bands parallel to posterior ter-
gite margins ... key in part II of this monograph

— Anterior surstyle lobe well-defined, moderately
large to large, the surstyle lobes are well differen-
tiated. The surstyle lobes are not all separated (by
two deep sulci), if a middle lobe occurs it is small
and coalescent with either anterior or posterior
lobe, and the lobes are not of equal size. Anterior
lobe not medially recurved and totally coalescent
with stylus mass, but free (although the apex may
be recurved). Abdomen relatively slender, T II
(more or less strongly) posteriorly tapering, abdo-
minal pruinose bands arcuate. Species fulfilling
all these requirements can be identified using the
following key.

1. Anterior surstyle lobe short, rounded, pubescent,
coalescent with surstyle mass (distinctus assem-
beretta 12

— Anterior surstyle lobe more elongate, often well
separated from surstyle mass, often with accesso-
malobes or. specialized. parts"... 2

2. Anterior surstyle lobe narrow, posterior lobe me-

10.

11.

122

Hurkmans: Monograph of Merodon

diolaterally aligned, often overlapping (alexeji
Te) ioni rari 17
Posterior lobe dorsoventrally aligned ................ 3
Basitarsi 3 swollen, with specialized (dense, short,
golden) pubescence on medial face; tibiae 3 often
with apical floccus; sternite IV often keeled or

strongly modified (tarsatus group) … 25
Sternite IV never strongly modified, if keeled the
basifarsy3imormal ae a 4

Legs all dark, many species with conspicuous pu-
bescence; anterior surstyle lobe wide, elongate,

Er (A OD) EE 32
Wepsiparthy yellow. "2... 7 a 5
Flanges present on outer face of aedeagus (elegans
PROUD) jets one eher aen seems’ 42

No such flanges present
Secondary, lateral lobe present on posterior sur-
style lobe; femora 3 much swollen, tibiae and tar-
si all yellow; anterior surstyle lobe large, rather el-
ongate, moderately wide; aedeagus with apical
shaft part normal (vandergooti group) .................
RE Re DEL vandergooti sp. n.
No secondary lateral lobe, or if present, femora 3
less swollen, anterior surstyle lobe narrow, elon-
gate, ventrally recurved, anterior surstyle lobe
narrow, ventrally recurved, apical shaft part of ae-

deagus)lenethenede. 2. u... 2 0 002 7
Anterior surstyle lobe separated from surstyle
mass/by.a sulcuss-. dda 8
Genitalia not with sulcate surstylus ................. 9

Large species, averaging 17 - 21 mm body length;
abdomen predominantly yellow, antennae nor-
mali(przzeronp) een pruni Rossi
Smaller species, 7 - 15 mm, abdomen dark with
crimson or dark red, antennae notably to ex-
tremely lengthened (longicornis group) ........... 45
Posterior surstyle lobe with large, ventral accesso-
walobeilerasssfemzonseroup) 22
ee crassifemoris Paramonov
This lobe without large ventral accessory lobe ....

Apical shaft part of aedeagus lengthened; anterior
surstyle lobe usually not apically widened but if

so, then bifid (alagoezicus group) .................... 46
Apical shaft part normal, anterior surstyle lobe
apically widened, mot bifid 2 11

Aedeagus with 2 pairs of spines on outer face, an-
terior surstyle lobe rectangular to diamond-
shaped (nigritarsis group)
Aedeagus smooth, anterior surstyle lobe ending
in spathulate-semicircular apex (avidus group) ...
EE EE er avidus (Rossi)
Femora 3 very much swollen .......................... 13
Femora 3 only moderately swollen ................. 14

153

15

14.

15:

16.

17.

20.

21.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Larger species, body length 13 - 15 mm; genital-
ia: aedeagus stouter, apical shaft of aedeagus
shorter
Smaller species, body length 10 - 13 mm; genital-
ia: aedeagus slenderer, apical shaft of aedeagus
JON LEE nn clunipes Sack
Thorax and abdomen densely yellow pubescent;
abdomen stout, T II strongly tapering; genitalia:
both surstyle lobes elongate; body length 15 mm
SIREN ALTRO Jee ull Aa. EEA ROIO mariae sp. n.
Pubescence less dense, abdomen slenderer; if
body length approx. 15 mm then abdomen very
slender
Genitalia: Posterior lobe very short; third anten-
nal article subacute - acute; pubescence overall
ver long ane) ASU Sono ottomanus sp. n.
Genitalia: Posterior lobe longer; third antennal
ATE CIETODTUSE e O PIE Eee .16
Body length 13 - 15 mm; both surstyle lobes co-
alescent; abdomen very slender, dark with large
yellow lateral spots ..................... distinctus Palma
Body length 9-12 mm; surstyle lobes separated by
a sulcus; abdomen yellow throughout ................
RR A]: NEE ENE AEE testaceoides sp. n.
Genitalia: Anterior surstyle lobe showing elon-
gate bald area; S IV deeply emarginate posterior-
lyrics dee marginicornis sp. n.
Anterior surstyle lobe without bald area ......... 18

. Genitalia: Posterior surstyle lobe with rounded

bald knob in antero-lateral part ...................... 19
No such knob present, or knob very weak ......21

. Both surstyle lobes short; cercus relatively large,

short,wide; T II with orange lateral spots bearing
submetallic lustre; apical part of aedeagus covered
by overlapping posterior surstyle lobes ...............
star «ple ERLE ARENA RE ch Po ER sophron sp. n.
Both surstyle lobes longer, cercus relatively small-
er, more elongate; no submetallic lustre on lateral
spots; aedeagus entirely visible ........................ 20
Pubescence on thorax and abdomen recumbent
to oblique, golden; triangular processus with clear
apical spur and short stout bristles .....................
BAALDE VUS. RER SALE RR altinosus sp. n.
Pubescence on thorax and abdomen less conspic-
uous, erect; no clear apical spur on triangular pro-
cessus which bears longer, less stout bristles … …
SARE SOIA IV) SACS TIGRI e alexeji Paramonov
Anterior surstyle lobe with length/width ratio 1.2
- 1.3 in lateral view; posterior surstyle lobe short,
rather spathulate in ventral view, coalescent … …
HAT ALARM LU ZM kawamurae Matsumura
Anterior surstyle lobe with length/width ratio of
2.0 or over; if posterior surstyle lobe short: acces-
sory lobe present where anterior and posterior
lobes meet, and anterior lobe strongly spathulate

DIA

23:

24.

DS)

26.

29)

30.

Antennal ratio 2.1; genitalia: cercus placed far
ventrad; accessory lobe present on posterior sur-
style lobe; anterior lobe very elongate (an incon-
spicuous bald knob present antero-medially on
posterior surstyle lobe) ........... alexeji Paramonov
Antennal ratio under 2.0; genitalia: cercus in nor-
mal position; no accessory lobe on posterior sur-
style loef e ee 23
Abdomen with strong golden lustre; genitalia: an-
terior surstyle lobe widest in its mid-region; ae-
deagus with two separate subapical cavities, apical
shaft part slightly lengthened ...4ypochrysos sp. n.
Abdomen without strong golden lustre; genitalia:
anterior surstyle lobe equally wide throughout, or
spathulate; aedeagus with one subapical cavity ...
D NE A I ah ak I Re Pre cd Ai MEE nti oa 24
Touchline of compound eyes incomplete, very
short; all legs with tarsi orange, tibiae orange with
wide dark distal band; genitalia: anterior surstyle
lobe equally wide throughout .....rufitarsis (Sack)
Touchline complete, tl-v ratio 0.9; all legs dark
except joints; genitalia: anterior lobe somewhat
Spatial ateo en an hirsutus Sack
Genital cap much elongate, with floccus ............

ARI ESTATE WERE eRe Clero le en caudatus Sack

S IV normal, slightly arched, emarginate posteri-
orly; basitarsi 3 with length/width ratio 2.5 and
dense, short, golden pubescence on lower face;
clear apical floccus present on tibiae 3 ................
PrO Lo A ri ri diga risi ia auronitens Sp. n.
S IV with small (single or paired) median projec-

ODA UE i SE ILE iis ilo Di
S IV with large paired appendages … 29
. Antennal ratio over 2.8 ............ tangerensis sp. n.
Antennal ratio under 2.3 ............................... 28

. Antennal ratio approx. 2.1; S IV with small pro-

jections; legs pale brown ........... xanthipous sp. n.
Antennal ratio approx. 1.3; S IV with single me-
dian projection ending in acute spine .................
deste Re REES ankylogaster sp. n.
S IV with long slender appendages; posterior
proximal face of basitarsus 3 showing ovoid area
of widely spaced setaceous pubescence bordered
by long setae; apical floccus of tibiae 3 inserted on
ENON PUS ee tarsatus Sack
S IV with less long, robust appendages; if special-
ized area present on basitarsi 3, pubescence in
centre and on margin similar... 30
Genitalia: anterior surstyle lobe widening apical-
ly, angular, its apical part projecting ventrad,
with a fringe of short yellow pubescence

PR Se nan tried enne persicus Sp. n.
Genitalia: anterior lobe not widened apically, not
Projecting MENACE een ee een 31

3.

SPA

34.

>:

36:

STA

38.

32.

Head: antennal ratio over 2.0; leg 3: length/width
ratio of basitarsus 2.0; genitalia: anterior surstyle
lobe tapering apically; aedeagus rather stout .......
MORN ce turkestanicus Paramonov
Head: antennal ratio approx. 1.6; leg 3: length/
width ratio of basitarsus 1.4; genitalia: anterior
surstyle lobe slightly spathulate; aedeagus slender-
CO? iy ee As. oidipous sp. n.
Wings strongly infuscate, deep brown except api-
cally; large species (body length 15 - 18 mm)
e antennen due crie velox Loew
Wings clear or slightly infuscate (yellowish at
most)

. Thorax and abdomen densely pubescent, obliter-

ating background colour; pubescence often red-

an Or VELO nue 38
Thorax and abdomen moderately dense pubes-
CENT surface well visiblen ev overs ten 34

Pubescence on thorax uniform; relatively stout
species, with T II only moderately tapering ....35
Pubescence on thorax with dark interalar band; T
II strongly tapering, slenderer species
Genitalia: anterior surstyle lobe with long stalk;
posterior lobe anteriorly with a low well-defined
projection bearing dense pubescence; aedeagus
rather stout; body length 14-15 mm
SIR RC E warnckei sp. n.
Genitalia: anterior surstyle lobe with short stalk;
dense hair-tuft anteriorly on posterior lobe not
inserted on a projection; aedeagus slender; body
lede mmie brevis Paramonov
Triangular processus modified, ending in large
hooked spur; 3rd antennal article orange; Body
ENEN na hamifer Sack
Triangular processus on femora 3 normally
shaped; abdomen slender, with bronze lustre;
body length 10-15 mm
Body length 11 - 15 mm; abdomen slender, with
bronze lustre; pubescence yellow, rather dense;
Éosdanithroughoutt een. na a ne
BEER u... aberrans ssp. aberrans Egger
Body length 12 - 14 mm; abdomen slender, with
bronze lustre; legs with yellow markings ............
Pate mere aberrans ssp. flavitibius Paramonov
Body length 10 - 11 mm; abdomen extremely
slender; with slate grey lustre and often purple re-
flections; pubescence grey to greyish yellow,
rather sparse; legs dark throughout .....................
RE aankaart ledee aberrans ssp. isperensis ssp. n.
Large species (body length 17 - 22 mm) .............
PPP RAMA Se eo sise nota clavipes Fabricius
Smaller species (body length up to 16 mm) ....39
Femora 3 normally swollen; triangular processus
with a strong, erect, apical spur; body length 12 -
Bani ei vad ch Pate tte 40
Femora 3 very much swollen; triangular proces-

40.

41.

42.

43.

44.

. Anterior surstyle lobe recurved mediad

Hurkmans: Monograph of Merodon

sus without apical spur, or this spur low, stout;
body length 14 - 16 mm
Tarsus 3: stout, with first article twice as long as
second; apical spur of triangular processus on fe-
mora 3 shorter; pruinose band on T IV chevron-
shapeda ss karadaghensis Zimina
Tarsus 3: slender, first article 1.4 x as long as sec-
ond; apical spur of triangular processus on femo-
ra 3 erect, longer; pruinose band on T IV round-
[N es Cee dzhalitae Paramonov
Antennal ratio 1.4; body length 14 - 16 mm; tri-
angular processus on femora 3 finely serrate, nor-
MARS. armadi. orione rn cupreus sp. n.
Antennal ratio 1.8; body length 13 mm; triangu-
lar processus strong, with slight apical spur ........
tee splendens sp. n.
Anterior surstyle lobe with rounded profile; sul-
cus between both surstyle lobes marked, but not
very deep or wide; aedeagus rather stout
Anterior surstyle lobe with angular profile; sulcus
between both surstyle lobes very wide; aedeagus
slenderer
Head: oa 35°; abdomen very slender; anterior
surstyle lobe with lateral ridge; flanges on outer
ace orales one 0 testaceus Sack
Head: oa 50°; abdomen moderately slender; an-
terior surstyle lobe without lateral ridge; flanges
onjaedeapusiighene See manicatus Sack
Pruinose bands on abdomen conspicuous; S IV
strongly arched; triangular processus on femora 3
bearing 6 - 9 bristles; posterior surstyle lobe with
free anterior part separated by an oblique sulcus;
cercus long, well projecting posteriorly ...............
Resana elegans sp. n.
Pruinose bands inconspicuous but visible; S IV
strongly arched; triangular processus bearing 10
bristles; posterior surstyle lobe without free ante-
rior part; sulcus rather quadrate; cercus shorter,
not much projecting ................... bequaerti sp. n.

. Antennal ratio 4.0 - 4.5 .............. longicornis Sack

Antennal ratio 2.5 - 3.0 kaloceros sp. n.

. Anterior surstyle lobe stalked, widening apically

Anterior surstyle lobe narrow throughout

. Both surstyle lobes separated by a deep sulcus...

EE TR A taniniensis sp. n.
Surstyle lobes coalescent; anterior lobe bifid .......
SR SR En eee PESTE toscanus sp. n.
stia i n nitidifrons sp. n.

Anterior lobe recurved ventrad to ventro-caudad

. Anterior lobe recurved ventro-caudad, apical part

more or less parallel with basal part ................ 50
Anterior lobe recurved ventrad, apical part pro-
HAINE entri elet sente doge Sil

50:

il

52:

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Tibiae 3 without large distal spur; posterior sur-
style lobe with semicircular ventral projection …
Secs ntl FEN PR LEHE lucasi sp. n.
Tibiae 3 with large, triangular spur; posterior
lobe without semicircular projection; lateral spots
on T II with postero-lateral digitiform dark in-
CENONE ccc ies alagoezicus Paramonov
Anterior surstyle lobe simple; surstyle guarded by
large lateral sclerotized plate, projecting far cau-
dad; posterior surstyle lobe straight; aedeagus ba-
SAIL eee eres en nn schachti sp. n.
Anterior lobe with accessory lobe; no guarding
lateral plate; posterior lobe twisted; aedeagus
slender throughout ................... satdagensis sp. n.
Anterior surstyle lobe with apical part rhomboid-
al; posterior lobe higher; aedeagus with outer face
all convex; trochanter 3 often with clear knob;
rather robust species … … … … nigritarsis Rondani
Anterior surstyle lobe suddenly widened apically,
profile of apical part quadrate; posterior lobe low-
er; aedeagus concave on basal outer face; tro-
chanter 3 smooth; slenderer species

Key to females of species

1.

Abdomen dark throughout on surface (the pu-
bescence may be brightly coloured!), often metal-
lic lustrous, without red or yellow lateral spots .

Abdomen showing yellow, orange or red lateral
spots on T II, or with even more extensive yellow
or red on abdomen
Legs all dark, at most femoral apices and tibial
bases very narrowly paler (but check 7: M. aber-
rans ssp. flavitibius)
Legs with yellow markings
Pruinose bands present on abdomen, pubescence

on posterior abdomen concolorous .................. 4
No pruinose bands present on abdomen ......... 10
Abdomen slenderer, in some cases with clear
BISI eee 5

Abdomen stouter, dark, not clearly bluish lus-
trous (but if clear grey lustre is present, see 7:

AUCTIGIESSS PENISDENETISL) SORTE eee 8
Femoral ore LEALYASWOILENE ese eee ee 6
Femora 3 normal, not much swollen ................ 7

Abdominal surface colour brownish, with nar-
rower pruinose bands ..................... clunipes Sack
Abdominal surface colour blackish, with wider

Abdomen with bluish metallic lustre, thorax of-
ten with inconspicuous pruinose bands; wings
apically slightly infuscate; femora straight; trian-
gular processus normally shaped; body length 13
- 15 mm (in the ssp. isperensis the metallic lustre

156

10.

16.

We

. Thorax with pruinose bands

is more greyish, while the abdomen is slenderer
than in the nominal subspecies; the ssp. flavitibi-
us, of which no records exist postdating the pub-
lication of the original description, shows yellow

zend mes). EE aberrans Egger
Habitus similar, triangular processus ending in
larger hacken en hamifer Sack

Thoracic pubescence unicolorous, species of
some 12 mm, bands on abdomen rather clear,
nearly continuous, paired pruinose spots on T II
BS LR oA a vain Pelee ti es hirsutus Sack
Thoracic pubescence bicolorous
Thoracic pubescence grey, dark interalar pubes-
CEN CED AND LES speoacodacnaconcecanesse cupreus sp. n.
Thoracic pubescence golden, but clearly paler
posteriorly; relatively stout species with marked
apical spur on triangular processus .… …
de nis och haben ee karadaghensis Zimina
Larger species, over 12 mm, densely pubescent at
least on thorax, abdomen dark, blackish pubes-
cent, with 2 pairs of outstanding, snow-white pu-
DESCESDOL n quadrinotatus (Sack)
Smaller species, sparsely pubescent throughout,
slight bluish lustre on thorax and abdomen, basi-
tarsus 3 swollen; body length 7.5 - 9 mm ...........
Berane I tert aie turkestanicus Paramonov

Thorax without bands, tibiae and tarsi partially
dark, basitarsi 3 rather swollen

. Species strikingly black-and-white with strong

pruinose bands on abdomen
Duller coloured species, bands inconspicuous ....
ee hal warnckei sp. n.

. Bluish lustre on thorax and abdomen, pruinose

abdominal bands nearly continuous
el le Air me nal ankylogaster sp. n.
No bluish lustre, bands interrupted ....................

. Well marked whitish pruinose bands present on

HOTA rosie tee cee eet ee ee nee 15
Pruinose bands on thorax very weak, or longitu-
dinal metallic bands pe or bands of different
OO ee eee 32
Thorax unicolorous, without traces of bands...36

. Tergite II posteriorly strongly tapering, thorax

with strong pruinose bands and extremely short,
even pubescence; habitus very slender, body
IRR n 16
Tergite II less tapering or habitus stouter, or
smaller or larger species, thoracic pubescence
longer sn eee ere er er 107
Femora 3 reddish to orange; midstripe on frons
ExXtremielyznanrow ee distinctus Palma
Femora 3 dark, midstripe normal .......................
"atea biarcuatus Curran

19:

Dijk

28.

29.

Antennal ratio 2.0 or less nn. 21

. Abdomen relatively stout, tapering posteriorly ...

Abdomen slenderer, more cylindrical, lateral
spots extended over much of 2nd tergite at least,
in many cases over 3rd as well, crimson coloured

Femora slender, abdomen with grey hue pro-
duced by overall whitish pubescence; no blue lus-
tre on abdomen … kaloceros sp. n.
Femora swollen, abdomen dark with bluish lustre
MM ee eee lusitanicus sp. n.

. Antennal ratio 3.4 to 5.0 ............ longicornis Sack

Antennal ratio 2.2 to 2.8 ..erivanicus Paramonov

MSec ansthrouglioltss. tnt tn... 22

Legs yellow marked, or all yellow/orange ....... 25

. Large, robust species (17 mm), wings often infus-

cate, thorax with dense pubescence ...velox Loew
Smaller species (8 - 11 mm), wings clear ............
e ian alexeji Paramonov

. Large species (17 - 21 mm), wings slightly infus-

cate, abdomen clearly conoid in outline .............

MEI et. pruni Rossi
Smaller species up to 15 mm, abdomen more cy-
Fina bce En no. via 24

. Abdomen mainly pale orange, slender, pruinose

bands on abdomen arcuate, posterior end of ab-

CPU ER testaceus Sack
Nbdomcnimamiyidark in 25
. Antennal ratio 2.0, triangular processus strongly
SOM nt affinis Gil Collado
Arena ton NME 26

. Tarsi 3 with dark upper face and contrasting,

golden pubescence on sides; pruinose bands on
thorax inconspicuous ............ nigritarsis Rondani
Tarsi without this contrast or if so, thorax with
RISAIE ANAS init 27
Legs nearly all orange ............... aureotibia sp. n.
Legs with extensive dark surface .................... 28
Abdomen relatively lustrous, thorax with strong
pruinose bands, often a fifth anteromedial band
BESSER een alagoezicus group
Comprising the following species: alagoezicus
Paramonov, /ucasi sp. n., nitidifrons sp. n., satda-
gensis sp. n., schachti SP. n., taniniensis sp. n. and
toscanus sp. n. These species cannot at present be
well separated in the female sex, as the pairing
with the males is not certain. M. toscanus is
known only from Italy, the other species are
mainly known from Turkey.

Abdomen duller, usually no 5th band on thorax

DO a oe ae dee ARRE La A TR (RE RA CRT COREE 29
Small species, about 10 mm body length, thorax
clearly but weakly banded .................. tener Sack
Larger species, 12 mm or over, bands more
men Si EN Sh DE a 30

Hurkmans: Monograph of Merodon

30. Frontal depression present; 2nd tergite bright
CNEL Tobey, ERRO Mae EOS elegans sp. n.
= Naldepressio MON {[0Nst ete ee ek ne, 31
Sle Femorasiswolleny curved nse eee
PRE SS OLE ITER BaD femoratoides (Paramonov)

— Femora 3 less swollen, straighter … avidus Rossi
S2igltepsallidariso ne ce, pies 33
— Legs at least with yellow markings, thorax rather

dulledark aero, A RE SIA 34

33. Femora 3 strongly swollen, abdomen bluish lus-
trous, body length 10 - 13 mm ./usitanicus sp. n.
— Femora3 slenderer, if in doubt: no blue abdomi-
nal lustre; approx. 17 mm, wings infuscate ........

BERS | Anche MARI Pee Ne SERRE A VAE. VE velox Loew
34. Large species, 17 - 22 mm, wings often infuscate
A TT SIRIA SIRO DS NLL PES pruni Rossi
VAI E 35

35. Abdomen mainly orange, legs dark .....................
PO LEA DEM SIRIA manicatus Sack
— Abdomen mainly dark, third antennal article
slender, acute to subacute, size 7.5 - 10 mm, prui-
nose abdominal bands conspicuous ....................
aE eee cate he kawamurae Matsumura
36. Lateral spots on abdomen fiery orange or red,
basitarsi 3 widened ........................ caudatus Sack
— Lateral spots less conspicuous, abdominal pubes-
cence bands clear, large to very large species ...37
37. Large species, 12 - 15 mm, abdomen stout with
parallel, often yellow pruinose, conspicuous
Dandie esis ie MN Ro mariae sp. n.
— Very large species, 17 - 22 mm, bands whitish,
less conspicuous, or dense pubescence obliterat-

ID ED EUROS EE e eno clavipes (Fabricius)

SYSTEMATIC PART

The distinctus assemblage

This set is a paraphyletic group sharing no other
than the ‘general’ synapomorphies of this section of
the genus, viz. the tapering T II and arcuate pruinose

bands.

Merodon biarcuatus Curran (figs. 21 a-c, 103)

Merodon biarcuatus Curran, 1939: 7. Holotype 9 : ‘Forest of
Namora, Robot, Morocco’ [near Rabat, approx. 33° 55’
N, 7° W] (AMNH) [examined].

Material examined. — France: 22 (BMNH, RMNH);
Greece: 19 (ZMAN); Italy: 32 (NHMW, ZMAN, WH);
Morocco: 22 (RMNH); Turkey: 122 (JLR, ZMAN, BMNH,
WH).

Description
Female. — Head: Antennae orange, antenna 3 with
upper margin convex, apex rounded, antennal ratio

157

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

2.2, ocellar angle 55°; pubescence pale yellow, rather
sparse, erect, but moderately dense and darker on me-
dian, lustrous 1/3-1/6 of frons and on vertex; face
densely pruinose except on narrow median line.

Thorax: Katepisternum, anepisternum and ane-
pimeron, lateral dorsum and scutellum lustrous, bear-
ing very short yellow even oblique pubescence; dark
intaralar band present. Wings clear; halteres, squamae
and antisquamae pale orange to yellow.

Legs: Trochanters brown, femora dark, tibiae and
tarsi orange to yellow; pubescence yellow, trochanter
3 smooth, femora 3 swollen with normal triangular
processus bearing 7-9 bristles on serrate distal margin,
apical bristle conspicuous.

Abdomen: Extremely slender; T II orange, some-
times with small dark medio-anterior lunule, strongly
tapering; T III with conspicuous wide arcuate whitish
pruinose band, background orange anterior, dark
posterior to band; T IV-V dark, with conspicuous ar-
cuate pruinose band on T IV; pubescence on all ter-
gites concolorous to pale throughout.

Body length 13-17 mm.

Male. — Unknown.

Diagnosis. — The extremely slender habitus will
distinguish this species from other species, but M. bi-
arcuatus is apparently closely related to distinctus. The
pubescence on the frons of distinctus female is denser,
oblique and golden; the lustrous midstripe is narrow-
er, occupying 1/8 of its width; the pubescence on the
thoracic dorsum is darker and shorter; the femora 3
are orange (dark in biarcuatus); the pruinose bands
on the abdomen are narrower.

Period of flight and distribution (fig. 103). — This
species seems to occur throughout the mediterranean
where it flies from May to August.

Merodon clunipes Sack
(figs. 22 a-c, 104)

Merodon clunipes Sack, 1913a: 444. d Holotype: ‘Smyrna
49417’ ‘Billar (on lower face of same label) [= Izmir,
Turkey] (ZMHB) [examined].

Merodon clunipes, Peck 1988: 169.

Lampetia clunipes, Sack 1931: 311.

Material examined. — Austria: 1d, 19 (RMNH); France:
154,29 (RMNH); Greece: 156,59 (NHMW, KBIN, RMNH,
JLR, WH, MNHN); Hungary: 12 (RMNH); Italy: 146, 79
(NHMW, RMNH); Lebanon: 2d, 12 (NHMW, KBIN); Spain:
13 (RMNH); Switzerland: 1d (RMNH); Turkey: 148, 129
(BMNH, NHMW, ZMAN); Yugoslavia: 10d, 19 (RMNH).

Description
Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apex rounded, antennal ratio 1.2,

158

pubescence dense, especially on face, erect, pale yel-
low; vertex angle 35°, ocellar angle 45°, tl-v ratio 0.5.

Thorax: Dark, without conspicuous lustre; pubes-
cence dull yellow, pruinose bands clear. Wings very
slightly infuscate; halteres, squamae and antisquamae
yellow.

Legs: Femora dark, tibiae 1-2 apically brownish,
tibiae otherwise dark; tarsi brownish, darker apically;
trochanter 3 with ridged projection; femora 3 very
much swollen, curved, with normally shaped triangu-
lar processus bearing 8-9 bristles on strongly serrate
distal margin.

Abdomen: Dark; slender to very slender; T II-IV
bearing arcuate somewhat vague rather wide pruinose
bands; pubescence dark, paler laterally and on T IV;
densest on pruinose bands of T II; S IV narrowly
emarginate posteriorly.

Genitalia: Anterior surstyle lobe rather short,
rounded, bearing dense, short, yellow, even pubes-
cence throughout, showing a ridge bearing 5-6 strong
bristles on medial face; posterior surstyle lobe coales-
cent, elongate, posteriorly with dense, moderately
long, yellow pubescence. Cercus rounded, bearing
long, yellow pubescence. Aedeagus moderately long,
apical shaft part slightly lengthened, fringed plates on
thecal apex recumbent.

Body length 8.5-13 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Ocellar angle 50°,
frons with lustrous midstripe occupying 1/3 of width.
Thorax: Pruinose bands quite conspicuous, pubes-
cence less dense, rather recumbent on sides. Legs:
Apices of femora paler, femora 3 less swollen, but if
compared to females of other Merodon species still
very much so. Abdomen: T II largely reddish to
orange laterally, this colour may extend over T III as
well; dark parts brown, not black.

Body length 9-12 mm.

Diagnosis. — The males of this species are rather
similar to femoratus from which they differ in being
smaller (range 8.5-13 vs. 12-16 mm) and having a
more slender aedeagus with slightly lengthened apical
shaft part. In the females the size ranges are 9-12 and
12-17 mm in clunipes and femoratus, respectively; the
females of clunipes have a more slender abdomen with
clear orange markings (all dark in femoratus females)
with vague bands; the bands in the females of femor-
atus are conspicuous and sharply bordered.

Variation. — The ridge on trochanter 3 is rather
variable in size, and seems correlated with specimen
size: small specimens have small ridges, large speci-
mens having large ones.

Period of flight and distribution (fig. 104). — This

species occurs through central Europe and the north-
ern Mediterranean where it flies in June and July.

Discussion. — One of the NHMW specimens identi-
fied as clunipes has an identification label ‘M. striatus
Sack det. Sack’. As far as could be traced, this name is

unpublished.

Merodon distinctus Palma
(figs. 23 a-d, 103)

Merodon distinctus Palma, 1863: 46. Holotype d : ‘from the
field of Sanseverino’ [in the vicinity of Naples, Italy] [not
located, not examined].

Lampetia dimorpha Szilady, 1940: 66. Holotype d: (and 9
Paratype) ‘Karacabey bei Brussa’ [West of Bursa, Turkey
on coast of Marmara inland sea] [lost, formerly in TMA,
not examined]. Syn. n.

Merodon distinctus, Rondani 1868a: 22; Sack 1913a: 444;
Peck 1988: 169.

Lampetia distincta, Sack 1931: 313.

Material examined. — France: 16 (BMNH); Greece: 236,
62 (KBIN, RMNH, BMNH, NHMW, JLR, WH); Italy: 16
(BMNH); Turkey: 52 (LR); U. S. S. R.: 12 (RMNH);
Yugoslavia: 1d, 22 (LR, RMNH).

Description

Male. — Head: Antennae bright orange, antenna 3
with upper margin convex, apex obtuse, antennal ra-
tio 1.5, pubescence golden with dark tuft in ocellar
region; vertex angle 40°, ocellar angle 45°, tl-v ratio
0.6.

Thorax: Dark, velvety lustrous on sides produced
by recumbent pubescence, with clear pruinose bands
on dorsum; pubescence golden, short, even. Wings
clear; halteres, squamae and antisquamae yellow.

Legs: All femora dark, not black, femora 1-2 apical-
ly often brown to orangish; tibiae and tarsi yellow,
legs with conspicuous golden pubescence through-
out; trochanter 3 bearing distinct knob with apical
obtuse ridge; femora 3 slightly swollen, rather
straight, triangular processus with serrate distal mar-
gin; in some specimens vague orange spot present ba-
sally on femora 3; tibiae 3 rather abruptly widened
distally (best seen in posterior view).

Abdomen: Slender; T II-IV dark with rather wide
sharply defined arcuate yellowish pruinose bands,
narrowly to hardly interrupted, the band on T IV
widened laterally and in some specimens slightly re-
curved; T II with conspicuous yellow lateral spots;
pubescence concolorous with surface; sternites I-III
yellowish orange, IV darker, narrowly emarginate
posteriorly, clearly though not strongly vaulted.

Genitalia: The anterior surstyle lobe short, round-
ed, with even, yellow, short, dense pubescence, co-
alescent with posterior surstyle lobe which is low and

Hurkmans: Monograph of Merodon

elongate and bears long, erect, yellow pubescence.
Cercus rounded to obtusely angular, with same pu-
bescence. Aedeagus rather stout, slightly tapering, ap-
ical shaft part somewhat lengthened, fringed plates on
thecal apex rather wide and recumbent.

Body length 12-16 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Midstripe on frons
occupying 1/8 of width or even less; antenna 3 with
dark upper margin in some specimens. Thorax:
Pruinose bands more conspicuous, a thin praesutural
crossband present in some specimens; pubescence ex-
tremely short, as if shaven; pubescence in mid region
of dorsum often dark. Legs: Deep orange to yellow all
through, but triangular processus often darker.
Abdomen: Even more slender than in d ; lateral spots
larger, deeper coloured, pruinose bands less wide; T V
bearing arcuate wide pubescence band; pubescence
shorter than in d.

Body length 13-15 mm.

Diagnosis. — The combination of the very slender
habitus, clear, sharply bordered pruinose bands, large
lateral spots and genital structure will easily separate
the males of distinctus from any other Merodon. The
females are separated from the females of most other
species by their relatively large size and extremely
slender habitus. They can be separated from the fe-
males of the alagoezicus group by the more cylindrical
abdomen, and from the very similar females of biar-
cuatus by the narrower frontal midstripe and the
orange femora 3.

Period of flight and distribution (fig. 103). — M.
distinctus occurs from May through August in the
mediterranean parts of Europe and Turkey.

Discussion. — M. dimorphus Szilady, 1940 is tenta-
tively considered a synonym of distinctus. In the orig-
inal description dimorphus is stated to be allied to clu-
nipes Sack, but the description points to very apparent
differences with that species. From the description it
can be inferred that dimorphus is very similar to, if not
conspecific with, distinctus, the only difference appar-
ent from the description is the colour of the pubes-
cence on the head which is said to be pale in dimor-
phus. Regrettably several features are not or very
superficially described; the ‘spots’ on the abdomen
may either be lateral spots or pruinose bands, or both.
The figure of the thoracic dorsum as given by Szilady
closely resembles this element in distinctus. No mate-
rial of dimorphus was examined, as the type-material
(formerly in TMA), the only material known, was lost
in 1956 (Dr. A. Dély-Draskovits, in litt.).

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Merodon femoratus Sack
(figs. 24 a-b, 104)

Merodon femoratus Sack, 1913a: 446. Syntypes: ‘Korsika,
Griechenland und Kleinasien’ [Corsica, Greece and
Turkey]; depository unknown [not examined].

Merodon femoratus, Van der Goot 1981: 216, 218; Peck
1988: 169.

Lampetia femorata, Sack 1931: 315; Séguy 1961: 179.

Material examined. — Greece: 26, 52 (ZMUC, BMNH,
KBIN, ZMAN, WH); Italy: 48 (ZMAN); Turkey: 156, 359
(JLR, BMNH, ZMAN, WH).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apex obtuse, antennal ratio 1.8;
pubescence pale yellow, face and occiput densely
white pruinose; vertex angle 35°, ocellar angle 45°,
tl-v ratio 0.5.

Thorax: Dark, katepisternum, anepisternum, ane-
pimeron, lateral dorsum and scutellum metallic lus-
trous, hardly pruinose except four longitudinal bands
on dorsum; pubescence pale brown, moderately
dense, erect, more even on dorsum but less so on
sides. Wings very slightly infuscate; halteres, squamae
and antisquamae pale brown.

Legs: Dark, except for apices of femora and bases of
tibiae of legs 1-2; trochanter 3 showing conspicuous
hump, femora 3 very strongly swollen, strongly
curved, with normally shaped triangular processus
with strongly serrate, sometimes undulating margin
bearing 5-10 bristles.

Abdomen: Dark; rather slender; T II tapering,
without lateral spots; T II-IV with interrupted mod-
erately wide strongly arcuate whitish pruinose bands;
pubescence concolorous, densest laterally on T II; S
IV strongly vaulted, deeply narrowly emarginate pos-
teriorly.

Genitalia: The anterior surstyle lobe short, round-
ed; bearing even, erect, short, dense, yellow pubes-
cence apically; coalescent with posterior surstyle lobe,
which shows long yellow pubescence posteriorly, is
elongate and medio-laterally aligned apically. Cercus
wide, short, rounded, not far protruding beyond pos-
terior surstyle lobe, bearing dense, long, yellow pu-
bescence. Aedeagus short, stout, smooth on outer
face, fringed plates on thecal apex rather wide, recum-
bent.

Body length 12-16 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Ocellar angle = 50°
to 60°, frons with lustrous midstripe occupying 1/4
of width. Thorax: Pruinose bands wider and clearer,
often striking. Abdomen: Distinctly wider, typically
black-and-white patterned, pruinose bands slightly
less arcuate.

160

Body length 12-17 mm.

Diagnosis. — M. femoratus is nearly similar to clu-
nipes from which it is distinguished in both sexes by
the greater size (12-17 mm vs. 9-12 in clunipes), in
the males by the stouter aedeagus, and in the females
by the black and white abdomen (brownish red in
clunipes). From other species it differs in having a
combination of very strongly swollen and curved fe-
mora 3 and very simple anterior surstyle lobe in the
males, and by a combination of strongly curved and
swollen femora 3 and a striking black-and-white ab-
domen in the females.

Period of flight and distribution (fig. 104). — M. fe-
moratus occurs in southern Europe and Turkey from
May through August, both in lowland and mountain
habitats (up to over 2000 m). It is absent from the
western Mediterranean.

Merodon mariae sp. n.
(figs. 25 a-c, 105)

Type-material. - d Holotype ‘France, Corse V. S. van
der Goot / Asco 620 m [42. 15’ N, 9. 10° E] 11.vii.1967
(ZMAN). — Paratypes: 296, 102: same data as holotype 24,
12 (ZMAN); topotypic, 20.vii.1956, leg. v.d. Goot, 19
(ZMAN); topotypic, 5/6.vii.1961, leg. J. A. W. Lucas, 7d
(JLR); topotypic, 9 /11.vii.1967, leg. Lucas, 3? (JLR); topo-
typic, 1/11.viii.1959, leg. P. M. F. Verhoeff, 18 (ZMAN);
topotypic, 25.vii.1977, leg. P. Goeldlin, 2d (coll. Goeldlin,
Lausanne); topotypic, 27.vii.1977, leg. P. Goeldlin, 54
(coll. Goeldlin, Lausanne); “Corsica, La Foce de Vizzavona,
30.vi/ 31.vii.1893, Col. Yerbury 56 (BMNH); ‘Corte,
Corsica, 2.vii.1893, Col. Yerbury 18, 12 (BMNH); ‘Evian,
Corsica, 19.vii.1899 Col. Yerbury 16 (BMNH); Corsica,
Vizzavona 900 1000 m, 2.x.1906, leg. F. Guglielmi’ 1d
(BMNH); ‘Italia Sardegna Alghero 27/28.iv.1957, leg. C. A.
W. Jeekel’ 16 (ZMAN); ‘Italia Sicilia Misterbianco 215 m,
27.v.1966 leg. H. J. P. Lambeck 16 (LR); “Environs de
Salonique [Thessaloniki, Greece], Region du Mt. Profitis
Ilias 768 m, Dr. A. Berton v.1918’ 12 (MNHN); Greece,
Phokis, Delphi 19/22.v.1984, 700-850 m, leg. W.
Hurkmans’ 2d, 12 (wn).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin straight, slightly sulcate longitudinally,
apex acute, antennal ratio 1.6; vertex angle 25 to 30°,
ocellar angle 40°; pubescence whitish yellow with
dark tuft in ocellar region, very dense above antennae,
causing silky lustre; tl-v ratio 0.35.

Thorax: Dark, katepisternum, anepisternum, ane-
pimeron, lateral dorsum and scutellum slightly lus-
trous; pubescence rather dense, deep yellow, very
even, short; vestigial longitudinal bands present in

‘most specimens. Wings slightly infuscate; halteres,

squamae and antisquamae pale yellow.
Legs: Dark, but tibial bases yellow marked; basitar-

sus 2 brown, not black in many specimens; trochan-
ter 3 smooth, femora 3 slightly swollen, rather
straight, triangular processus with distal margin wavy,
weakly serrate, but apically with two strong pedestals,
holding 6-9 bristles; tibiae 3 with short rounded api-
cal spur on inner face, basitarsi 3 somewhat swollen.

Abdomen: Mainly yellow; moderately slender, T II
conspicuously tapering; T II-IV bearing slightly arcu-
ate, narrowly interrupted to continuous pale yellow
pruinose bands and showing dark brown anterior
strips and posterior lunules; pubescence erect to
oblique, rather dense and long, pale orange to golden
throughout; S IV strongly vaulted, posterior margin
entire.

Genitalia: Anterior surstyle lobe apically bearing
short, even, yellow pubescence; subapical-medial ac-
cessory lobe seen as acute angle in lateral view; coales-
cent with posterior surstyle lobe; this lobe very elon-
gate, low, posteriorly showing long, erect, rather
dense, yellow pubescence and bearing a subapical,
medial accessory lobe as well. Cercus large, elongate,
with membraneous ventral plates basally, with dense,
long, erect, yellow pubescence apically; Aedeagus
rather stout, the outer face with elongate subapical
cavity showing paired elongate projections; apical
shaft part slightly lengthened; fringed plates near the-
cal apex suberect.

Body length 15 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Habitus: slightly stouter
than males. Head: Midstripe on frons occupying 1/3
of width, noticeably tapering posteriorly. Thorax:
Longitudinal bands slightly clearer; less lustrous on
sides. Abdomen: T II hardly tapering; pruinose bands
wider, less arcuate, strongly marked; apart from the
quite different colour, not unlike those in femoratus.

Body length 15-16 mm.

Diagnosis. — The males of mariae can be distin-
guished from those of other species by their possess-
ing accessory lobes on both anterior and posterior
surstyle lobe and robust habitus with typical dark yel-
low to orange abdominal pubescence. The females are
stout and show a very sharply defined pattern of prui-
nose bands on the abdomen; they are unlikely to be
confused with other species.

Period of flight and distribution (fig. 105). — M.
mariae is apparently common on Corsica and is
known also from Sardinia, Sicily and Greece. This
distribution pattern is not known from other
Merodon. The flight period is from May through

October, probably in more than one generation.

Etymology. — It is a pleasure to dedicate this species

HURKMANs: Monograph of Merodon

to my wife, Marijke Wester, to commemorate her
support and stimulation of my work. A noun in gen-
itive case.

Merodon ottomanus sp. n.
(figs. 26 a-c, 106)
Type-material. - d Holotype ‘Turkey, Hakkari, Tanin-

Tanin pass 2200 m, 12.vi.1984 leg. J. A. W. Lucas’ (ZMAN).
Paratypes: 7d, 39, all with same data as holotype (JLR).

Description

Male. — Head: Antennae orange, antenna 3 with
upper margin straight, apex subacute, antennal ratio
1.7: vertex angle 50°, ocellar angle 65°; pubescence
long, pale ochre yellow, moderately dense, with dark
tuft in ocellar region; pubescence on compound eyes
pale yellow to white, moderately dense, conspicuous-
ly long; tl-v ratio 0.5.

Thorax: Dark, faintly blue metallic lustrous to
naked eye, under magnification showing very strong
lustre partially dulled by disperse whitish pruinosity,
the lustre strongest on katepisternum, anepisternum
and anepimeron; pubescence as on face but longer
still. Wings slightly yellow tinged, halteres, squamae
and antisquamae yellow to pale yellow.

Legs: All femora dark with yellow apices, tibiae yel-
low with wide dark distal band, tarsi yellow; trochant-
er 3 smooth, femora 3 slender, triangular processus
low and bearing 4-6 bristles on serrate distal margin.

Abdomen: Slender; dark; bluish metallic lustrous
laterally on T II, there showing dense pubescence,
otherwise pubescence moderately dense, pale yellow,
and conspicuously long; T III with vestigial narrow
arcuate pruinose band in some specimens; S IV
strongly vaulted, posteriorly hardly emarginate, par-
tially hidden by curved sides of T IV and its pubes-
cence.

Genitalia: Anterior surstyle lobe strongly yellow,
short even pubescent, somewhat angular, coalescent
with the short posterior surstyle lobe bearing long pu-
bescence posteriorly. Cercus quadrate, with long
dense yellow pubescence. Aedeagus with apical shaft
part lengthened, fringed plates on thecal apex recum-
bent, aedeagus subapically constricted in lateral view.

Body length 9 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: ocellar angle 70°,
frons lustrous, diffusely pruinose laterally. Thorax:
Less lustrous. Abdomen: Pubescence slightly less
dense and long, but still most conspicuous.

Body length 9 mm.

Diagnosis. — Apart from differences in the male
genitalia, ottomanus can be distinguished from other

161

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

species by the long, pale erect pubescence of all ex-
posed parts of the body; viewed with the naked eye
this pubescence produces a hazy fringe around the in-
sect.

Period of flight and distribution (fig. 106). — The
species occurs in south-eastern Turkey where it flies
in June.

Etymology. — The name ottomanus (an adjective)
refers to the former Ottoman empire and was chosen
to indicate the region of occurrence of the species, viz.

Turkey.

Merodon testaceoides sp. n.
(figs. 27 a-c, 105)

Type-material. — d Holotype ‘Yugoslavia, 30 km ZW
Gradsko, 41. 26 N, 21. 50’ E, 13.viii.1963 exc. Leiden’
(RMNH). — Paratypes: 16 (ZMAN), 19 (RMNH), 22 (LR),
same data as holotype.

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin slightly convex, apex obtuse, antennal ra-
tio 1.5; vertex angle 25°; ocellar angle 40°; pubescen-
ce pale yellow; tl-v ratio 0.5.

Thorax: Dark, rather lustrous on katepisternum,
anepisternum, anepimeron, lateral scutellum and
dorsum; pubescence moderately dense, pale yellow;
dorsum showing inconspicuous pruinose bands.
Wings clear; halteres, squamae and antisquamae pale
yellow.

Legs: All femora dark with yellow apices, tibiae yel-
low with broad dark distal band, tarsi yellow to red;
trochanter 3 with low ridge; femora 3 swollen,
curved, bearing normally shaped triangular processus
with 8 bristles on slightly serrate distal margin, widest
spaced basally.

Abdomen: Slender; conspicuously orange, T II ta-
pering considerably, with dark anterior lunule in
some specimens; T II-IV with arcuate pruinose
bands, interrupted on T II-III; pubescence pale
throughout, some dark pubescence often present on
dark anterior lunule of T II; S IV orange, strongly
vaulted, incised up to 2/3 its length from posterior
margin.

Genitalia: Anterior surstyle lobe low, rounded-el-
ongate, with short, dense, even, yellow pubescence
throughout, separate from posterior surstyle lobe by
moderately deep and wide sulcus; posterior surstyle
lobe rounded, rather low, bearing moderately dense,
erect, yellow pubescence throughout, slightly denser
apically. Cercus rounded, rather small, bearing dense
yellow pubescence. Aedeagus showing paired humps
basally on outer face,and marked subapical cavity; ap-

162

ical shaft part slightly lengthened, fringed plates on
thecal apex recumbent to somewhat suberect.
Body length 10-12 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Lustrous midstripe
occupying 1/3 of frons, ocellar angle 50°. Abdomen:
pale reddish to yellow throughout, T II less tapering.

Body length 10-12 mm.

Diagnosis. — The relatively small size and striking
orange abdomen in both sexes precludes confusion
with all but a few species. M. testaceoides is similar ex-
ternally to testaceus from which it is distinguished in
the males by the very different genitalia; the females
of testaceoides can best be distinguished by the paler
colour of the abdomen and longer, denser abdominal
pubescence. Both sexes differ from the species of the
longicornis group by the all red to orange abdomen,
which in that species group is partially crimson, par-
tially black; moreover, the antennal ratio is less in tes-
taceoides.

Period of flight and distribution (fig. 105). — This
species is known only from Macedonia, where it was
collected in August.

Etymology. — The epitheton testaceoides refers to
Merodon testaceus to which this species is externally
similar; the suffix -ides means ‘in likeness of. An ad-
jective.

The alexeji group

Apomorphies: Anterior surstyle lobe elongate; pos-
terior surstyle lobe high to very high, curving medial-
ly.

Description. — Head: Antennae dark, 3rd article
red or orange in some species, antennal ratio 1.5 to
2.4; eyes with touchline incomplete to equalling
0.15-0.9 vertex height; pubescence golden to pale yel-
low with more or less conspicuous dark tuft in ocellar
region. vertex angle and ocellar angle variable, oral
margin well defined.

Thorax: Dark with some lustre and in some species
with traces of longitudinal pruinose bands; wings
mostly clear, veins brown, paler in rufitarsis and soph-
ron; halteres, squamae and antisquamae yellow.

Legs: All dark or with yellow markings on tibiae
and tarsi; trochanters 3 smooth, with faint (Aypochry-
sos) or strong (marginicornis, rufitarsis) edge or knob;
femora 3 mostly rather swollen, straight; basitarsi 3
somewhat swollen in marginicornis.

Abdomen: Rather slender; T II moderately taper-
ing, with slightly to more strongly arcuate pruinose
bands of variable clarity; lateral spots yellow to

orange, or bluish, or lacking; S IV normal.

Genitalia: The anterior surstyle lobe elongate to ex-
tremely elongate, bearing short, dense pubescence at
least apically, separated from the posterior surstyle
lobe by a shallow sulcus, or coalescent; the posterior
surstyle lobes much extended medially, often overlap-
ping, hiding part of the aedeagus from view.
Aedeagus smooth on outer face, mostly slender, api-
cal shaft part slightly lengthened, subapical cavity well
defined.

Merodon alexeji Paramonov
(figs. 28 a, b, 29 a, 81)

Merodon alexeji Paramonov, 1925: 155. Holotype d (and
? paratype) ‘Kechanowka, Balta, Odessa, Ukraine, U. S.
S. R., 1.vi.1924, leg. A. Paramonov’ [not examined, pre-
sumably lost].

Merodon alexeji, Paramonov 1935: 164, 166; Liepa 1969:
20; Van der Goot 1981: 215, 218; Violovitsh 1983: 124;
Peck 1988: 167.

Lampetia alexeji, Sack 1931: 304.

Material examined. — Algeria: 28 (ZMAN, WH); France:
56, 42 (MNHN, KBIN, ZMAN); Greece: 58, 42 (BMNH,
KBIN, ZMUC, NHMW); Italy: 14 (NHMw); Mongolia: 19
(TMA); Morocco: 66, 19 (MNHN); Spain: 2d (ZMAN, WH);
Tunisia: 28 (LR); U.S. S. R. : 54, 39 [Id identified by
original author, in KBIN] (BMNH, KBIN, ZMHB, H. Hippa,
Helsinki).

Description

Male. — Head: Antenna dark brown, antenna 3 ta-
pering, upper margin rather straight, apex subacute,
antennal ratio 2.1; pubescence whitish to pale yellow,
rather sparse with dark tuft in ocellar region; vertex
angle 40°, ocellar angle 50°, ocelli rather large; tl-v ra-
tio 0.45; pubescence on eyes pale, inconspicuous.

Thorax: Dark; katepisternum, anepisternum, ane-
pimeron, dorsum and scutellum metallic lustrous,
with short moderately dense, erect, uneven yellow
pubescence, with variably clear interalar band; in
some specimens vague pruinose bands present. Wings
clear, brown veined; halteres, squamae and antisqua-
mae yellow.

Legs: All dark; trochanters 3 smooth, femora 3
rather swollen, with normal triangular processus bear-
ing 6-12 bristles, basitarsi 3 and other tarsi 3 normal,
tibiae 3 rather slender.

Abdomen: Rather slender, with tapering T II,
somewhat triangular, dark with slight metallic lustre
overall (strongest laterally), bearing dark orange later-
al spots on T II; T I-IV with rather narrow, arcuate,
interrupted whitish pruinose bands, pubescence pale,
inconspicuous medially.

Genitalia: The anterior surstyle lobe much elon-
gate, slightly spathulate, with short, even, dense,
erect, yellow pubescence apically and along dorsal

Hurkmans: Monograph of Merodon

margin; separated from posterior surstyle lobe by a
sulcus; posterior surstyle lobe divided into basal and
apical parts by a suture, anteriorly with smooth knob;
apical part mediolaterally aligned, rather high, these
parts on both sides overlapping; posterior surstyle
lobe with rather dense, yellowish, erect pubescence
but sparser on apical margin. Cercus conspicuous,
rectangular, with dense, long, uneven, yellow pubes-
cence. Aedeagus rather slender, smooth on outer face,
apical shaft part slightly lengthened, fringed plates on
thecal apex oblique to recumbent.

Body length 9-11 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Ocellar region with
darker pubescence; ocellar angle = 60°; lustrous mid-
stripe on frons occupies all but fringes. Thorax:
Pruinose bands clearer, less lustrous. Legs: The femo-
ra 3 rather variable in the degree of swelling.
Abdomen: Triangular, pruinose bands wider, lateral
spots variable, not always clear. T V acute, almost
bald, mostly inconspicuous.

Body length 9-11 mm.

Variation. — Especially in the males M. alexeji is
variable in the clearness of the abdominal lateral
spots, the number of bristles on the triangular proces-
sus, and slightly so in the elongation of the knob on
the posterior surstyle lobe and the position (not the
shape) of the cercus.

Diagnosis. — M. alexeji can be separated from other
species in the alexeji group by the presence of both the
knob on the posterior surstyle lobe and the rectangu-
lar well protruding cercus. The latter feature separates
from altinosus (cercus inconspicuous) and sophron
(cercus short, very wide) which also have the posteri-
or surstyle lobe knob. alexeji is separated from altino-
sus by the lack of dense abdominal pubescence as well.

Period of flight and distribution (fig. 81). — The
imagines occur from May in the Mediterranean to
July in the northern parts of the range, which in-
cludes the mediterranean countries and the temperate
parts of western and central Eurasia.

Discussion. — Although type-material of this spe-
cies was not available there is no doubt about the
identity of alexeji. The original description mentions
a number of diagnostic characters; moreover material
examined and identified by S. J. Paramonov has been
examined.

163

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Merodon altinosus sp. n.
(fig. 29 b, 102)
Type-material. — d Holotype ‘Turkey, Hakkari,

Suvarihalil pass 1250 m., W. side near Habul Deresi [37°
35° N, 43° 20° E], 13.vi.1984, leg. Lucas (ZMAN).

Description

Male. — Head: Antennae blackish, antenna 3 with
upper margin slightly concave, apex subacute, anten-
nal ratio 2.4, pubescence golden, a dark tuft present
in ocellar region; vertex angle 30°, ocellar angle 45°,
tl-v ratio 0.6.

Thorax: Dark, slightly metallic lustrous on katepis-
ternum, anepisternum, anepimeron, dorsum and scu-
tellum, with moderately dense, longish, uneven,
golden pubescence, without pruinose bands. Wings
slightly troubled evenly, especially basally and on an-
terior margin; halteres, squamae and antisquamae yel-
low.

Legs: All dark; trochanters 3 smooth, femora 3
swollen, rather straight, bearing normal triangular
processus with strong apical spur and 8 extremely
short stout bristles; tibiae 3 and tarsi 3 normal.

Abdomen: Dark, tapering posteriorly; T II with
yellow lateral spots; dense, golden, oblique pubes-
cence present throughout; vestigial arcuate pruinose
bands present on T II-IV, clearest on T III.

Genitalia: The anterior surstyle lobe elongate, ante-
riorly with dense, short, even, yellow pubescence, co-
alescent with posterior surstyle lobe; that lobe medio-
laterally aligned in its apical part, overlapping; an
elongate smooth knob present anteriorly on the pos-
terior surstyle lobe; posterior part with long, rather
dense, erect, uneven, yellow pubescence. Cercus
moderately large, not far protruding, bearing dense,
erect, long, yellow pubescence. Aedeagus rather slen-
der, smooth on outer face, fringed plates on thecal
apex suberect.

Body length 9.5 mm.
Female. — Unknown.

Diagnosis. — M. altinosus resembles alexeji from
which it is separated by the conspicuous golden pu-
bescence on the abdomen, the extremely short bristles
on the triangular processus with its strong apical spur
and the less protruding cercus; it is distinguished
from sophron by the same features, and from other
species in the alexeji group by the presence of a knob
on the posterior surstyle lobe.

Period of flight and distribution (fig. 102). — The
species is known from the holotype only, collected in

eastern Turkey in June.

Etymology. — The name altinosus (an adjective) de-

164

notes the golden habitus of the species, and is derived
from the Turkish altin, meaning gold, and the Latin
suffix -osus, meaning full of.

Merodon hirsutus Sack

(figs. 30 a-c, 82)

Merodon hirsutus Sack, 1913a: 435. Lectotype d (here des-
ignated): ‘Djebel Akra [35° 35’ N, 36° 15° E] vi.85
[1885], N. Syria, Dr. E. Leuthner/ Lampetia hirsuta Sack
det. Sack’ (NHMW) [examined].

Merodon hirsutus, Peck 1988: 171.

Lampetia hirsuta, Sack 1931: 318; Séguy 1941: 14.

Material examined. — Israel: 32 (BMNH, RMNH); Jordan:
29 (RMNH); Syria: 9d Paralectotypes, with same data as lec-
totype (NHMW, ZMAN) [examined]; 1% Paralectotype,
‘Syria/ hirsutus Sack/ coll. J. Villeneuve: Lampetia hirsuta
Sack RMHN Belg. 15.392’ (KBIN); Turkey: 22 (BMNH).

Description

Male. — Head: Antennae blackish, antenna 3 with
straight upper margin, apex acute, antennal ratio 1.9;
pubescence yellow, dark tuft in ocellar region weak;
vertex angle 40°, ocellar angle 50°, tl-v ratio 0.9; pu-
bescence on eyes inconspicuous.

Thorax: Dark, slightly metallic lustrous on katepis-
ternum, anepisternum, anepimeron, dorsum and scu-
tellum; pubescence dense, erect, uneven, yellow,
moderately long. Wings clear, brown veined; halteres,
squamae and antisquamae yellow.

Legs: Dark with yellow markings on both ends of
tibiae, rather slender; bearing normal triangular pro-
cessus with short stout bristles, tibiae 3 and tarsi 3
normally shaped.

Abdomen: Dark, rather slender to somewhat stout,
lacking strong lustre (not due to surface sculpturing);
laterally slight micaceous lustre present; T II-IV with
slightly arcuate, moderately wide whitish interrupted
pruinose bands; pubescence rather long, yellowish,
densest laterally, dark and sparser on posteromedian
parts of T III-IV; sternites dark, with scattered whit-
ish pubescence; S IV slightly vaulted.

Genitalia: Anterior surstyle lobe extremely elon-
gate, anteriorly rather rectangular, bearing dense,
short, yellow pubescence, slightly spathulate, separat-
ed from posterior surstyle lobe by shallow but clear
sulcus; posterior surstyle lobe rounded, showing un-
even, long, dense, yellow pubescence especially poste-
riorly. Cercus conspicuously protruding, with dense,
long, yellow pubescence. Aedeagus smooth on outer
face, stout basally, slender apically, apical shaft part
slightly lengthened, fringed plates on thecal apex sub-
erect.

Body length 10 mm.

Female. — Except for sexual dimorphism, differing

from the male as follows. Head: Ocellar angle 55°,
shining midstripe on frons occupies 0.6 of width.
Thorax: Pubescence comparatively shorter, more
even. Legs: femoral apices very narrowly brownish.
Abdomen: Slightly more slender; pruinose band on T
II reduced; those on T III-IV clearer, wider, nearly
continuous, more arcuate.

Body length 9.5-10.5 mm.

Variation. — There is little variation in the males
(all specimens examined are from same locality); the
females show slight size variation.

Diagnosis. — Merodon hirsutus is externally similar
to M. hirtus Sack from which it is easily distinguished
by its very different genitalia; study of the genitalia is
essential since both species fly together in the same re-
gion. From the other species of the alexeji group, hir-
sutus can be separated by the extremely long anterior
surstyle lobe, the relatively low posterior surstyle lobe
and lack of yellow lateral spots on T II.

Period of flight and distribution (fig. 82). — M. hir-
sutus occurs in the eastern Mediterranean region from

March through June.

Merodon hypochrysos sp. n.
(figs. 31 a-c, 82)

Type-material. . — 4 Holotype ‘Turkey, Adiyaman, 10
kms. North of Celikhan, Alu Haral Golii [38° 05’ N, 38°
22° E], 1450 m, 1.vii.1986, leg. Lucas’ (ZMAN). Paratypes:
206 ‘Turkey, Hakkari, Sat Daglari, Varegòs [37° 25’ N,
43° 55° E] 1600-1650 m, 15/18.vi.1984, leg. Lucas’ (JLR).

Description

Male. — Head: antennae dark, antenna 3 red, upper
margin convex, apex rounded, antennal ratio 1.5, ver-
tex angle 40°, ocellar angle 45°, tl-v ratio 0.2; pubes-
cence pale yellow, with dark tuft in ocellar region.

Thorax: Black, lustrous on katepisternum, anepis-
ternum, anepimeron, dorsum and scutellum; pubes-
cence rather sparse, golden, paler down sides and on
scutellum, rather even. Wings slightly tinged, veins
brown, halteres, squamae and antisquamae pale yel-
low.

Legs: Dark, apices of femora and bases of tibiae
paler; trochanters 3 with inconspicuous edge; femora
3 much swollen, straight, bearing normal triangular
processus with weakly serrate distal margin holding 5-
8 small bristles; tibiae 3 long, slender, bearing small
apical spur distally on medial face; tarsi 3 normal, rel-
atively small.

Abdomen: Dark, golden metallic lustrous with
bluish to bronze reflections on sides; T III-IV with
moderately arcuate, whitish to pale yellow, interrupt-

Hurkmans: Monograph of Merodon

ed, moderately wide pruinose bands; pubescence pale
golden, densest laterally.

Genitalia: Anterior surstyle lobe elongate, slightly
diamond-shaped, apically with short, even, dense,
yellow pubescence, posterior surstyle lobe more or
less coalescent but steeply rising, rather rounded,
slightly mediolaterally aligned apically; posterior por-
tion showing dense, erect, yellow pubescence. Cercus
well protruding, moderately large, rounded, bearing
dense yellow pubescence. Aedeagus basally stout, ta-
pering, subapical cavity elongate and showing con-
spicuous paired chitinous plates; an accessory triangu-
lar cavity present just below the recumbent fringed
plates on the thecal apex (which acutely protrudes
well beyond the plates).

Body length 7 mm.

Female. — Unknown.

Diagnosis. — M. hypochrysos can be distinguished
from other species in the alexeji group by its golden
surface and pubescence of the abdomen; the lack of a
knob on the posterior surstyle lobe distinguishes from
the larger but superficially similar altinosus.

Period of flight and distribution (fig. 82). — This
species is known from the mountains of eastern
Turkey where it occurs in June and July.

Etymology. — The epitheton is derived from the
Greek words hypo, i.e. under, less than, and chrysos,
meaning gold, on account of the moderate golden
lustre of the male. An adjective.

Merodon kawamurae Matsumura

(figs. 32 a-e, 79)
Merodon kawamurae Matsumura, 1916: 257. Described af-

ter an unknown number of specimens from Kumamoto,
Kyushu, Japan, leg. Kawamura. [not examined].

Lampetia micromegas Hervé Bazin, 1929: 111. Lectotype d
(here designated): ‘Tchen-Kiang, 13.iv.1918/ Lampetia
micromegas H. B. type’ [examined] (MNHN). Syn. n.

Merodon kawamurae, Paramonov 1925: 160; Shiraki 1930:
206.

Lampetia kawamurae, Hervé Bazin 1929: 111; Sack 1931:
319.

Merodon kawamurai; Shiraki 1968: 198; Peck 1988: 171.

Merodon micromegas, Peck 1988: 172.

Material examined. — China: Paralectotypes of micromeg-
as: 198, 399: Topotypic 9d, 182 (MNHN); ‘Shia-Shu’
62 (MNHN, BMNH); ‘Che-Mo’ 96, 102 (MNHN, BMNH);
‘Nanking’ 39 (MNHN, BMNH); ‘Hoa-Chan 1d, 29
(MNHN), Collection dates from 1.iv.1918 to 16.v.1918. —
Non types: China: 19 (NHMW); Japan: 2d, 19 (BMNH).

165

TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Description

Male. — Head: Antennae orange, antenna 3 with
upper margin convex, apex acute to rounded, most
variable, antennal ratio 1.6; vertex angle 35°, ocellar
angle 50°, tl-v ratio 0.15 to 0.3; vertex sides straight
to slightly concave; pubescence pale yellow, with dark
tuft in ocellar region.

Thorax: Metallic lustrous on katepisternum, ane-
pisternum, anepimeron, dorsum and scutellum, bear-
ing moderately dense and long, uneven, yellow to
white pubescence. Wings slightly tinged, halteres,
squamae and antisquamae yellow; very faint longitu-
dinal striation present in some specimens.

Legs: All femora dark brown, tibiae yellowish with
broad dark band in distal half (in some specimens
more extended), tarsi, especially tarsi 3 dark above,
with golden pubescence underneath; trochanters 3
smooth to distinctly edged (seen as little spine in end-
on view), femora 3 rather swollen, with normal trian-
gular processus bearing 12-15 spinules of which the
apical 1-3 are robust, the others rather fine; tibiae 3
and tarsi 3 normal.

Abdomen: Dark; T II-IV bearing widely to hardly
interrupted, slightly arcuate, white, laterally widening
pruinose bands; T II with rather large orange lateral
spots, dark-centred in some specimens; pubescence
inconspicuous, pale yellow to white, densest laterally
on T II-IV and abdominal tip; S IV weakly vaulted,
posterior margin entire to shallowly emarginate.

Genitalia: The anterior surstyle lobe rather large,
slightly spathulate, elongate, rounded, apically with
dense, short, even, yellow pubescence; posterior sur-
style lobe coalescent, strongly mediolaterally aligned,
overlapping, posteriorly with dense, erect, yellow pu-
bescence. Cercus wide and short, bearing dense, erect,
long, yellow pubescence. Aedeagus smooth on outer
face, apical shaft part short, fringed plates on thecal
apex large, suberect to erect.

Body length 7-11.5 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Antenna 3 generally
less acute than in males; ocellar angle 60°; frons
strongly golden lustrous throughout, diffusely prui-
nose on margins, showing white pubescence; vertex
slightly but clearly edged anteriorly in some speci-
mens. Thorax: Clearly more lustrous; pubescence less
dense; traces of striation clearer. Legs: Often relative-
ly more yellow, especially on tibiae. Abdomen:
Slenderer, lateral spots larger.

Body length 6-12 mm.

Variation. — M. kawamurae is one of the most var-
iable species in Merodon. The variation observed
seems to be size-correlated: the largest specimens
show the strongest lustre, the most acute antenna 3,

166

most edged trochanters 3 and most vaulted and emar-
ginate S IV (males), or strongest edged vertex (fe-
males). In view of this variability, including size vari-
ation of 100%, more than one species would seem to
be involved. There are however several strong argu-
ments for considering all material conspecific: 1.
there are gradual transitions in all variable features; 2.
all males are completely similar in the genitalia; 3.
many cases of such extreme variation are known in
other insect species, some of which have been shown

to be the direct effect of food availability.

Diagnosis. — M. kawamurae can be distinguished
by its clearly spathulate anterior surstyle lobe in the
males and by the golden lustrous vertex, slender ab-
domen and (some specimens) the edge on the vertex.
Moreover it is the only small-sized Merodon species
currently known from China and Japan, the only
larger-sized species being equestris Fabricius. Of other
Merodon species the only one possibly occurring in
this region is alexeji which has a much longer anterior
surstyle lobe and more protruding cercus (fig. 28 a).
M. alexeji has been found in Mongolia but more east-
ward localities are unknown.

Period of flight and distribution (fig. 79). — M.
kawamurae is known from eastern China and Japan,
and occurs in April and May.

Discussion. — The synonymy of M. micromegas
with M. kawamurae might be questioned since no
type-material of M. kawamurae has been examined.
After Horn and Kahle (1936) the Matsumura types
might be at the Agricultural Department of
Hokkaido University, Sapporo, Japan, but inquiries
there have not confirmed the presence of the kawam-
urae types at this institute. The illustrations and de-
scriptions of kawamurae and micromegas leave hardly
any doubt as to their conspecificity; moreover,
Hervé-Bazin himself already stated that micromegas
might be conspecific with kawamurae and described
micromegas since he ‘thought the description of kaw-
amurae as given by Matsumura (1916) insufficient’
(Hervé-Bazin 1929). The Japanese material currently
known is in the BMNH.

Merodon marginicornis sp. n.
(figs. 33 a-c, 82)

Type-material. — d Holotype ‘Iran, Chiraz [26° 40° N,
52° 30’ E], 11.iv.1937, Coll. F. H. Brandt (RMNH).

Description
Male. — Head: antennae large, antenna 3 orange,
apex acute, upper margin edged, concave, antennal

ratio 2.4; vertex angle 35°, ocellar angle 65°, tl-v ratio
0.4, pubescence pale yellow.

Thorax: Dark; katepisternum, anepisternum, ane-
pimeron, dorsum and scutellum bronze lustrous; pu-
bescence rather sparse, short, uneven, pale yellow; no
traces of pruinose bands present. Wings slightly
tinged, brown veined; halteres, squamae and anti-
squamae pale yellow.

Legs: Dark, but tarsi brownish orange; trochanters
3 with distinct knob; femora 3 swollen, rather
straight, with low, strongly serrate triangular proces-
sus bearing 8 spinules; tibiae normal, basitarsi 3
slightly swollen, other tarsi 3 normal.

Abdomen: Dark, vaguely bluish lustrous, with
rather vaguely bordered but well marked, slightly ar-
cuate, interrupted whitish pruinose bands on T II-IV;
T II bearing strongly lustrous bluish lateral spots; pu-
bescence rather sparse, whitish, densest antero-lateral-
ly; S IV deeply emarginate posteriorly, somewhat
vaulted.

Genitalia: Anterior surstyle lobe elongate, bearing
short, even, yellow, erect pubescence dorsally and
along margins, showing elongate bald area, bordered
by a rim, on lateral face; the posterior surstyle lobe
separated by a shallow sulcus, elongate, posteriorly
with yellow, dense, erect pubescence. Cercus incon-
spicuous, with long, dense, yellow pubescence.
Aedeagus smooth on outer face, with long subapical
cavity, showing paired elongate, heavily sclerotized
structures basally; fringed plates suberect, apical shaft
part short.

Body length 9 mm.

Female. — Unknown.

Diagnosis. — M. marginicornis can be easily distin-
guished by the bald, rimmed area on the lateral face of
the anterior surstyle lobe.

Period of flight and distribution (fig. 82). — M.
marginicornis is known only from the holotype, col-
lected in south-western Iran in April.

Discussion. — M. marginicornis has been assigned
to the alexeji group in view of the structure of its male
genitalia; it has somewhat swollen basitarsi 3 [a syn-
apomorphy for the tarsatus group], but lacks the
modification of S IV [the other synapomorphy for
the tarsatus group]. Therefore this species may in fact
occupy a position in between these two groups.

Etymology. — The epitheton marginicornis refers to
the edged upper margin of the antenna 3; it is derived
from the latin margo, meaning edge, and cornu,
meaning horn, in insects often used in referring to an-
tennae. A noun in apposition.

Hurkmans: Monograph of Merodon

Merodon rufitarsis Sack stat. n.

(figs. 34 a-c, 83)

Merodon fulcratus Sack, 1913, subsp. rufitarsis Sack, 1913a:
438. Holotype d: ‘Alai mont [the Alajsski Mountains,
Tadzhikistan, approx. 39° 30’ N, 71° to 73° E] (ZMHB)
[examined].

Merodon (Exmerodon) fulcratus ssp. rufitarsis Peck 1988:
165.

Description

Male. — Head: Antennae dark, antenna 3 red, apex
rounded, upper margin convex, antennal ratio 1.6;
vertex angle 30°, ocellar angle 50°, pubescence pale
yellow; touchline incomplete, tl-v ratio 0.15; comp-
ound eyes with inconspicuous white pubescence.

Thorax: Dark; katepisternum, anepisternum, ane-
pimeron, dorsum and scutellum slightly lustrous,
bearing moderately long and dense, yellow, rather un-
even pubescence, without traces of pruinose bands.
Wings clear, veins pale brown, halteres, squamae and
antisquamae pale yellow.

Legs: All femora dark, tibiae orange with wide dark
distal band, tarsi orange; trochanters 3 with low sharp
edge, femora 3 moderately swollen, rather straight,
bearing normal triangular processus with 10 spinules;
tibiae 3 and tarsi 3 normal.

Abdomen: Dark, without lateral spots or pruinose
bands; pubescence rather dense, uneven, yellow, sub-
erect; S IV strongly keeled; abdominal surface slight-
ly lustrous throughout, strongest laterally.

Genitalia: Anterior surstyle lobe much elongate,
with short yellow even pubescence apically and down
margins; posterior surstyle lobe separated by shallow
sulcus, large, strongly mediolaterally aligned, showing
rather long, yellow pubescence apically and posterior-
ly. Cercus rather short, wide, with dense, long, yellow
pubescence. Aedeagus smooth on outer face, with
paired chitinous plates in middle region of shaft; api-
cal shaft part rather short, subapical cavity well
marked, fringed plates on thecal apex suberect.

Body length 9 mm.

Female. — Unknown.

Diagnosis. — M. rufitarsis can be distinguished by
the combination of lacking abdominal bands and lat-
eral spots with the mediolaterally aligned posterior
surstyle lobe.

Period of flight and distribution (fig. 83). — M. ru-
fitarsis is known from Tadzhihistan; the flight period

is unknown.
Discussion. — M. rufitarsis cannot be considered a

subspecies of M. fulcratus Becker as originally estab-
lished by Sack (1913a). From their respective descrip-

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TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

tions it is clear that Becker and Sack described two
different species, although both were based on infor-
mation provided by Loew. Type material belonging
to the nominate form of M. fulcratus Sack has not
been recovered. Merodon fulcratus Sack, 1913 (pub-
lished August, 1913) is a junior homonym of
Merodon fulcratus Becker, 1913 (published April,
1913), and should be renamed when its identity has
become clear. The male genitalia are very different.
For instance, in M. fulcratus Becker the aedeagus is
abruptly tapering basally and the surstyle lobes are
smaller. M. rufitarsis has completely touching eyes in
the male, whereas in males of fulcratus Becker the eyes
are widely separated. These species are assigned to dif-
ferent species groups, and are not considered closely
related.

Merodon sophron sp. n.
(figs. 35 a-c, 81)

Type-material. — 4 Holotype ‘Morocco, Azrou [33° 25°
N, 05° 20’ W], 29.v.1925, E. Hartert’ (BMNH).

Description

Male. — Head: Antennae dark, antenna 3 with
slightly convex upper margin, apex acute, antennal
ratio 2.0; vertex angle 30°, ocellar angle 60°, tl-v ratio
0.6; pubescence pale yellow, with dark tuft in ocellar
region.

Thorax: Dark, katepisternum, anepisternum and
anepimeron lustrous; pubescence greyish yellow,
slightly uneven; no pruinose bands present. Wings
clear, orange veined; halteres, squamae and antisqua-
mae pale orange.

Legs: All dark; trochanters 3 smooth; femora 3
swollen, with normal triangular processus bearing 6
bristles; tibiae 3 and tarsi 3 normal.

Abdomen: Dark, comparatively stout; T II-IV with
slightly arcuate, narrow, widely interrupted whitish
pruinose bands and slight metallic lustre overall; T II
with clear metallic lustrous lateral spots bearing rath-
er dense, golden pubescence; pubescence otherwise
paler, less conspicuous.

Genitalia: Anterior surstyle lobe elongate, rectan-
gular, markedly curving dorso-anteriorly, spathulate,
with dense, yellow, even pubescence apically; posteri-
or surstyle lobe separated by small clear sulcus, rather
large, very high, apical parts mediolaterally aligned
and largely overlapping; the posterior surstyle lobe
bears a bald, slightly elongate knob anteriorly; from
here a suture runs toward the posterior margin, divid-
ing the posterior surstyle lobe in basal and apical
parts; the posterior margin bears dense, long, erect,
yellow pubescence. Cercus very large with same pu-
bescence. Aedeagus smooth on outer face, excavated
on this face in hardly lengthened apical shaft part;

168

fringed plates on thecal apex rather large, erect; in
natural position aedeagus entirely covered by posteri-
or surstyle lobe.

Body length 9.5 mm.

Female. — Unknown.

Diagnosis. — M. sophron is distinguished from oth-
er species by its very large overlapping posterior sur-
style lobe and its golden pubescence on the metallic
lustrous lateral spots on T II.

Period of flight and distribution (fig. 81). — M.
sophron is known only from the holotype, collected in
Morocco in May.

Etymology. — The epitheton sophron is derived
from the greek words soos, meaning chaste, and
phronein, meaning to act, and refers to the ‘chaste’
covering of the aedeagus in this species. A noun in ap-
position.

Merodon tener Sack
(fig. 81)

Merodon tener Sack, 1913a: 443. Lectotype 2 (here desig-
nated): ‘Sarepta [= Krasnoarmeysk near Volgograd, after
Peck 1988]/ M. tener Sack det. Sack/ coll. Lichtwardt/
coll. D. E. I. Eberswalde’ [examined].

Merodon tener, Becker 1921: 54; Van der Goot 1981: 215.

Lampetia tenera; Sack 1931: 329.

Material examined.-® Paralectotype with same data as
lectotype (DEIC).

Description

The lectotype and paralectotype are quite similar to
alexeji females. The main differences are: ocellar ang-
le 70° in the tener types, 60° in alexeji, coloration of
abdomen reddish throughout in tener and slightly
stouter than in alexeji. Lectotype and paralectotype
differ slightly in the degree of swelling of the femora
3 but are otherwise similar among themselves and to
alexeji.

Period of flight and distribution (fig. 81). — M. ten-
er is known from the southern and eastern parts of
Russia, where it has been collected in June (Sack

19315529);

Discussion. — Since no male type-material of tener
has been traced it is impossible to ascertain the status
of tener as a separate species. In view of the differences
stated tener is here considered to be a separate species.
In view of the similarity in the females the species is
tentatively assigned to the alexeji group.

Merodon trizonus (Szilady)

Lampetia trizona Szilady, 1940: 67. Syntypes d and 9: La
Calle [el Kala], Algeria and Ain Draham, Tunisia [not ex-
amined].

Discussion. — The extremely brief description pro-
vided by Szilady is insufficient to separate trizonus
from other Merodon. The original description ex-
pressly states that the species is related to hirsutus
Sack. Therefore the species is tentatively assigned to
the alexeji group which also includes hirsutus. The
types of this species, formerly stored in TMA, have
been lost in 1956 (Dr. A. Dely-Draskovits, in litt.).

No other material assigned to this species is known.

The tarsatus group

Apomorphies: Basitarsus 3 strongly swollen, spe-
cialized pubescence present on inner face; S IV at least
keeled, mostly strongly modified.

Description. — Head: Antennae orange to brown,
antennal ratio 1.3 to 1.7, compound eyes in male
usually touching with tl-v ratio 0.1 to 0.3, pubescence
yellow to white, oral margin well marked. Thorax:
Dark, metallic lustrous. Wings clear to slightly
tinged, orange to dark veined, halteres, squamae and
antisquamae yellow. Legs: Femora 3 with normally
shaped triangular processus, tibiae 3 in male often
bearing apical floccus inserted on distal spur, basitar-
si 3 swollen strongly to enormously, less conspicuous-
ly so in females; showing short thick-set pubescence
on inner face. Abdomen: Tergites metallic lustrous, S
IV strongly modified in males of all species except au-
ronitens and caudatus where it is only keeled.
Genitalia: Anterior surstyle lobe simple, often with
short, dense pubescence distally, separated by small
suture from higher posterior surstyle lobe bearing
long posterior pubescence. Cercus well protruding.
Aedeagus with apical part short.

Merodon ankylogaster sp. n.
(figs. 36 a-b, 84)

Type-material. - d Holotype ‘Iran, Fars, Road Chiraz-
Kazeroun, Ft. Sine-Sefid [Southwestern Iran, about 29° 35’
N, 52° E, in mountains], 29.iv.1937, coll. F. H. Brandt’
(RMNH). Paratype 9, same data as holotype (RMNH).

Description

Male. — Head: Antennae orange, antenna 3 with
upper margin convex, apex obtuse, antennal ratio 1.3;
pubescence pale yellow, dark hairs present in ocellar
region; vertex angle 40°, ocellar angle 45°, vertex lar-
ge, d-v ratio 0.15.

Thorax: Coppery lustrous on katepisternum, ane-
pisternum, anepimeron, dorsum and scutellum, dor-

Hurkmans: Monograph of Merodon

sal surface rather roughly sculptured, showing yellow,
erect, uneven pubescence. Wings orange veined,
clear.

Legs: Dark, tibiae basally yellow; tarsi with dense
golden pubescence; trochanters 3 smooth, femora 3
much swollen, slightly metallic lustrous, with normal
triangular processus; tibiae 3 without apical floccus;
basitarsi 3 strongly swollen, bearing dense, golden,
short pubescence medially; other tarsi 3 normally
shaped.

Abdomen: T II much tapering posteriorly, without
lateral spots; pruinose bands interrupted, slightly ar-
cuate; pubescence moderately dense, golden, oblique;
S IV with conspicuous median projection; abdominal
surface roughly sculptured on medial region of all ter-
gites.

Genitalia: The anterior surstyle lobe elongate,
slightly spathulate, apically bearing short, dense,
erect, yellow pubescence, posterior surstyle lobe sep-
arated from anterior surstyle lobe by shallow sulcus,
steeply rising, posteriorly with long dense yellow pu-
bescence. Cercus subangular, oblong, with similar
pubescence. Aedeagus stout, with fringed plates on
thecal apex recumbent, apical part screened by the ap-
ical portion of the posterior surstyle lobe in natural
position.

Body length 7.5 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Ocellar angle 60°;
frons lustrous throughout, laterally yellowish metallic
lustrous. Thorax: Stouter than in male; sculpturing
less strong, pubescence more whitish. Abdomen:
Stouter, pruinose bands nearly continuous; pubes-
cence whiter; S IV normal.

Body length 9 mm.

Diagnosis. — The male is unmistakable in view of
the morphology of S IV; the female ankylogaster can
be separated from turkestanicus by the partially orange
legs, and from caudatus by the lack of lateral spots on
the abdomen. :

Period of flight and distribution (fig. 84). — The
species is known only from the types, collected in Iran

in April.

Discussion. — The shape of the anterior surstyle
lobe and inward curve of the posterior surstyle lobe in
ankylogaster suggest a certain degree of relationship
with alexeji and its allies [note the discussion of mar-
ginicornis above]; in view of the modified S IV and
the swollen basitarsi 3, ankylogaster has been assigned
to the tarsatus group.

Etymology. — The epitheton ankylogaster is derived

from the Greek ankylos, meaning spine, and gaster,

169

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

meaning belly, hence abdomen, and refers to the pro-
jection of S IV in the male of this species. The epithe-
ton is to be treated as a noun in apposition.

Merodon auronitens sp. n.
(figs. 37 a-c, 84)

Type-material. — Holotype d ‘Israel, Mt. Carmel, Bat
Slomo 32° 35’ N, 35° 00’ E, 22.iii.1981, W. B. M. Brantjes’

(ZMAN). The antennae and the right leg 3 on cardboard tag,
both anterior surstyle lobes attached to another cardboard

tag.

Description

Male. — Head: Antennae black, 3rd article with up-
per margin concave, apex subacute, antennal ratio
1.7, vertex angle 35°, ocellar angle 40°, tl-v ratio
0.35; pubescence very dense, long, glossy golden on
frons, slightly shorter and sparser on face and vertex.

Thorax: Dark; golden metallic lustrous laterally,
bearing moderately dense, even, golden brown pubes-
cence. Wings clear, halteres, squamae and antisqua-
mae yellow.

Legs: Dark throughout, tibial bases and tarsi slight-
ly lighter; trochanters 3 smooth, femora 3 much swol-
len, showing normally shaped triangular processus
bearing 8 bristles; tibiae 3 show a clear, golden apical
floccus; basitarsi 3 swollen, with very dense, short,
golden pubescence on inner face.

Abdomen: T II slightly tapering, with traces of lat-
eral spots; T II-IV with arcuate greyish pruinose
bands, that on T II inconspicuous; pubescence rather
long, even, yellow, moderately dense; sternite IV
keeled, posteriorly slightly emarginate, but not de-
formed.

Genitalia: Anterior surstyle lobe somewhat elon-
gate, with dense orangish pubescence throughout;
connection to posterior lobe severed; posterior lobe
rather high, curving mediad apically, with scattered
yellow pubescence on apical margin; posterior end
with dense yellow pubescence. Cercus well protrud-
ing, bearing dense, long, yellow pubescence.
Aedeagus smooth on outer face, fringed plates on the-
cal apex recumbent.

Body length 11.5 mm

Female. — Unknown.

Diagnosis. — M. auronitens can be distinguished
from other species in this group by the lack of a de-
formed S IV, like caudatus, but differs from that spe-

cies by not having a deformed genital cap.
Period of flight and distribution (fig. 84). — M. au-

ronitens is known only from the holotype, collected in
Israel in March.

170

Etymology. — The epitheton auronitens is latin, and
derived from the word aurum, meaning, gold, and
the verb nitere, meaning, to shine, the construction
meaning, shining with gold, as a reference to the
glossy pubescence on the head. The epitheton is to be
treated as an adjective.

Merodon caudatus Sack
(figs. 38 a-d, 84)

Merodon caudatus Sack, 1913a: 446. Holotype: Asia Minor
[Turkey] [not examined, considered lost].

Merodon caudatus, Paramonov 1926b: 322; Peck 1988: 168.

Lampetia caudata; Sack 1931: 309.

Material examined. — Israel: 26, 12 ‘Mt. Carmel, Beth
Slomo, 22.3.1981 leg. Brantjes’ (LR); 12 ‘Palestine, Beth
Hakerem [near Jerusalem], 17.3.1951 leg. O. Theodor’
(RMNH).

Description

Male. — Head: Antennae dark, antennal ratio 1.8,
antenna 3 with apex obtuse, somewhat edged; vertex
large, metallic lustrous, vertex angle 50°, ocellar ang-
le 65°, pubescence yellow, whitish on face and occipi-
tal sides; compound eyes showing grey pubescence,
tl-v ratio 0.2-0.25.

Thorax: Dark, brassy lustrous on katepisternum,
anepisternum and anepimeron, dorsum and scutel-
lum, showing dense, erect, yellowish, even pubes-
cence, diffusely pruinose on lateral dorsum, scutellum
and sides. Wings quite clear, halteres, squamae and
antisquamae pale yellow.

Legs: All femora dark, apically paler, tibiae yellow
with dark distal band, tarsi orange to brown, apical
articles darkest; pubescence dense on tibiae and tarsi,
yellow underneath; trochanters 3 smooth, femora 3
rather straight, triangular processus somewhat re-
duced, forming only slight knob; tibiae 3 curved and
twisted, bearing apical floccus on drawn-out distal
medial spur; basitarsi 3 short, swollen, with furrow
giving twisted aspect, bearing short, even, dense, yel-
low pubescence; other tarsi 3 normal.

Abdomen: Tergites dark, metallic lustrous, T II
with orange lateral spots and erect, orange pubescence
(densest and longest on lateral spots); pruinose bands
arcuate, interrupted on T III, continuous on T IV; S
normal but S VIII (the genital cap) strongly modified,
elongate, with apical floccus conspicuous to naked
eye (hence the name caudatus).

Genitalia: The anterior and posterior surstyle lobes
separated by deep wide sulcus; anterior surstyle lobe
rounded-triangular, with short, even, dense, yellow

pubescence, showing ridge running from ventro-pos-

terior angle to mid region; posterior surstyle lobe
short and high, with longish, uneven, yellow pubes-
cence apically. Cercus well protruding, angular, with

very long, yellow pubescence. Aedeagus with short
apical shaft part, the fringed plates on thecal apex re-
cumbent.

Body length 12 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Ocellar angle 70°,
frons metallic lustrous, diffusely pruinose on sides;
frons clearly tapering toward vertex. Thorax:
Markedly stouter than in male. Legs: Slenderer than
in males, basitarsi 3 quite similarly modified but pu-
bescence duller. Abdomen: Stouter, rather triangular,
with conspicuous lateral spots bearing fiery orange
hair-tufts; pruinose band on T III nearly continuous.

Body length 9.5 mm.

Diagnosis. — The males are easily separated by their
unique genital cap; the females are similar to those of
ruficornis Meigen and allied species from which they
are separated by the swollen basitarsi 3.

Period of flight and distribution (fig. 84). — The
species is known from Israel and Turkey where it oc-
curs in March.

Discussion. — In spite of the fact that no type-ma-
terial was available, the identification of this species is
certain in view of the extraordinary features present.
The assignment of the female of caudatus as present-
ed here seems justified in view of the striking similar-
ities (basitarsi 3, lateral spots on T II) and the concur-
rent flight in the same area.

Merodon oidipous sp. n.
(figs. 39 a-d, 84).
Type-material. — dé Holotype ‘Turkey, Hakkari,

Suvarihalil pass 2100 m, 4.vi.1984, leg. J. A. W. Lucas’
(ZMAN). Paratypes: 2d , same data as holotype (JLR).

Description

Male. — Head: Antennae orange to brown, anten-
nae III paler than I-II, upper margin straight to slight-
ly concave, apex rounded, antennal ratio 1.6; vertex
angle = 40°, ocellar angle = 60°, tl-v ratio 0.25; face
and occiput slightly, vertex strongly blue metallic lu-
strous; pubescence whitish to pale yellow, face more-
over diffusely pruinose; compound eyes with greyish
pubescence.

Thorax: Dark, with some steel blue lustre on later-
al dorsum and scutellum, stronger on sides; pubes-
cence erect, moderately dense, rather short, uneven.
Wings clear; halteres with knob yellow, stalk darker,
squamae and antisquamae yellow.

Legs: All femora dark with narrow yellow apices,
tibiae yellow with broad dark distal band, tarsi dark

Hurkmans: Monograph of Merodon

above, with dense pale pubescence underneath, upper
faces orangeish to brown on legs 1-2, darker on 3; tro-
chanter 3 bearing low projection with yellow hair-
tuft; femur 3 swollen, bluish metallic lustrous; trian-
gular processus rather large, with conspicuous apical
spur, otherwise finely serrate, bearing some 9 bristles;
tibiae 3 with long yellow apical floccus inserted on
drawn-out distal inner angle; basitarsi 3 enormously
swollen, almost globular, brownish, with dense gold-
en short erect pubescence medially; other tarsi 3 nor-
mal.

Abdomen: Rather slender, dark; T II tapering pos-
teriorly, T I-IV bearing slightly arcuate moderately
wide interrupted pruinose bands; pubescence whit-
ish, rather sparse, slightly denser laterally and on prui-
nose bands, darker medially; at certain angles vague
metallic blue lustrous lateral spots on T II may be
seen; S IV with paired, conspicuous appendages.

Genitalia: The anterior surstyle lobe somewhat
spathulate, bearing dense, yellow, even, erect pubes-
cence on distal margin; separated from the slightly el-
ongate rounded posterior surstyle lobe by a shallow
sulcus; posterior surstyle lobe incised dorsally, bearing
dense, erect, yellow pubescence. Cercus conspicuous,
elongate, with same pubescence. Aedeagus rather
slender with short apical shaft part, fringed plates on
thecal apex recumbent to suberect.

Body length 8.5 mm.
Female. — Unknown.

Diagnosis. — M. oidipous can be separated by its
enormously swollen basitarsi 3 which are differently
shaped than in other species of the tarsatus group, by
the clear blue lustre on the vertex, and the more pro-
truding cercus.

Period of flight and distribution (fig. 84). — The
species, known only from the types, has been found
in south-eastern Turkey in June.

Etymology. — The epitheton oidipous derives from
the Greek and means, swollen foot. It refers to the
enormously swollen basitarsi 3, conspicuous to the
naked eye.

Merodon persicus sp. n.

(figs. 40 a-d, 85)

Type-material. — d Holotype ‘Iran, Fars, Ft. Sine-Sefid,
Rd. Chiraz Kazeroun [29° 35’ N, 52° E], Coll. F. H. Brandt
25.v.1937 / Lampetia smirnovi det. P. H. van Doesburg’
(RMNH).

Description
Male. — Head: Antennae brown, antenna 3 orange,

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

upper margin straight to slightly concave, apex acute,
antennal ratio 2.3; pubescence pale yellow, with dark
hair-tuft in ocellar region; oral margin more protru-
ding than in the other species in this group; face
slightly diffusely pruinose; vertex angle 40°, ocellar
angle 60°; tl-v ratio 0.25; compound eyes with whi-
tish pubescence.

Thorax: Rather dull, surface dark brown, bearing
uneven, short, erect yellow pubescence on dorsum,
scutellum, katepisternum, anepisternum and anepim-
eron, with weak bronze lustre on these elements.
Wings clear, halteres, squamae and antisquamae yel-
low.

Legs: Dark but bases of tibiae and tarsi brown; tro-
chanter 3 showing well-defined low hump bearing
rather short black bristly pubescence; femur 3 swol-
len, curved, with rather small triangular processus,
strongly serrate on distal margin; tibiae 3 short, stout,
with distinct apical floccus on distal medial angle;
basitarsi 3 strongly swollen, showing dense, golden,
short, erect pubescence throughout (densest on medi-
al face). A slightly elevated, well defined area proxi-
mally on lower face bearing erect, dense dark bristly
short pubescence obliterating the surface of this area;
other tarsi 3 normal.

Abdomen: Moderately slender, all dark, with rath-
er strong olivaceous metallic lustre; vaguely defined,
arcuate pruinose bands present on T III-IV; pubes-
cence golden throughout, some dark hairs present in
posteromedian parts of T II-III. All S rectangular, S
IV bearing conspicuous, paired appendages visible to
naked eye.

Genitalia: The anterior surstyle lobe elongate, api-
cal part widening and projecting dorsally, showing
short, yellow, erect pubescence along ventral and dis-
tal margins; separated from posterior surstyle lobe by
shallow suture, from which the rounded-rectangular
posterior surstyle lobe steeply rises; posterior surstyle
lobe with yellow, scattered, erect pubescence, longest
on ventral and posterior margins. Cercus largish,
bearing dense, long, yellow, erect pubescence.
Aedeagus stout, fringed plates on thecal apex recum-
bent, apical shaft part short; in natural position the
apical portion of the aedeagus is covered by the poste-
rior surstyle lobe tops.

Body length 9.5 mm.

Female. — Unknown.

Diagnosis. — M. persicus seems to be most closely
related to tarsatus, from which it differs in the smaller
size, the entire basitarsi 3 with a small raised area (as
opposed to excavated basitarsi 3 with large area in tar-
satus), the elongate posterior surstyle lobe with dorsal-
ly projecting apical part, and the stronger inward
curve of the apical posterior surstyle lobe region. M.

172

persicus differs from the description of smirnovi in the
smaller size, the smaller tl-v ratio and the colour of
the antennae.

Period of flight and distribution (fig. 85). — The

species is known to occur in Iran; it flies in May.

Etymology. — The epitheton persicus is derived
from the classical name Persia, indicating the region
of occurrence of the species.

Merodon tangerensis sp. n.
(figs. 41 a-d, 87)

Type-material. — d Holotype ‘Environs de Tanger
[Morocco], Fevrier 1859’ (MNHN).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apex rounded, antennal ratio 3.3;
pubescence pale yellow with dark hair-tuft in ocellar
region; vertex angle 30°, ocellar angle 35°, touchline
incomplete, tl-v ratio 0.15; face slightly, diffusely
pruinose, occiput densely pruinose down sides, me-
tallic lustrous; compound eyes showing whitish pu-
bescence.

Thorax: Dark, slightly metallic lustrous on lateral
dorsum and scutellum; katepisternum, anepisternum
and anepimeron steel grey lustrous; dorsum with
double paired anteriorly coalescent whitish pruinose
bands; pubescence erect, yellow, moderately dense,
distinctly uneven throughout, paler on sides. Wings
clear, pale brown veined; halteres, squamae and anti-
squamae yellow.

Legs: Dark but apices of femora and bases and api-
ces of tibiae lighter; trochanter 3 smooth, femur 3
swollen, black metallic lustrous, bearing normal tri-
angular processus holding 6 bristles on serrate distal
margin; tibiae normal, showing drawn-out distal me-
dial angle bearing dense somewhat lengthened pubes-
cence (this might be termed a subfloccus); basitarsi 3
swollen, with dense, yellowish, erect pubescence
(densest medially); other tarsi 3 normal.

Abdomen: Dark, slender, tergites rectangular,
slightly transverse, with white, slightly arcuate mod-
erately wide pruinose bands, interrupted on T II,
continuous on T III-IV; pubescence short, white on
bands, dark elsewhere; S IV strongly keeled, deeply
emarginate posteriorly.

Genitalia: The anterior surstyle lobe elongate, with
apical part bearing short, even, yellow, dense pubes-
cence, more scattered elsewhere; separated from pos-
terior surstyle lobe by deep sulcus; posterior surstyle
lobe high, apically curving medially; with marked an-
tero-ventral angle, posteriorly rounded, there with
moderately dense rather long yellow pubescence.

Cercus inconspicuous, bearing long dense yellow pu-
bescence. Aedeagus slender, fringed plates recum-
bent, subapical cavity well marked.

Body length 8.5 mm.

Female. — Unknown.

Diagnosis. — M. tangerensis is distinguished from
other species in this group by its very long antenna 3,
the relatively small projection of S IV and the relative-
ly large somewhat dorsally curving anterior surstyle

lobe.

Period of flight and distribution (fig. 87). — This
species is known only from the holotype, collected in
northern Morocco in February.

Etymology. — The species is named after the site
where the holotype was collected. The epitheton is to
be used as an adjective.

Merodon tarsatus Sack

(figs. 42 a-e, 83).

Merodon tarsatus Sack, 1913a: 437. Holotype d: ‘Pamir
49409/ Mer. tarsatus Sack det. Sack’ (ZMHB) [examined].

Merodon smirnovi Paramonov, 1926b: 320. Holotype d:
‘Ak-Tash, Prope Tashkent, Turkestan’ [not examined,
see discussion]. Syn. nov.

Merodon tarsatus, Paramonov 1926b: 320; Paramonov
1927: 76, 78; Peck 1988: 174.

Lampetia tarsata, Sack 1931: 328.

Merodon smirnovi, Paramonov 1927: 76, 78; Liepa 1969:
20; Peck 1988: 173.

Lampetia smirnovi, Sack 1931: 326.

Material examined. — U. S. S. R.: 16 ‘Ak-Tash [41° 37°
N, 69° 40° E], 9 vii 1923, Tashkent A.Zecholovtsev/
Lampetia smirnovi Param., S. Paramonov det. mm/ coll. J.
Villeneuve’ (KBIN); 12 ‘Berkana, Turkestan, 24.vi.1925,
Kuznetsov leg.” (ZMAS); 19 ‘Outsj. Koydara, dol. R.
Warzob, Tadzj. [river Warzob valley in Tadzhikistan,
North of Dushanbe in 68° 50’ E, between 38° 20’ and 38°
55’ N] (ZMAS).

Description

Male. — Head: Antennae brown, antenna 3 with
upper margin edged, straight, apex subacute, anten-
nal ratio 1.9; pubescence yellow, with dark hair-tuft
in ocellar region; vertex angle 40°, ocellar angle 70°;
d-v ratio 0.3; face and occiput metallic greenish lus-
trous; central ocellus only half as large as the lateral
ones; compound eyes with whitish pubescence.

Thorax: Strongly lustrous on lateral dorsum and
scutellum, katepisternum, anepisternum and anepim-
eron; dorsum and scutellum with diffuse golden pu-
bescence and scattered pruinosity, with 4 faint prui-
nose bands also showing. Wings tinged brown on
central portions, clear marginally, veins very dark;

Hurkmans: Monograph of Merodon

halteres, squamae and antisquamae yellow.

Legs: All joints and tibial bases yellow, otherwise
dark; trochanter 3 sharply keeled, femur 3 swollen,
metallic lustrous, triangular processus normal, bear-
ing 6-8 bristles on serrate distal margin; tibiae 3 rath-
er strongly curved, bearing large lamella on distal me-
dial face with ruddy floccus inserted apically; floccus
extends over half the length of basitarsi 3; these swol-
len, asymmetrical, with lower face showing large flat
metallic lustrous area bearing stout bristle-like pubes-
cence, bordered by very stout long pubescence; inner
face of basitarsi 3 showing dense orange-yellow pu-
bescence.

Abdomen: Dark, greenish yellowish metallic lus-
trous, rather slender; pubescence dense, moderately
long, pale orange, partially obliterating roughly punc-
tated surface; no pruinose bands; S IV modified,
bearing large, slender paired appendages.

Genitalia: The anterior and posterior surstyle lobes
separated by shallow sulcus, of about equal size; ante-
rior surstyle lobe apically with short, dense, erect, yel-
low pubescence, of rectangular outline; posterior sur-
style lobe moderately high, regularly rounded,
bearing erect, long, yellow, uneven pubescence. Cer-
cus of moderate size, with similar pubescence, slight-
ly angular. Aedeagus robust, heavily sclerotized, api-
cal shaft part short, fringed plates recumbent.

Body length 10.5-13 mm.

Female. — Rather different from the males, as not-
ed by e.g. Sack, 1931; therefore a full description is
given.

Head: Antenna 3 dark orange, subacute; antennal
ratio 2-2.3; pubescence whitish; ocellar angle 70°; the
central ocellus clearly smaller than the lateral ones;
frons diffusely pruinose laterally, lustrous for median
one-third; metallic lustre absent.

Thorax: Weakly greenish metallic lustrous on later-
al dorsum and scutellum, katepisternum, anepister-
num and anepimeron; pubescence rather short, erect,
even, pale yellow. Wings as in male, slightly less
tinged.

Legs: All femora dark, tibiae orange with dark dis-
tal band, femur 3 slightly swollen, triangular proces-
sus bearing 6 bristles on serrate distal margin; tibiae 3
short, otherwise normal; basitarsi 3 clearly swollen,
not deformed, bearing dense golden pubescence;
other tarsi 3 normal.

Abdomen: Dark, slightly metallic lustrous; prui-
nose bands on T II-IV narrow, whitish, arcuate, well
defined, interrupted; pubescence yellow to white,
deepest coloured and densest laterally on T II.

Body length 12 mm.

Diagnosis. — M. tarsatus can be distinguished in the
males, by its deformed basitarsi 3 with the lustrous

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

area at the lower face which at once separates it from
all other species; in the females, by the clearly swollen
basitarsi 3 and the great antennal ratio.

Period of flight and distribution (fig. 83). — M. tar-
satus occurs in Tadzhikistan, and adjacent regions,
where it flies in June and July.

Discussion. — In the original description the colour
of the pubescence of the holotype is said to be foxy-
red, its size as 12 mm; the colour has apparently
faded, while the specimen was measured recently at
10.5 mm only. The synonymy of smirnovi with tarsa-
tus was established after examination of one of the
specimens (originating from the type-locality of smir-
novi) identified as smirnovi by Paramonov, and
matching the description as well as the holotype of
tarsatus. Moreover, the females, all identified previ-
ously as smirnovi, match the description of the © tar-
satus as provided by Sack. Furthermore Sack (1931)
noted the specialized setaceous area on the basitarsus
3 of smirnovi which he failed to observe in tarsatus,
but which is easily seen on the holotype of tarsatus.
Sack gave a difference in the tl-v ratios between tarsa-
tus and smirnovi (the ratio of tarsatus being greater) in
his key to the species (Sack, 1931), but checking of
the holotype of tarsatus showed that the tl-v ratio in
that species is about 0.3, whereas the ratio given by
Paramonov for smirnovi is 0.5. Therefore Sack’ s
interpretation that two species are involved is not fol-
lowed here; in view of the above arguments smirnovi
is considered synonymous with tarsatus.

Merodon turkestanicus Paramonov

(figs. 43 a-e, 86)

Merodon turkestanicus Paramonov, 1926b: 319. Holotype à :
‘Tashkent, Tadzhikistan, 12.v.1915’ [U. S. S. R.] [not ex-
amined].

Merodon turkestanicus, Paramonov 1927: 75; Liepa 1969:
20; Peck 1988: 175.

Lampetia turkestanica; Sack 1931: 330.

Material examined. — Iran: 1d ‘Kuh-e-Hazaran, S. Rayen
Kerman [29° 24’ N, 57° 22’ E], 3800 m, 25.v.1978 leg. K.
Warncke’ (LR); U. S. S. R.: 24 ‘Outsj. Koydara, dol. R.
Warzob Tadzj. [68° 50’ E, 38° 20’ to 38° 50’ N], Warzob
river, Tadzhikistan ] Stackelberg 6.v.43’ (ZMAS); two 9
from same locality and collector, dated 2.v.44 (ZMAS,
BMNH); 19 ‘Outsj. Kwak. Werch. Koydarye Tadzh. [in
same region as holotype] Stackelberg 27.v.43’ (RMNH); 1d
‘Hissar mountains Tadzhikistan [same region], Stackelberg

3.v.44’ (RMNH).

Description
Male. — Head: Antennae dark, antenna 3 with up-
per margin straight, apex obtuse, antennal ratio 2.1;

174

vertex conspicuously large; vertex angle 45°, ocellar
angle 50°, ocelli large, tl-v ratio 0.15, touchline in-
complete; pubescence pale yellow, dark in ocellar re-
gion, on compound eyes sparse, yellowish.

Thorax: Moderately strong metallic lustrous on lat-
eral dorsum, scutellum and sides, background colour
dark; pubescence rather sparse, uneven, pale yellow.
Wings clear throughout; halteres, squamae and anti-
squamae pale yellow.

Legs: All dark, tibiae and tarsi 1-2 with dense yel-
low pubescence giving false impression of light
colour; trochanter 3 with small rounded knob, femur
3 swollen, strongly metallic lustrous, with triangular
processus strongly serrate apically, tibiae 3 with con-
spicuous apical spur bearing strong floccus; basitarsi 3
very much swollen, bearing dense, short, yellow, even
pubescence medially; other tarsi 3 normal.

Abdomen: Outline rather stout, background
colour dark, obliterated by strong metallic lustre and
moderately dense, yellowish, even pubescence; no
traces of pruinosity or lateral spots. S IV bears con-
spicuous paired appendages.

Genitalia: Anterior and posterior surstyle lobes sep-
arated by deep narrow sulcus; anterior surstyle lobe
bearing rather dense, yellow, erect, even pubescence
apically; posterior surstyle lobe showing scattered,
long, yellow pubescence, densest posteriorly. Cercus
rectangular, well protruding, bearing long, erect, yel-
low pubescence. Aedeagus stout, apical shaft part
short, fringed plates on thecal apex recumbent.

Body length 9 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Eyes separate widely,
ocellar angle 55°, frons lustrous throughout, partially
dulled laterally by light pruinosity. Legs: The basitar-
si 3 slightly but distinctly swollen, with a length/
width ratio about 3.0. Abdomen: Appendages on S
IV lacking.

Body length 9 mm.

Diagnosis. — The males of turkestanicus can be dis-
tinguished from other species in this group by the
large vertex, all dark legs, structure of tibiae 3 and
basitarsi 3 and strong metallic lustre; the females by
the swollen basitarsi 3 and dark legs.

Period of flight and distribution (fig. 86). — M.
turkestanicus occurs in Tadzhikistan and Turkestan,
and Iran, in mountain areas, flying there during May
and June.

Discussion. — The identification of the specimens,
some by Paramonov, seems to be correct; all speci-
mens match the original description.

Merodon xanthipous sp. n.

(figs. 44 a-c, 85)

Type-material. — & Holotype ‘Iran, Chiraz, 11.iv.1937,
coll. F. H. Brandt / L. crassicornis Sack, det. v. Doesburg’
(ZMAN). 19 Paratype ‘Iran, Rd. Chiraz Kazeroun, Ft. Sine-
Sefid, 19.iv.1937, coll. F. H. Brandt / L. crassicornis Sack,
det. v. Doesburg’ (ZMAN) [both localities at about 29° 30°
N, 52° E].

Description

Male. — Head: Antennae brown, antenna 3 orange,
upper margin slightly concave, apex acute, antennal
ratio 2.1; vertex angle = 35°, ocellar angle = 45°; pu-
bescence pale yellow, with hardly any dark pubescen-
ce in ocellar region; tl-v ratio 0.15, touchline incom-
plete; head without metallic lustre.

Thorax: Dark, lacking metallic lustre; katepister-
num, anepisternum and anepimeron with some dull-
ish blue tinge; pubescence sparse, brown, uneven,
moderately long; some diffuse pruinosity present on
sutures; vestigial pruinose bands can be seen at certain
angles on posterior dorsum. Wings clear; halteres,
squamae and antisquamae pale yellow.

Legs: All pale brown; trochanter 3 with low knob
bearing yellow hair-tuft, femur 3 slightly curved,
swollen, with rather high triangular processus show-
ing apical spur, bearing one strong and 5-6 lesser bris-
tles; tibiae 3 with apical floccus reaching down basal
half of basitarsi 3, inserted on drawn-out medial dis-
tal angle; basitarsi 3 swollen, bearing rather dense,
brownish yellow, erect, short, even pubescence
throughout, densest on medial face; other tarsi nor-
mal.

Abdomen: Very dark brownish red, tapering poste-
riorly, especially T II; slightly arcuate, faint, narrowly
interrupted whitish pruinose bands present on T II-
IV; pubescence pale brown, erect, uneven; S IV bear-
ing robust large paired appendages that are smaller
than in other species of this group, but still very con-
spicuous.

Genitalia: The anterior surstyle lobe rather spathu-
late, with dense, yellow, erect, even pubescence dor-
sally and apically, more scattered on other parts; sep-
arated from posterior surstyle lobe by shallow sulcus;
posterior surstyle lobe steeply rising, anteriorly high,
with distinct slight anteroventral angle and rather
dense, erect, yellow, uneven pubescence posteriorly;
cercus somewhat elongate, rather wide, with dense,
long, yellow pubescence; aedeagus moderately slen-
der, with marked subapical cavity, fringed plates on
thecal apex suberect, apical shaft part slightly length-
ened.

Body length 7 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Antenna 3 rounded,

Hurkmans: Monograph of Merodon

more obtuse; frons lustrous (not metallic) through-
out, eyes widely separate, ocellar angle 60°. Thorax:
Showing dull bluish-greenish lustre on dorsum, scu-
tellum and sides. Legs: Deeper brown than in male.
Abdomen: Stouter than in males, darker, pruinose
bands more marked; T II-V with a micaceous lustre.

Body length 7 mm.

Diagnosis. — The species can be distinguished in
both sexes by the relatively pale, uniformly coloured
legs; the males are also distinguished by the relatively
smaller appendages on S IV and the steeply rising
posterior surstyle lobe as well as by the absence of me-
tallic lustre.

The former identification as crassicornis Sack, pre-
viously applied to the holotype and paratype, is incor-
rect in view of contradictions of characters in the
types of xanthipous with the description of crassicor-
nis. In the females of the latter species the abdominal
pruinose bands are stated to be wide and clear, the an-
tennae are dark, the compound eyes bearing very
dense pubescence, the legs are black with yellow
markings; moreover bands of long white pubescence
are stated to be present on the posterior tergite mar-
gins. The male of crassicornis has not been described.

Period of flight and distribution (fig. 85). — The
species, known only from the types, occurs in south-
western Iran in April.

Etymology. — The epitheton xanthipous, to be
treated as a noun in apposition, refers to the golden
brown leg coloration, and is derived from the greek
xanthos, meaning blonde, and pous, meaning foot.

The clavipes group

Apomorphies: Anterior surstyle lobe elongate, den-
sely pubescent and strongly to enormously enlarged.

Description. — Head: Antennae with antennal ratio
1.4 to 2.0, antenna 3 mostly subacute; pubescence on
head whitish to yellow, usually dense on frons; face
often bluish lustrous; tl-v ratio in the males 0.3 to 0.5.
Thorax: Dark, lustrous on dorsum and scutellum,
mostly with strong blue lustre on katepisternum, an-
episternum and anepimeron; pubescence most vari-
able in colour density and length; some species have
slightly to strongly infuscate wings. Legs: Mostly all
dark, in some cases tibiae 1 and 2 basally with orange
markings; femur 3 usually much swollen, curved, in
some cases straight and slender; triangular processus,
tibiae 3 and tarsi 3 normally shaped. Abdomen:
Moderately slender to slender or stout; T II tapering
(less strongly in the females); arcuate pruinose bands
present on T II-IV which may in some species be dif-
ficult to see through the dense abdominal pubes-

175

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

cence. Genitalia: The anterior surstyle lobe very large,
rounded-rectangular, ridged laterally, usually with
short even pubescence throughout; posterior surstyle
lobe variously shaped, with dense pubescence, with
specialized hair-tufts in warnckei, aberrans, hamifer
and brevis, cercus variable; aedeagus mostly slender,
apical shaft part short to slightly lengthened, fringed

plates recumbent to oblique.

Merodon aberrans aberrans Egger

(figs. 45 a-d, 50 b, 89)

Merodon aberrans Egger, 1860: 10. Lectotype d (here desig-
nated): ‘Austria Alte Sammlung/ Merodon aberrans
Egger’ (NHMW) [examined].

Merodon obscuripennis Palma, 1863: 47. [No type material
mentioned in original description].

Merodon kneri Mik, 1867: 415. Syntypes: “Halicia’ [not ex-
amined]. ;

Merodon aberrans, Mik 1867: 417, 1883: 182; Bezzi 1900:
89; Sack 1913a: 435; Paramonov 1925: 153; Liepa 1969:
20; Gaunitz 1969: 82, 86; Van der Goot 1981: 215, 218;
Peck 1988: 166.

Lampetia aberrans, Oldenberg 1919: 388; Sack 1931, 302;
Séguy 1961: 174.

Merodon obscuripennis, Peck 1988: 166.

Merodon knerit; Mik 1883: 182; Peck 1988: 166.

Lampetia kneri; [sic!] Sack 1931: 302.

Material examined. — Albania: 8¢ (NHMW, RMNH);
Austria: 86 paralectotypes and 2? paralectotypes of aber-
rans, with same label as lectotype; Czechoslovakia: 19
(ZMAN); France: 476, 89 (MNHN); Greece: 236, 119
(KBIN, BMNH, JLR, ZMUC, WH, RMNH, ZMAN); Hungary: 3d
(NHMW, RMNH); Italy: 646, 462 (NHMW, BMNH, JLR,
ZMAN); Lebanon: 1d, 1% (KBIN, NHMW); Poland: 14, 19
(BMNH); Romania: 2d (BMNH); Turkey: 1366, 459
(BMNH, JLR, WH); U. S. S. R.: 156, 142 (NHMW, ZMAS);
Yugoslavia: 236 , 59 (NHMW, JLR, BMNH, ZMAN, RMNH).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apex obtuse to subacute, antennal
ratio 1.6; pubescence yellow, with dark tuft in ocellar
region; vertex angle 40°, ocellar angle 50°, tl-v ratio
0.4; compound eyes showing white pubescence; face
variably bluish lustrous, mostly only weakly.

Thorax: Blue lustrous on lateral dorsum and scutel-
lum, katepisternum, anepisternum and anepimeron;
pubescence yellow, moderately dense, with dark
interalar band mostly clear. Wings troubled evenly in
distal half, anterior veins pale; halteres, squamae and
antisquamae pale yellow.

Legs: All dark, all femora slender; femur 3 straight;
trochanter 3 smooth, femur 3 bearing normally
shaped triangular processus bearing 6-11 bristles; bas-
itarsi 3, tibiae 3 and tarsi 3 normal. in most specimens
tibiae and tarsi with bright yellow pubescence.

Abdomen: Dark, with characteristic rather strong
bluish overall lustre; T II strongly tapering; T II-IV

176

with slightly arcuate, well interrupted pruinose
bands; vague mainly yellow pruinose lateral spots on
T II present in some specimens; S IV moderately
arched, deeply, narrowly emarginate posteriorly.
Genitalia: Anterior surstyle lobe large, anteriorly
bearing dense, short, even, yellow pubescence, with
scattered pubescence throughout, showing lateral
ridge; coalescent with posterior surstyle lobe; the lat-
ter bearing low anteroventral projection with dense,
long, yellow pubescence; posterior surstyle lobe with
long, dense, yellow pubescence. Cercus conspicuous,
rather quadrate with rounded corners. Aedeagus
smooth on outer face, moderately slender, apical shaft
part short, fringed plates on thecal apex recumbent.
Body length 12-16 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Antenna 3 more ob-
tuse; frons laterally pruinose, lustrous on median one-
third or just over; ocellar angle = 40°, pubescence on
frons less dense. Thorax: Pubescence brownish, inter-
alar band less clear; vague longitudinal pruinose
bands often present; wings in some specimens more
clearly infuscate distally. Abdomen: T Il less tapering,
pubescence very dense in some specimens; T V rela-
tively narrow, posteriorly more or less acute.

Body length 12-15 mm.

Diagnosis. — As the slender femora 3 are not found
in other species and the blue lustre is uncommon in a
species of this large size, aberrans is hardly ever con-
fused with other species; only hamifer is very similar
but can be separated at once by its deformed triangu-
lar processus; brevis is superficially similar but stouter,
lacks the interalar band in the males which moreover
have slightly different genitalia (see fig. 45); splendens
has a wider, shorter abdomen, denser pubescence and
the femora 3 are much more swollen than in aber-
rans.

Period of flight and distribution (fig. 89). — The
species occurs from May through July in central
Europe and the Mediterranean, in southern parts of
Russia, Ukraine and in Asia Minor.

Merodon aberrans ssp. flavitibius Paramonov

Merodon aberrans ssp. flavitibius Paramonov, 1925: 153.
Syntypes: ‘Armenia’ [not examined].

Merodon aberrans ssp. flavitibius, Liepa 1969: 20; Peck
1988: 166.

Lampetia aberrans ssp. flavitibius, Sack 1931: 302.

Description
Subspecies flavitibius differs from the nominate
subspecies only in the colour of the legs, which in the

present subspecies show yellow markings on the basal
parts of tibiae 1 and 2.

Period of flight and distribution. — This subspecies

occurs in Armenia.

Merodon aberrans isperensis ssp. n.
(fig. 90)

Type-material. — d Holotype “Turkiye, Rize, Ovit Pass
[near Ispir], S side 1800 m, 16.vii.1987, leg. J. A. W. Lucas’
(ZMAN). 436 , 249 paratypes, all from Turkey, all in (LR)
and all leg. Lucas, except where noted: Topotypic 19,
31.vii.1983; 1d, 19, 16.vii.1986; 15 ‘Turkey Hakkari,
Suvarihalil pass 1250 m W side nr. Habul Deresi
13.vi.1984’; 49 Turkey Hakkari, Tanin-Tanin pass W side
1700 m, 12.vi.1984; 12 ‘Turkey, Kars, Handere 2100
2200 m, 20 km W of Sarıkamıs, 1.vili.1983’; 26, 19
‘Turkey, Siirt 21 km W of Uludere, 12.vi.1984’; 3d
‘Turkey, Kars, 25 km W. of Sarikamis 5.viii.1985, 2100 m,
C. J. Zwakhals leg.’; 36 “Turkey, Kars, Bahnstat. Soganli
W. Sarıkamıs 2100 m 5.vii.1985 W. Schacht’; 19 “Turkey,
Hakkari, Sat Dagi, Varegös, SW Yiiksekova 1700 m
29.vi.1985 W. Schacht’; 134, 49 ‘Turkey, Bingöl, Buglan
Gegidi 1640 m 5.vii.1986’; 16, 19 “Turkey, Kars 11 km E
of Karakurt 1450 m 6.vii.1986; 6, 29 “Turkey, Kars, 8
km W Sarikamis, 2000 2050 m 6-9.vii.1986’; 2d, 19
“Turkey, Kars, Handere 20 km W Sarıkamıs 2000 m
7.vii.1986; 18,1% “Turkey, Kars, 12 km S Sarikamis 2050
m, 7.vii.1986; 106, 62 ‘Turkey, Erzurum (Kayak)
Palandöken Dagi 2200-2300 m 1.vii.1986’. See map show-
ing Turkish localities (fig. 79).

Description

Male. — Head: Pubescence more grey than in no-
minate subspecies, otherwise similar.

Thorax: Slenderer than in nominate subspecies,
bearing less dense pubescence; interalar band less
clear.

Legs: Similar to nominate subspecies, but pubes-
cence pale yellow, not golden.

Abdomen: Clearly less wide than in nominate sub-
species, showing slate grey instead of bluish lustre;
pubescence slightly less dense.

Genitalia: completely similar to nominate subspe-
cies.

Body length 10-15 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Thorax: The infuscation of
the apical wing portions is less strong than in females
of the nominate subspecies.

Period of flight and distribution (fig. 90). — This
subspecies occurs in June through August in the
mountains of eastern Turkey. The distributions of
the nominate form of aberrans and of the ssp. isperen-
sis are shown in figs. 89-90.

Hurkmans: Monograph of Merodon

Etymology. — The adjective isperensis refers to the
type-locality in the vicinity of the town of Ispir in
north-eastern Turkey.

Merodon brevis Paramonov

(figs. 46 a-c, 86)

Merodon brevis Paramonov, 1925: 157. Holotype d:
‘Inaklu, Bez. Etschmiadzin, Gouv. Erivan, (Armenien),
Siidlicher Abhang des Alagéz, 4500’ [Gora Aragac, 40°
29’ E, 44° 12’ E]’ [Armenia] [not examined, considered
lost].

Merodon brevis, Peck 1988: 168.

Lampetia brevis, Sack 1931: 308.

Material examined. — U. S. S. R.: 1¢, Armenia, Abaran
[Aparan, 40° 34’ N, 44° 21 E], M. Karny-larich, 8000’,
10.vii.26. A. Shelk / Lampetia brevis Param. d, S.
Paramonoy det./ Coll. J. Villeneuve, Lampetia brevis Param.
(KBIN).

Description

Male. — Head: Antenna with articles 1-2 brown, 3
conspicuously orange, with upper margin convex, ap-
ex subacute, antennal ratio 1.7; pubescence white,
long, dense on frons, sparser on face, golden on occi-
put, darker on vertex; face blue lustrous, partially
with dense, pale pruinosity, vertex angle 35°, ocellar
angle 50°, tl-v ratio 0.5.

Thorax: Dark, lateral dorsum and scutellum, kate-
pisternum, anepisternum and anepimeron greenish
lustrous, showing rather dense, golden, erect pubes-
cence; two faint longitudinal pruinose bands present
on dorsum; no interalar band. Wings rather clear,
very pale brown veined; halteres, squamae and anti-
squamae yellow, halteres deepest coloured.

Legs: Dark, but tibiae 1 and 2 orange on basal half.
Trochanters 3 smooth, femora 3 swollen, steel grey
lustrous, bearing rather large triangular processus
with strongly serrate distal margin bearing 5-6 bris-
tles; tibiae and tarsi normally shaped, tarsi 3 with
golden pubescence laterally, dark on upper face.

Abdomen: Relatively small, rather stout; dark, with
interrupted well-defined arcuate interrupted pruinose
bands; pubescence pale anterior and dark posterior,
longest laterally; no lateral spots; S dark, S IV vaulted,
moderately deep emarginate posteriorly.

Genitalia: Anterior surstyle lobe elongate, recurved
dorsally, apically wide, stalked, with clear lateral
ridge, with dense, short, erect, yellow pubescence
throughout, bearing small dorsal basal projection;
posterior surstyle lobe rather quadrate, two more scle-
rotized areas on ventral margin bearing dark bristle-
like pubescence; pubescence otherwise as on anterior
surstyle lobe but longer laterally. Cercus rather small,
bearing short, dense, erect, yellow, uneven pubes-
cence. Aedeagus moderately slender, apical shaft part
short, fringed plates recumbent.

177

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Body length 12 mm.
Female. — Unknown.

Diagnosis. — M. brevis most closely resembles aber-
rans from which it is distinguished by the shorter
body, more elongate anterior surstyle lobe and uni-
formly coloured thoracic pubescence; the areas of spe-
cialized pubescence on the posterior surstyle lobe are
not found in aberrans. Moreover, this species differs
from aberrans by the orange markings on the legs. In
this respect brevis resembles aberrans ssp. flavitibius.
No material of the latter taxon has been examined,
but the other characters mentioned should be suffi-
cient to distinguish these taxa.

Period of flight and distribution (fig. 86). — M. bre-
vis has been found in the mountains of Armenia [ap-
prox. 40° 30° N, 44° 20° E], where it was collected in

July.

Discussion. — The identification as brevis, by
Paramonov, is considered reliable; the specimen ex-
amined originates from a site less than 20 km from
the locus typicus.

Merodon clavipes (Fabricius)
(figs. 47 a-c, 89)

Syrphus clavipes Fabricius, 1781: 427. Lectotype d (here
designated): ‘clavipes in Sehestedt & Tonder Lund col-
lection (ZMUC) [examined].

Merodon clavipes var. albus Paramonov, 1927: 90. Syntypes:
‘Bezirk Balta, Gouv. Odessa’ [Ukraine, U. S. S. R.] [not
examined].

Merodon clavipes var. ater Paramonov, 1927: 91. Syntypes:
locality unknown [not examined].

Merodon clavipes var. niger Paramonov, 1927: 90. Syntypes:
‘Bezirk Balta, Gouv. Odessa [not examined].

Merodon canipilus Rondani, 1865: 131. Lectotype d (here
designated): ‘52’ [number refers to description of canipi-
lus] in coll. Rondani (LSF) [examined].

Musca clauda Villers, 1789: 463. Syntype(s): ‘Gallia
Australiori’ [the southernmost of France] [not examined].

Musca curvipes Gmelin, 1790: 2871. Published in synonymy
with Syrphus clavipes Fabricius, 1781.

Syrphus gravipes Rossi, 1790: 286. Syntypes: ‘in provinciis
Florentina et Pisana’ [Toscana, Italy] [not examined].
Lampetia sacki Paramonov, 1937a: 3. Lectotype © (here
designated): ‘Chiklana’ [Chiklana de la Frontera, near

Jerez, southern Spain] (ZBSM) [examined]. Syn. n.

Merodon senilis Meigen, 1822: 356. Lectotype 9 (here des-
ignated): ‘senilis’ (NHMW) [examined].

Syrphus clavipes, Fabricius 1787: 337; Rossi 1790: 286;
Fabricius 1794: 292.

Lampetia clavipes, Meigen 1800: 34; Coquillett 1910: 557;
Oldenberg 1919: 388; Sack 1931: 310; Séguy 1961: 176.

Lampetia clavipes var. alba; Sack 1931: 311.

Lampetia clavipes var. atra; Sack 1931: 311.

Lampetia clavipes var. nigra; Sack 1931: 311.

178

Merodon clavipes Fabricius 1805: 195; Latreille 1804: 443;
Meigen 1822: 351; Macquart 1828: 291; Macquart
1834: 513; Westwood 1840: 137; Macquart 1842: 70;
Rondani 1845: 256, 259; Macquart 1849: 466; Walker
1849: 597; Rondani 1857: 54, 62; Schiner 1857: 410;
Schiner 1862: 344; Palma 1863: 46; Portschinsky 1877:
184; Strobl 1893: 76; Bezzi 1895: 16; Bezzi 1900: 89;
Strobl 1900: 593; Verrall 1901: 559; Villeneuve 1903:
115; Sack 1913a: 433; Paramonov 1925: 146; Gil
Collado 1930: 243, 246; Gaunitz 1969: 83, 86;
Delfinado & Hardy 1975: 343; Marcos Garcia 1985:
197; Hurkmans 1985: 69; Bradescu 1986: 123; Peck
1988: 165, 168 (see for additional references Kertész,
1907: 274).

Merodon clavipes var. albus, Liepa 1969: 20; Peck 1988: 169.

Merodon clavipes var. ater. Liepa 1969: 20; Peck 1988: 169.

Merodon clavipes var. niger. Liepa 1969: 20; Peck 1988: 169.

Milesia clavipes, Latreille 1810: 331.

Merodon canipilus, Peck 1988: 169.

Lampetia canipila, Sack 1931: 310 (as syn. of clavipes).

Musca clauda; Kertész 1907: 274; Peck 1988: 168.

Musca curvipes, Peck 1988: 168.

Merodon curvipes, Meigen 1803: 74 (erroneous); Peck 1988:
168.

Lampetia curvipes, Sack 1931: 310 (as syn. of clavipes).

Syrphus gravipes, Peck 1988: 168.

Merodon senilis, Peck 1988: 168.

Material examined. — Algeria: 328, 59 (MNHN, BMNH,
ZMAN, ZMUC); Austria: 106, 22 (NHMW); France: 1436,
612 (MNHN, NHMW,Bonn, BMNH, ZMUC, ZMAN); Greece:
1178, 372 (ZMUC, den Hollander, NHMW, Bonn, BMNH,
Thessaloniki, ZMAN, JLR, WH); Hungary: 5d, 42 (BMNH,
ZMUC); Italy: 1168, 579 (ZMUC, JLR, NHMW, BMNH, ZMAN,
WH); Morocco: 19 (BMNH); Spain: 2 Paralectotype of
Lampetia sackî: ‘Chiklana’ (ZSBM) and 206, 52 (ER,
ZMUC, BMNH, Bonn, ZMAN); Sweden: 12, Bohuslan, leg.
Malm, 1862 (MNHN); Switzerland: 16, 22 (ZMUC);
Yugoslavia: 31d, 52 (BMNH, NHMW, JLR, ZMAN).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin concave, apex subacute, vertex angle 50°,
ocellar angle 60°, antennal ratio 1.8; pubescence yel-
lowish, darker in ocellar region; face dense white
pruinose; tl-v ratio 0.4.

Thorax: Dark, slightly metallic lustrous on lateral
dorsum, scutellum, katepisternum, anepisternum and
anepimeron, pubescence yellow to brown, erect, rath-
er dense, with dark interalar band. Wings troubled
evenly, halteres, squamae and antisquamae pale yel-
low.

Legs: All dark, trochanters 3 with high ridge
shaped like chisel edge; femora 3 curved, very strong-
ly swollen, often with oily lustre, and golden pubes-
cence; triangular processus normally shaped, tibiae 3
and tarsi 3 normal.

Abdomen: Dark; T II-IV with interrupted arcuate
wide pruinose bands and dense pubescence, yellow
on T II and III (anteriorly) and orange on posterior
abdomen; T II clearly but not very strongly tapering,

normally without lateral spots; S IV strongly vaulted,
posterior margin widely and deeply emarginate.

Genitalia: The anterior surstyle lobe extremely
large, bearing short, dense, yellow, even pubescence
throughout, laterally ridged; posterior surstyle lobe
coalescent, steeply rising, rather high, long, uneven,
with less dense pubescence. Cercus well protruding,
angular, with dense, long, yellow pubescence.
Aedeagus slender, smooth on outer face, somewhat
concave subbasally, apical shaft part slightly length-
ened, fringed plates on thecal apex recumbent-
oblique.

Body length 17-23 mm.

Female. — Quite different from the males, as they
lack the dense abdominal pubescence. Apart from
sexual dimorphism the main other differences are:
Head: Frons whitish pruinose, leaving lustrous mid-
stripe occupying one-fifth of width. Thorax:
Pubescence greyish, shorter than in males, rather
even, no interalar band. Legs: The femur 3 slightly
less swollen, often pubescence duller; compared with
other Merodon females, still very much swollen.
Abdomen: Pubescence overall much less conspicu-
ous; dense only in anterolateral areas of T II, and on
pruinose bands; in some specimens large portions of
the tergites bald altogether; colour of the pubescence
mainly dark yellow to pale; in many specimens pale
orange lateral spots can be seen.

Body length 15-22 mm.

Varieties of clavipes and variation. — In the var. 4/-
bus the pubescence is white throughout, except for a
black interalar band in both sexes with blue back-
ground lustre. In the var. ater the thoracic dorsum
shows only black pubescence; of this variety only the
males are known. In the var. niger the pubescence is
black throughout, except on the face, while the prui-
nose abdominal bands are almost or completely ab-
sent in both sexes. No types of either of the varieties
have been examined; they are considered to be lost
(Liepa 1969: 4).

Apart from the described varieties and size varia-
tion the specimens of this species are remarkably uni-
form. A few aberrant individuals totally lack the pu-
bescence on the abdomen and have strangely vaulted
posterior tergite margins. These specimens are not
considered to belong to a separate variety. Possibly
they have suffered from frost damage, known to pro-
duce aberrances in insects.

Diagnosis. — M. clavipes is easily distinguished
from other species in this group by its size. Moreover,
the combination of enormously swollen femora 3 and
chisel-edged trochanters 3 does not occur in males of
any other species; the degree of swelling in the femo-

Hurkmans: Monograph of Merodon

ra 3 in the females far exceeds that of any other
Merodon currently known.

Period of flight and distribution (fig. 89). — M. cla-
vipes occurs from late April through August in south-
ern Europe, the Mediterranean and Asia Minor,
though it may occasionally occur more northward;
the specimen from Sweden seems unquestionable; re-
ports of clavipes from Great Britain have been serious-

ly doubted by Verrall (1901).

Biology. — The territorial behaviour of the males
has been described by Hurkmans (1985). In the field
the males visit flowers, mainly of Umbelliferae, and
are conspicuous, but the females fly close to the soil
through the vegetation.

Discussion. — The types of canipilus, sacki and sen-
ilis have been examined, and found conspecific with
clavipes. The types of clauda, curvipes and gravipes
have not been examined. The original description of
clauda leaves hardly any doubt that this is a synonym
of clavipes this synonymy has been considered by
Kertész (1907: 274), while Peck (1988: 168) also
names clauda as synonym of clavipes. Musca curvipes
was originally published as a synonym of Syrphus cla-
vipes. Syrphus gravipes refers to the publication of the
female of clavipes, a possibility already considered in
the original description.

Merodon cupreus sp. n.
(figs. 48 a-c, 88)

Type-material. - & Holotype “Turkey, Kars, Handere
2100-2200 m, 20 km W of Sarıkamıs, 1.viii.1983, leg. J. A.
W. Lucas’ (ZMAN). Paratypes: 298, 59: 244,49 topotyp-
ic, with dates ‘1.viii.1983’ and ‘2.viii.1983’ (LR), 12 with
same data (ZMAN); 36 ‘Turkey, Hakkari S. of Yiiksekova
28.vi.1986 W. Schacht’ (LR); 26 “Turkey, Kars, 8 km W.
of Sarıkamıs 2000-2050 m, 9.vii.1986' leg. J. A. W. Lucas
(JLR).

Description

Male. — Head: Antennae dark, antenna 3 cordate,
apex subacute; antennal ratio 1.4; vertex angle 30°,
ocellar angle 35°, pubescence yellow, often with dark
tuft in ocellar region; face and occiput white pruino-
se, somewhat bluish lustrous; tl-v ratio 0.35.

Thorax: Dark, metallic lustrous on lateral dorsum
and scutellum, katepisternum, anepisternum and an-
epimeron; pubescence deep yellow to brown, even,
rather dense, with dark interalar band. Wings clear;
halteres, squamae and antisquamae pale yellow.

Legs: Dark, but tarsi laterally with golden pubes-
cence, contrasting; trochanter 3 sharply keeled, with
chisel edge, femur 3 rather swollen, slightly lustrous,
with normally shaped triangular processus bearing 6-

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

7 bristles on very strongly serrate distal margin; tibiae
3 normal, basitarsi 3 slightly swollen.

Abdomen: Dark, no lateral spots on T II; T II-IV
with white pruinose bands, straight on T II, arcuate
on T III-IV; interrupted on T II-III, continuous on T
IV; pubescence golden to brassy or orange, rather
dense, recumbent; T II tapering; S IV vaulted, deep-
ly, rather narrowly emarginate posteriorly.

Genitalia: The anterior surstyle lobe large, round-
ed, with basal projection on ventral margin, and
dense, short, even, yellowish pubescence anteriorly;
laterally weakly ridged; the posterior surstyle lobe sep-
arated by shallow sulcus, low, elongate, bearing yel-
low, moderately long, uneven pubescence along ven-
tral and posterior margins. Cercus small but well
protruding, with dense, long, yellow pubescence.
Aedeagus moderately slender, fringed plates on thecal
apex oblique, apical shaft part short.

Body length 12-15.5 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons tapering poste-
riorly, sides white pruinose, lustrous midstripe opccu-
pying one-third of width; pubescence grey, but dark
tuft present in ocellar region; ocellar angle 60°.
Thorax: Dorsum dull, pubescence grey, dark interalar
band present. Legs: Dark, with grey pubescence; on
tibiae and tarsi yellow-grey pubescence with some
contrast as in d but weaker; trochanter 3 smooth; tri-
angular processus as in males. Abdomen: Stouter than
in male, with rather dense grey pubescence, pruinose
bands moderately wide, continuous, arcuate; tergite
V dark, with scattered grey pubescence.

Body length 13 mm.

Diagnosis. — At first sight, cupreus seems to be a
small clavipes. Differences between those species are:
antennal ratio in cupreus 1.4, in clavipes 1.8; antenna
3 with upper margin convex in cupreus, concave in
clavipes, triangular processus stronger serrate in cupre-
us than in clavipes, in the males of clavipes the anteri-
or surstyle lobe is much larger than in those of cupre-
us, in the female cupreus lateral abdominal spots
lacking in, present in clavipes, size in clavipes 15-23
mm, as opposed to 12-15.5 mm in cupreus. The dif-
ferences between dzhalitae (discussed hereafter) and
cupreus are: antenna 3 with upper margin convex in
cupreus, concave in dzhalitae, on the hind legs the tar-
si Il in dzhalitae are almost as long as the basitarsi; in
cupreus they are less than half as long. Moreover, in
the males of cupreus the S IV is deeply emarginate
posteriorly while this element is entire in dzhalitae.

Differences with the description of M. clavipes var.
albus, stated to have all black and white pubescence in
both sexes, are: The size in clavipes var. albus is great-
er, in cupreus the pubescence is not black and white in

180

both sexes. Moreover, the differences (with respect to
clavipes) already mentioned apply. The 2 cupreus are
of the same size and habitus as those of femoratus.
They are distinguished by their lack of pruinose
bands on the thorax and much denser overall pubes-
cence; also the shape of the antenna 3 is different:
ovoid to cordate in cupreus, more elongate in femora-
tus.

Period of flight and distribution (fig. 88). — M. cu-

preus occurs in July and August in eastern Turkey.

Etymology. — The adjective cupreus refers to the
colour of the larger portion of the abdominal pubes-
cence in the male of this species.

Merodon dzhalitae Paramonov

(fig. 99)

Merodon dzhalitae Paramonov, 1927: 89. Holotype d:
‘Hissar, Yalta’ [south slope of the Crimea mountains,
Crimea, Ukraine] [not examined, considered lost].

Merodon dzhalitae, Liepa 1969: 20; Van der Goot 1981:
215; Peck 1988: 169; Zimina 1989: 24.

Lampetia dzhalitae, Sack 1931: 313.

Description

The description of dzhalitae presented here is based
on the original description, on Sack (1931: 313) and
on Zimina (1989: 24) who described recently collec-
ted material of both sexes.

Male. — Head: Antennae dark, antenna 3 yellow-
brownish, upper margin concave, lower margin con-
vex, apex acute; face, frons and vertex with yellow pu-
bescence, face sparsely pruinose; compound eyes with
yellow pubescence; tl-v ratio 0.5.

Thorax: Dorsum dark with some weak blue lustre,
with conspicuous orange pubescence throughout,
scutellum concolorous, duller; wings slightly trou-
bled; reddish pubescence clear, interalar band some-
times present.

Legs: All dark, with yellow pubescence, femur 3
normal, resembling this element in avidus [therefore
less swollen than in e.g. clavipes|, triangular processus
with strong apical pedestal (cf. Zimina 1989: fig. 2);
on hind legs the second tarsi almost as long as basitar-
Si.

Abdomen: T II slightly tapering only, clearly bluish
lustrous, T III-IV with white pruinose bands, just
interrupted on T III, just continuous on T IV; pubes-
cence reddish, but black, short and recumbent on
posterior margins of T II-III. S IV with posterior
margin entire, keeled.

Body length 13 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Habitus: Pubescence overall

paler than in male. Head: With broad lustrous mid-
stripe on frons. Antenna 3 less long than in d, ovoid,
dark with paired yellow spots ventrally. Legs: Tibiae
brownish basally. Abdomen: T II showing nearly tri-
angular greenish metallic lustrous lateral spots which
are reddish translucent mediad. Pubescence dark,
short and recumbent on T II-III, reddish elsewhere.
Body length 13 mm.

Diagnosis. — The differences with respect to cupre-
us, which seems to resemble dzhalitae most closely,
are as follows. Third antennal article concave in
dzhalitae, convex in cupreus tarsi II on hind leg as
long as basitarsi in dzhalitae, only half as long in cu-
preus, and S IV entire in dzhalitae, deeply emarginate
in cupreus.

Period of flight and distribution (fig. 99). — The
species occurs in August and probably is endemic to
the Crimea.

Discussion. — In view of the extensive original de-
scription and the full account by Zimina (1989) it
seems clear that dzhalitae is a distinct species. The de-
scription of the recently collected material by Zimina
(1989) matches the original description on all signifi-
cant points. M. dzhalitae has been tentatively assigned
to the clavipes group on account of its external simi-
larities with e.g. clavipes and cupreus.

Merodon hamifer Sack
(figs. 49 a-b, 88)

Merodon hamifer Sack, 1913a: 436. Lectotype d (here des-
ignated): ‘Anatolien Ak-chehir [Aksehir] Korb 1900/
Mer. hamifer Sack det. Sack/ coll. Lichtwardt/ coll. D. E.
I. Eberswalde’ (DEIC) [examined].

Merodon hamifer, Paramonov 1926b: 322; Paramonov
1927: 78; Peck 1988 : 171.

Lampetia hamifera; Sack 1931: 318.

Material examined. — Greece: 1d ‘Samos, Ampelos
22.vi.1932, Werner (NHMW); Turkey: Paralectotype 9,
same data as lectotype.

Description

M. hamifer is similar to aberrans except in the fol-
lowing details.

Male. — Head: Antennal ratio 2.0; antennae all
orange. Thorax: Pubescence less dense. Legs: All tib-
iae and tarsi orange, tibiae 3 basally brown; femora 3
bearing modified triangular processus, basitarsi 3
showing apical projection on medial face. Genitalia:
Pubescence on posterior surstyle lobe denser.

Body length 13.5-14.5 mm.

Female. — Except for sexual dimorphism, differing

HURKMANS: Monograph of Merodon

from the male as follows. Head: Compound eyes sep-
arate, frons lustrous, slightly dulled down sides; an-
tennae orange, antennal ratio 2.8. Legs: Similar to
those of hamifer male, triangular processus slightly
smaller. Abdomen: Pruinose bands wider; tergite V
narrower.

Body length 14 mm.

Diagnosis. — This close relative of aberrans can be
distinguished from all other species by the shape of
the triangular processus in both sexes.

Period of flight and distribution (fig. 88). — M. ha-
mifer occurs in the Greek Aegean islands and western
Turkey, where it flies in June.

Merodon karadaghensis Zimina

Merodon karadaghensis Zimina, 1989: 24. Holotype d:
‘Karadagh, Crimea, 5. ix. 1984, leg. Zimina’. Moscow
Zoological Institute [not examined].

It appears that this species may be related to M.
dzhalitae, with which it is compared by Zimina
(1989). The differences are stated to be in the relative
length of the tarsal joints, the outline of the triangular
processus on the hind femora, while the abdominal
pruinose bands also slightly differ.

Diagnosis. — After Zimina (1989), M. karadaghen-
sis may be distinguished from M. dzhalitae by the
length ratio of the basal tarsal joints (II:I) on tarsus 3,
which is about 0.5 in karadaghensis and 0.8 in dzhal-
itae,and also by the abdominal pruinose band on T
IV being chevron-shaped in karadaghensis while this

is more rounded in dzhalitae.

Discussion. — This species is provisionally regarded
as belonging to the c/avipes group, on the grounds of
resemblance to M. dzhalitae.

Merodon lusitanicus sp. n.
(figs. 50 a, 93)
Type-material. — 2 Holotype ‘Portugal, Algarve,

Quarteira 27.iv.1985, J. A. W. Lucas’ (ZMAN); paratype 9,
same data, but locality: ‘Vilamoura’ (JLR).

Description
Female. — Head: Antennae black, antenna 3 with
upper margin concave, apex rounded, antennal ratio
2.8; pubescence white on face, black on frons and ver-
tex, pale brown on occiput; frontal sides whitish prui-
nose, leaving lustrous midstripe on median one-third;
ocellar angle 60°.
Thorax: Dark, dorsum with 4 vague pruinose

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

bands and yellow pubescence; pubescence duller on
scutellum and sides, katepisternum, anepisternum
and anepimeron steel grey lustrous. Wings clear; hal-
teres, squamae and antisquamae yellow.

Legs: All dark, joints lighter; femora 3 extremely
swollen for a female, triangular processus showing
strongly serrate distal margin bearing 10 bristles.

Abdomen: Dark with some purplish lustre, trian-
gular in outline, with clear lateral spots on T II com-
bined with dense erect white pubescence; pubescence
otherwise sparse; T II-IV bearing rather narrow, con-
spicuous widely interrupted somewhat arcuate prui-
nose bands; T V acute, all dark.

Body length 11-12 mm.

Male. — Unknown.

Diagnosis. — M. lusitanicus can be separated from
aberrans by the lateral spots on T II and the swollen
femora 3; /usitanicus is too dark to be confused with
females of avidus, nigritarsis or distinctus, its habitus
can be compared with that of alexeji from which it
differs in larger size, longer antennae and more swol-
len femora 3.

Period of flight and distribution (fig. 93). — This
species, known only from the types, has been found
in Portugal in April.

Discussion. — In view of the shape, width and posi-
tion of the abdominal pruinose bands, relative bald-
ness of the abdomen with purplish lustre, and gener-
al habitus, /usitanicus is provisionally regarded as
closely related to aberrans, and therefore assigned to
the clavipes group.

Etymology. — The adjective /usitanicus refers to the
classical name for Portugal, Lusitania.

Merodon quadrinotatus (Sack)
(figs. 51, 85)

Lampetia quadrinotata Sack, 1931: 324. Holotype 2:
‘Mesopotamia’ [Iraq] [not examined, considered lost].
Merodon quadrinotatus, Peck 1988: 173.

Material examined. — Turkey: 19 ‘Hakkari, Suvarihalil
Pass W side 2300-2400 m 11.viii.1983 leg. J. A. W. Lucas’
(ZMAN).

Description

Female. — Similar to the female of clavipes, but with
the following differences. Head: Frons much tapering
posteriorly, laterally whitish pruinose leaving a lus-

trous midstripe occupying a quarter of the width; pu- _

bescence white, but grey on occiput. Thorax: Grey
pubescence on dorsum, with conspicuous black inter-

182

alar band. Legs: Pubescence grey to black, back-
ground colour deep black.

Abdomen: Black, pubescence dense and black
throughout but with two pairs of very conspicuous
white spots, one each on T HI-IV, and weak latero-
posterior triangular spots of grey pubescence on same
tergites.

Body length 14 mm.

Diagnosis. — The dense black, white-dotted pubes-
cence of the abdomen is unique and at once separates
quadrinotatus from all other species currently known.

Period of flight and distribution (fig. 85). — The
species is known from ‘Mesopotamia’ [Iraq] (month
not reported by Sack) and the eastern Turkish moun-
tains where it was collected in August.

Discussion. — Sack (1931) did not indicate a de-
pository of the holotype. He did provide a reasonably
clear description of guadrinotatus. Given the unique
features of the species the identification of the
Turkish specimen is unquestionable. Since quadri-
notatus is believed to be a close relative of clavipes the
species is assigned to the c/avipes group.

Merodon splendens sp. n.
(fig. 47 d, 91)

Type-material. — ¢ Holotype ‘Sardaigne St. Ussassai
16.v.1977 P. Goeldlin’ (LAU).

Description

Male. — Head: Antennae blackish, antennae 3
rounded-subacute, upper margin about straight, ver-
tex angle 30°, ocellar angle 40°, tl-v ratio 0.4; pubes-
cence pale yellow, face greenish lustrous, yellow prui-
nose; slight dark tuft present in ocellar region; oral
margin weakly protruding.

Thorax: Dorsum and sides dark, weakly bluish lus-
trous on anepisternum, katepisternum and anepime-
ron; pubescence moderately dense, erect, pale yellow,
with dark interalar band; no pruinose bands present.
Wings clear, halteres, squama and antisquama pale
yellow.

Legs: Black, with yellow pubescence; trochanters 3
with curved low chisel-edge; femora 3 very much
swollen, with some oily lustre, curved.

Abdomen: T II strongly tapering, T III-IV cylin-
drical. Lateral spots on T II weak, micaceous; prui-
nose bands on T II-IV very clear, interrupted, arcu-
ate; pubescence moderately dense, yellow on T II,
deep orange on T III-IV.

Genitalia: Anterior surstyle lobe reniform, pubes-
cence, although dense, so extremely short that it is in-
conspicuous; posterior surstyle lobe coalescent, much
bulging outward anteriorly, rounded, with short,

erect, moderately dense pubescence. Cercus slightly
protruding, bearing short, pale pubescence. Aedeagus
slender, with smooth outer face, subapical cavity well
marked, somewhat concave subbasally; fringed plates
on thecal apex recumbent.

Body length 14.5 mm.
Female. — Unknown.

Diagnosis. — M. splendens is a close ally of clavipes
and velox. From the latter it is distinguished by the
clear wings. The differences with clavipes, apart from
the smaller size, are: Antennae more slender, pubes-
cence on head, thorax and abdomen less dense and
shorter, T II more tapering, III-IV cylindrical.
Differences with both species are: Anterior surstyle
lobe more curving dorsad, differently shaped (cf. figs.
46 a and d), pubescence much less dense and very
much shorter on anterior and posterior surstyle lobes
and cercus. M. splendens can be distinguished from
cupreus and dzhalitae by the swollen femora 3. The
same character also separates sp/endens from brevis, to
which it is slightly similar on account of its habitus.
Moreover, the anterior surstyle lobe is quite different-
ly shaped in brevis. It differs from aberrans by the
strongly swollen femora 3 and denser pubescence.

Period of flight and distribution (fig. 91). — M.
splendens is known only from the holotype, collected
in Sardinia, in May.

Etymology. — The adjective splendens is latin and
means splendid or beautiful. It was given in view of
the colourful pubescence of this species.

Merodon velox Loew

(figs. 47 a-c, 91)

Merodon velox Loew, 1869: 253. Lectotype d (here desig-
nated): ‘Rhodus/ Alte Sammlung’ (NHMW) [examined] .

Merodon velox var. anathema Paramonov, 1925: 149.
Holotype 2: ‘Mons Takalta, prope Kulp,
Transkaukasien’ [in Turkey, approx. 38° 30 N, 41° 05 E]
[not examined, probably lost].

Merodon velox var. armeniacus Paramonov, 1925: 147.
Syntypes: 126 and 19 ‘Eriwan’ and ‘Ordubad (Erivan)’
[Armenia] [not examined, probably lost].

Merodon velox, Sack 1913a: 434; Paramonov 1925: 147;
Paramonov 1926a: 15; Peck 1988: 175.

Merodon velox var. anathema, Liepa 1969: 20.

Merodon velox var. anathemus, Peck 1988: 175.

Merodon velox var. armeniacus, Liepa 1969: 20; Peck 1988:
175.

Lampetia velox, Sack 1931: 331.

Material examined. — Greece: 2 Paralectotype with same
label as lectotype; 1006, 732 (BMNH, NHMW, RMNH,
ZMAN, JLR, WH); Italy: 26, 12 (NHMW); Turkey: 16d and
32 paralectotypes ‘As. Minor/ Alte Sammlung’ (NHMW);

Hurkmans: Monograph of Merodon

496, 372 (MNHN, RMNH, NHMW, BMNH, JLR); U. S. S. R.:
38,12 (NHMW, RMNH); Yugoslavia: 25,32 (NHMW).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin concave, apex subacute, antennal ratio
1.8; pubescence pale yellow, with few dark hairs in
ocellar region; face and occiput densely white to yel-
low pruinose; vertex angle 40°, ocellar angle 70°. to
85°, tl-v ratio 0.5.

Thorax: Dark, lateral dorsum and scutellum, kate-
pisternum, anepisternum and anepimeron weakly
blue lustrous; pubescence rather dense, erect, brown-
ish, dark interalar band present. Wings strongly in-
fuscate, except apical-posterior margin; halteres,
squamae and antisquamae yellow.

Legs: All dark, with yellow pubescence; trochanter
3 sharply ridged, femur 3 strongly swollen, triangular
processus normally shaped, tibiae 3 and tarsi 3 nor-
mal;

Abdomen: Rather slender; all dark; T II tapering,
bearing vague pale orange lateral spots, T I-IV with
widely interrupted arcuate whitish pruinose bands,
pubescence moderately dense, orange; S IV vaulted,
posteriorly deeply, widely emarginate.

Genitalia: Completely similar to those of clavipes.

Body length 17-23 mm.

Female. — Except for the rather marked sexual di-
morphism, differing from the male as follows. Head:
Frons with lustrous midstripe occupying one-third of
width, laterally white pruinose. Thorax: Incon-
spicuous longitudinal pruinose bands present, pubes-
cence brown to grey throughout; no interalar band;
wings less infuscate, strongest on surface bordering
veins. Legs: The trochanter 3 smooth, femur 3 slen-
derer than in & but still much swollen, pubescence
duller. Abdomen: Pruinose bands conspicuous,
stronger arcuate, less interrupted, rather variable
between specimens; pubescence almost restricted to T
II where dense; lateral spots clearer.

Body length 17-18 mm.

Diagnosis. — M. velox is easily distinguished by the
infuscate wings in the male; in the female it could be
confused with clavipes and pruni from which it is dis-
tinguished by the veins bordered with infuscation (in
clavipes wings clear; in pruni wings more evenly trou-
bled), from pruni also by the all dark legs, less slender,
much darker coloured abdomen and larger size.
Moreover, in pruni the genitalia are quite different.

Period of flight and distribution (fig. 90). — M. ve-
lox occurs in the eastern Mediterranean where the im-
agines are found from May through August; the spe-
cies may be bivoltine.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Biology. — The males of velox, observed by the au-
thor in May of 1984 in several Greek localities, were
flying close to the soil on rather dry sunny hillsides
with short vegetation.

Discussion. — Although the similarity of the geni-
talia with respect to clavipes is clear, the following ar-
guments against synonymizing velox can be listed: (1)
there are clear external differences; (2) the distribu-
tion of velox is quite restricted and purely
Mediterranean, whereas clavipes occurs over all of
southern and central Europe; (3) the flight period of
velox is more restricted; (4) there is some evidence
that velox, where it occurs together with clavipes, pre-
fers more open, drier biotopes (author’s observations;
J. Lucas, pers. comm.); therefore, the ecological nich-
es might be different; (5) behavioural differences
seem to exist. When the velox males were observed in
Greece, they did not show territorial behaviour.
Males of clavipes observed at the same locality and
date displayed territorial behaviour (cf. Hurkmans,
1988). This may indicate that either velox shows no
territorial behaviour, or this behaviour takes place in
a different period of the season.

Merodon velox vat. anathemus Paramonov

Material examined. — Greece: 12 (WH); Turkey: 19
(WH).

Diagnosis. — Completely similar to the nominate
form, except for the all dark pubescence. Only fe-
males are known.

Merodon velox var. armeniacus Paramonov

Material examined. — Turkey: 12 (wr)

Diagnosis. — Completely similar to the nominate
form except in the abdomen where lateral spots are
lacking and pruinose bands are reduced, in both sex-
es.

Discussion. — No recent male material is known; of
the syntypes, all but one were males. Possibly some of
the material might be present in the collection of the
Museum of Armenia, Erivan.

Merodon warnckei sp. n.

(figs. 52 a-d, 88)

Type-material. — d Holotype ‘Turkey, Hakkari,
5.vili.1983, Gavaruk-lake Mt. Sat, 2900 m, leg. Warncke’
[37° 20° N, 43° 35° E] (ZMAN). Paratypes: 48, 19; 1d
‘Turkey, Hakkari, Sat Mountains north of Mt. Gavaruk

184

2900 m, 7.viii.1983 leg. Warncke’ (LR); 2d, 19 “Turkey,
Hakkari, Sat Mountains, south of Sat Lake, 2800 m,
7.viii.1983, leg. Warncke’ (LR); 16 ‘Turkey, Van,
Mengene Dagi north of Baskale, 2700-3000 m,
27/28.vi.1986, leg. Hurkmans’ (wr).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apax acute, antennal ratio 2.0;
pubescence yellow throughout, pruinosity slight on
face, dense on occiput; face strongly blue lustrous, oc-
ciput greenish lustrous; vertex angle 25°, ocellar ang-
le 60°, tl-v ratio 0.3.

Thorax: Dark, greenish to blue lustrous on lateral
dorsum and scutellum, katepisternum, anepisternum
and anepimeron; slightly pruinose on sides; pubes-
cence rather even, moderately dense, dull yellow.
Wings slightly tinged; halteres, squamae and anti-
squamae yellow.

Legs: Dark except joints on legs 1-2, leg 3 all dark;
trochanters 3 smooth, femora 3 bluish lustrous, swol-
len, with normally shaped triangular processus bear-
ing 6-9 bristles, tarsi 3 normally shaped; pubescence
yellow.

Abdomen: All tergites dark with some micaceous
purple lustre; vague lateral pruinose spots present on
T II; T ILIV with arcuate widely interrupted moder-
ately wide yellowish pruinose bands; pubescence dull
yellow, moderately dense throughout; S IV shallowly
emarginate posteriorly, rather flat.

Genitalia: The anterior surstyle lobe very large,
rather stalked, bearing dense, even, yellow pubes-
cence apically, coalescent with posterior surstyle lobe;
the latter with anterior low elongate projection bear-
ing yellow, rather dense pubescence, posteriorly with
long, dense, yellow pubescence. Cercus rather large,
showing dense, very long, yellow pubescence.
Aedeagus moderately slender, slightly widening api-
cally, apical shaft part short, fringed plates on thecal
apex recumbent.

Body length 14-15 mm.

Female. — Except for the conspicuous sexual di-
morphism, differing from the male as follows. Head:
Lustrous midstripe on frons occupying one-third of
width; slight dark hair-tuft in ocellar region; face
weakly lustrous, apex of antenna 3 subacute; pubes-
cence grey. Thorax: Pruinose bands on dorsum more
distinct; pubescence greyish yellow. Legs: With grey
pubescence. Abdomen: Pruinosity more extensive on
T II, weaker on T III-IV; pubescence grey, less dense
than in male; lustre weaker; T V tapering, apex
rounded.

Body length 14 mm.

Diagnosis. — M. warnckei males can be distin-
guished from those of other species in this group by

its relatively stout build, the stalked anterior surstyle
lobe and the strong metallic lustre of the face; the fe-
males of warnckei are stouter and smaller, and have
less swollen femora 3 than clavipes or velox, are much
stouter than pruni or the species in e.g. the avidus and
alagoezicus groups from which they differ as well in
having much less conspicuous pruinose bands.

Period of flight and distribution (fig. 88). — The
species is known from eastern Turkey; it has been col-
lected in high altitudes only, where it occurs in June
and August; in view of the few specimens collected,
the split occurrence does not necessarily indicate that
warnckei is bivoltine.

Etymology. — The species has been named after the
German entomologist K. Warncke, who collected the
holotype and most paratypes. A noun in genitive case.

The pruni group
Apomorphies: The d genitalia with surstyle sulca-

te; vertex angle very small; yellow coloration on abdo-
men extensive.

Merodon pruni (Rossi)
(figs. 53 a-d, 94)

Syrphus pruni Rossi, 1790: 293. Syntypes: ‘in provinciis
Florentina et Pisana’ [Toscana, Italy] [not examined].
Merodon pruni var. obscurus Gil Collado, 1929: 407.
Holotype d: ‘Tanger, Mz. Escalera/ M. pruni var. ob-
scurus Gil Tipo, Gil Collado det./ M. N. C. N. Madrid’

(IEE) [examined].

Merodon fulvus Macquart, 1834: 514. Syntypes ‘France
Meridionale’ [not examined].

Merodon fuscinervis Von Réder, 1887: 73. Syntypes: ‘bei
Elos [Elous, 35° 20’ N, 23° 36° E] in der Nahe von
Kisamos auf der Insel Kreta’ [not examined].

Merodon pallidus Macquart, 1842: 70. Lectotype d (here
designated): ‘Baghdad’ [examined].

Merodon sicanus Rondani, 1845: 258, 264. Lectotype 9
(here designated): ‘58’ [number referring to description
of M. sicanus in catalogue by Rondani] [examined].

Merodon pruni, Schiner 1857: 415; Bezzi 1900: 90; Becker
1912: 602; Sack 1913a: 434; Paramonov 1925: 152;
Paramonov 1926a: 15; Hurkmans 1985: 69.

Merodon pruni var. obscurus, Peck 1988: 173.

Lampetia pruni, Sack 1931: 323; Séguy 1961: 181; Van der
Goot 1964a: 431.

Merodon pallidus, Brunetti 1923: 217.

Lampetia pallida; Sack 1931: 323.

Merodon sicanus, Rondani 1857: 55, 65; Schiner 1857: 411;
Schiner 1862: 347; Palma 1863: 46; Strobl 1893: 76;
Strobl 1900: 593.

Lampetia sicana; Sack 1931: 323.

Material examined. — Algeria: 6d, 22 (BMNH, MNHN);
Cyprus: 104, 102 (BMNH, NHMW, KBIN); France: 1d, 12
(MNHN); Greece: 236, 112 (MNHN, NHMW, BMNH, JLR,

Hurkmans: Monograph of Merodon

Den Hollander, WH); Iran: ® paralectotype of pallidus.
‘Baghdad’ (MNHN); 1d, 12 (RMNH); Italy: 9. paralectotype
of sicanus: ‘58’; 796, 322 (BMNH, NHMW, ZMAN, RMNH,
JLR, WH); Israel: 166, 119 (KBIN, NHMW, MNHN, NHMW,
BMNH); Lybia: 6d, 72 (BMNH); Morocco: 5d (MNHN,
RMNH); Pakistan: 116, 109 (BMNH); Syria: 1d (NHMW);
Turkey: 88, 142 (JLR, ZMUC, MNHN).

Description

Male. — Head: Antennae brown to orange, antenna
3 with upper margin convex, apex subacute, antennal
ratio 1.4; pubescence yellowish, darker in ocellar re-
gion; face and occiput rather densely pale yellow prui-
nose, weakly blue metallic lustrous; vertex angle 25°,
ocellar angle 35°, tl-v ratio 0.4.

Thorax: Dark; katepisternum, anepisternum and
anepimeron weakly blue lustrous; pubescence on dor-
sum and scutellum rather short, very even, yellow, in
some specimens with more or less clear interalar band
and/ or with more or less vague pruinose longitudinal
bands; pubescence on sides concolorous, less even.
Wings troubled, strongest in anterio-basal portion;
halteres, squamae and antisquamae pale yellow.

Legs: The femora dark with yellow apices, tibiae
yellow with broad dark distal band, tarsi 1 and 2 yel-
low, tarsi 3 dark with striking golden pubescence on
sides contrasting dark upper face; trochanters 3 bear-
ing acute projection, femora 3 strongly swollen, bear-
ing much projecting triangular processus holding 6-9
bristles on serrate distal margin; all tarsi normal.

Abdomen: Rather slender; T II strongly tapering,
with large yellow lateral spots; background colour
dark brown, but in most specimens replaced with yel-
low to a large extent, in some specimens yellow
throughout; pruinose bands pale yellow, arcuate, in-
complete on T II, wide and hardly interrupted on T
III-IV where they may occupy most of the tergite sur-
face; pubescence rather dense, predominantly yellow,
longest laterally; sternites yellow, S I-II pale, III-IV
darker, the latter strongly vaulted, deeply, narrowly
emarginate posteriorly.

Genitalia: Anterior surstyle lobe rounded, bearing
short, erect, yellowish pubescence throughout except
basally; separated from posterior surstyle lobe by shal-
low sulcus; posterior surstyle lobe bearing long, un-
even, erect, yellow pubescence. Cercus conspicuous,
wide, with long, dense, yellow pubescence, somewe-
hat angular. Aedeagus rather short, moderately slen-
der, apical shaft part short, fringed plates on thecal
apex recumbent.

Body length 8-23 mm.

Female. — Very similar to the male. Except for sex-
ual dimorphism, differing as follows. Head: Frons
with lustrous midstripe occupying less than 0.2 of
frontal width; ocellar angle 45°. Thorax: Narrower
than in male; Pruinose bands often vaguely present

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TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

on dorsum; wings stronger infuscate. Legs: Femora 3
less swollen, but strongly swollen compared with fe-
males of other species of Merodon; trochanters 3
smooth. Abdomen: Pruinose bands wider medially,
narrower laterally compared with males; T V taper-
ing, all dark.

Body length 10-21 mm.

Variation. — Merodon pruni shows an immense
variation in size, which does not seem to be correlat-
ed with locality or flight period. The specimens as-
signed to pruni can be arranged in a continuous se-
ries; the expression of several characters seems to be
correlated with specimen size, e.g. relative size of pro-
jection on trochanters 3 and triangular processus.
This phenomenon is also found in other species, most
strongly in kawamurae. The differences between spe-
cimens may be the consequence of differing food
availability in the larval stage.

Diagnosis. — The usual size in both sexes (normal
range 17-21 mm) and predominantly yellow abdo-
men combined will distinguish pruni from all other
species except clavipes and velox, from these two the
males are easily distinguished by the more slender ab-
domen with less dense pubescence, moreover from
velox by the absence of deep infuscation on wings; the
females of pruni are paler and have more slender ab-
domina than those of clavipes and velox. Small speci-
mens can be distinguished from other Merodon, in
the males by the nearly all yellow abdomen and the
genitalia; the dwarf females may cause some problems
but can be distinguished by their triangular yellow
abdomina and darkened wings. In pruni females of all
sizes the darkening of the wings is black, without pur-
ple reflections.

Period of flight and distribution (fig. 94). — M.
pruni has been recorded by Sack (1931: 324) to fly
from May to August in southern Europe, north
Africa and Asia Minor. The species is common in
Toscana, Italy and in central Greece. From the speci-
mens in collections it appears that pruni may be bi-
voltine at least in Italy and Greece. M. pruni seems to
be rare in southern France and absent from Spain.

Biology. — M. pruni occurs in rather open habitats
with scattered tall plants, mainly Umbelliferae and
Liliaceae, on hillsides exposed to the sun (Hurkmans
1985); the males are territorial and aggressively expel
invaders; the females are much less conspicuous in the
field than the males as they fly close to ground level
(Hurkmans 1985). This difference in flight behavi-

our might explain the sex ratio found among pre- —

served material in museum collections studied, which
is about 1.7 males to a female.

186

Discussion. — Type-material of fulvus Macquart,
1834 and fuscinervis Von Réder, 1887 could not be
examined. To judge from the descriptions both taxa
seem to fall well within the variation range of pruni.
The synonymy with pruni was already given by Sack
(1913a, 1931) and Peck (1988).

Merodon pruni var. obscurus Gil Collado

Diagnosis. — Rather similar to the nominal form,
except for the following differences in the male (fe-
males unknown). Head: Antennae 3 dark brown [not
black as indicated in description], truncate. Thorax:
Pubescence very short, even, yellow, but darker in
middle of dorsum, transition gradual; no pruinose
bands present. Wings darkened along anterior mar-
gin, the brown veins bordered with slight infuscation.

Body length 18 mm.

The longicornis group

Apomorphies: Extreme lengthening of antenna 3;
male genitalia with surstyle deeply sulcate and very
slender aedeagus; vertex angle very small.

Description. — Head: Antennae 3 with upper mar-
gin straight, apex subacute to rounded, antennal ratio
2.2 to 4.8; vertex angle 25° to 30° in males, ocellar
angle some 35° in males, 60° in females; in females
lustrous midstripe on frons occupying 0.35 to 0.6 of
width. Thorax: Dark, rather narrow, bearing 4 longi-
tudinal, mostly clear pruinoise bands. Wings clear.
Legs: All dark, sometimes joints paler, trochanters 3
smooth, femora 3, triangular processus and basitarsi 3
normally shaped.

Abdomen: Rather slender; T II somewhat tapering,
T III-IV cylindrical in male, tapering in female; in er-
ivanicus and longicornis dark background coloration
partially replaced by crimson red colour typical of this
group; pruinose bands narrow, arcuate and conspicu-
ous. Genitalia; Anterior surstyle lobe quadrate to rec-
tangular, anteriorly showing dense, yellow pubes-
cence, separated from posterior surstyle lobe by deep
sulcus, stalked; the posterior surstyle lobe low, short
to elongate. Cercus well protruding. Aedeagus very
slender, fringed plates on thecal apex recumbent to
oblique.

Body length in both sexes 9-13 mm.

Merodon erivanicus Paramonov

(figs. 54 a-b, 55 a, 92)

Merodon erivanicus Paramonov, 1925: 151.
‘Erivan, Armenien [not examined].

Merodon erivanicus, Liepa 1969: 20; Peck 1988: 170.

Syntypes:

Lampetia erivanica; Sack 1931: 315; Sack 1938: 22.

Material examined. — Greece: 29 (NHMW); Israel: 19
(BMNH); Turkey: 192 (BMNH, JLR, WH); Yugoslavia: 19
(NHMW) .

Description

Female. — Head: Antennae dark brown, antenna 3
with upper margin convex to straight, apex obtuse,
antennal ratio 2.2 to 2.8; ocellar angle 60°, pubescen-
ce yellow, face and occiput with slight metallic lustre
obliterated by dense whitish pruinosity; frons densely
pruinose laterally, midstripe occupying 0.4 of width.

Thorax: Dark, dorsum with well marked pruinose
bands, pubescence greyish yellow, short, dense, even;
katepisternum, anepisternum and anepimeron green-
ish to bluish metallic lustrous, hardly pruinose, pu-
bescence concolorous. Wings clear; halteres, squamae
and antisquamae pale yellow.

Legs: Dark, tibiae 1 and 2 basally yellow; dense yel-
low pubescence on sides of tarsi, contrasting dark pu-
bescence on upper faces; trochanters 3 smooth, femo-
ra 3 slightly swollen, triangular processus normal.

Abdomen: Slender, dark coloration partially re-
placed by crimson; the extent of the crimson colour is
much variable between specimens; T II-IV with inter-
rupted arcuate white pruinose bands; pubescence very
short, inconspicuous, black on dark surface, yellowish
on crimson surface; S I-II red, III-IV dark red to
black.

Body length 10-12 mm.

Male. — Unknown.

Diagnosis. — M. erivanicus is distinguished from fe-
males of other Merodon species by the long antenna 3
and crimson colour on the abdomen; from longicornis
by the shorter antenna 3 (antennal ratio in longicornis

is 4.1 to 4.8).

Period of flight and distribution (fig. 92). — M. er-
ivanicus occurs in June and July, in the eastern
Mediterranean, in eastern Turkey and Armenia.

Merodon kaloceros sp. n.
(figs. 54 d, 55 c, 56 a-c, 87)

Type-material. — d Holotype “Turkey, Antalya, H.
Coene, J. Lucas & B. van Oorschot / Irmesan Gedigi 12 km
N of Akseki 1600 m 24.vii.1981’ (ZMAN). Paratypes: 24d,
19:19, same data as holotype (LR); 48 “Turkey, Hakkari,
Suvarihalil pass 1250 m, W side near Habul Deresi
13.vi.1984 leg. J. A. W. Lucas’ (LR); 2d “Turkey Hakkari,
Chilo Daglari N of Oramar 1400 m, 16/17.vi.1984, leg. J.
A. W. Lucas’ (LR); 28 “Turkey, Hakkari, Sat Dag, SW
Yüksekova 1600 m 18.vi.1984 J. A. W. Lucas’ (LR); 8d
‘Turkey pr. Hakkari, Sat Dag, Varegös SW Yiiksekova 28.
and 29. vi.1985’ (LR); 18 ‘Turkey, Van, 40 km SW

Hurkmans: Monograph of Merodon

Akdamar 1720 m, 17.vii.1986, leg. P. van Ooijen’ (ZMAN);
16 ‘Dalmatien [Yugoslavia], Gravosa vi.28 [1928] Zerny
(NHMW); 1d ‘Greece, Ithaca 6/11.vi.1965 F. J. Francois’
(KBIN); 24 ‘Hill Scrub 500 ft above lake/ Macedonia Prespa
geul Otesevo 20/27.vi.1958/ Yugoslavia R. L. Coe’ (BMNH);
1d ‘Yugoslavia Kroatie Bribir (50 100) 10.viii.1964 H. J. P.
Lambeck’ (JLR); 16 ‘Corfu, Erber 218’ [Greece] (JLR).

Description

Male. — Head: Antennae dark, antenna 3 brown
with base and apex paler, upper margin straight to
convex, apex subacute to obtuse, antennal ratio 2.3 to
2.7; pubescence greyish yellow with dark tuft in ocel-
lar region ; vertex angle 25-30°, ocellar angle 40°, tl-
v ratio 0.55.

Thorax: Dark, dorsum with inconspicuous prui-
nose bands, katepisternum, anepisternum and ane-
pimeron blue to green metallic lustrous; pubescence
yellowish, rather even and short on dorsum and scu-
tellum, longer, uneven on sides. Wings clear; halteres,
squamae and antisquamae yellow.

Legs: Dark, tarsi with contrasting golden pubes-
cence on sides and dark upper faces; trochanters 3
smooth, femora 3 slightly swollen, triangular proces-
sus normally shaped, distal margin much serrate; tib-
iae 3 and tarsi 3 normal.

Abdomen: Dark, slender, with conspicuous orange
lateral spots on T II; T II-IV showing white conspic-
uous narrowly interrupted arcuate pruinose bands;
pubescence concolorous with surface, yellow on later-
al spots; S brown, S IV darkest, flat, posterior margin
entire.

Genitalia: Anterior surstyle lobe large, more or less
rectangular, rounded anteriorly, on margin with
dense, yellow, even pubescence; the posterior surstyle
lobe separated by deep sulcus, rather short, low,
somewhat angular, showing dense, uneven, yellow
pubescence. Cercus rather short, wide, with dense
yellow pubescence. Aedeagus bearing paired humps
basally on outer face, apical shaft part short, fringed
plates on thecal apex recumbent to oblique.

Body length 13 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons with lustrous
midstripe occupying 0.3 of width. Thorax: Pruinose
bands more conspicuous; pubescence denser, grey,
even as if shaven. Legs: Pubescence greyish through-
out. Abdomen: Outline more triangular; pubescence
denser, greyish; pruinose bands wider, continuous;
lateral spots extended over all of T II and anterior
margin of T III; T V tapering, dark, with grey pubes-
cence.

Body length 13 mm.

Diagnosis. — M. kaloceros can be distinguished
from other Merodon except erivanicus and longicornis
by the large antennal ratio and structure of the male

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

genitalia; it is distinguished from the two species
mentioned by the coloration on the abdomen which
is orange in kaloceros but crimson in erivanicus and
longicornis.

Period of flight and distribution (fig. 87). — M.
kaloceros occurs in the eastern Mediterranean region
and Turkey in June through August. Given the col-
lection dates from eastern Turkey, where most of the
type material was collected, the species is univoltine.

Discussion. — In view of apomorphies shared with
longicornis, e.g. the shape of the d genitalia and large
antennal ratio, kaloceros is assigned to the longicornis

group.

Etymology. — The epitheton kaloceros is derived
from the greek words kalon, meaning good, beautiful,
and ceros, meaning horn; it refers to the large anten-
nae (not as extremely long as in longicornis). The
epitheton is to be treated as a noun in apposition.

Merodon longicornis Sack
(figs. 54 c, 55 b, 57 a-c, 93)

Merodon longicornis Sack, 1913a: 447. Lectotype © (here
designated): ‘longicornis det. Sack/ Merodon longicornis
Sack Type Turkestan/ longicornis det. Hermann’
(NHMW) [examined].

Merodon longicornis, Paramonov 1925: 154, 1926a: 15; Van
der Goot 1981: 214, 218; Peck 1988: 171.

Lampetia longicornis, Sack 1931: 320.

Material examined. — Greece: 26, 259 (KBIN, JLR, Coll.
Thessaloniki, ZMAN, BMNH, NHMW); Italy: 306, 249
(RMNH, JLR, ZMAN); Israel: 12 (RMNH); Morocco: 19
(BMNH, doubtful); Syria: 12 (KBIN); Turkey: 16d, 249
(BMNH, JLR, NHMW); Yugoslavia: 42 (BMNH, RMNH).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin rather straight, apex rounded, antennal ra-
tio 4.1 to 4.8; pubescence pale yellow, dark in large
ocellar region; ocelli placed far forward; pruinosity on
face and occiput dense, whitish; vertex angle 30°,
ocellar angle 35°; tl-v ratio 0.3.

Thorax: Dark; dorsum with pruinose bands, pu-
bescence rather sparse, even, moderately long; surface
of katepisternum, anepisternum and anepimeron
weakly lustrous, with some silky lustre produced by
yellow recumbent pubescence. Wings clear; halteres,
squamae and antisquamae pale yellow.

Legs: Dark, joints paler in specimens; trochanters 3
smooth, femora 3 rather strongly swollen, straight;
triangular processus and tarsi normally shaped.

Abdomen: Dark; moderately slender; T II taper-
ing; T II-IV showing conspicuous slightly arcuate
white narrow pruinose bands; pubescence concolor-

188

ous with surface; S IV moderately vaulted, posterior-
ly narrowly emarginate.

Genitalia: The anterior surstyle lobe rather rectan-
gular, with short, even, yellow pubescence, separated
from the rather long, low posterior surstyle lobe by a
deep sulcus; the posterior surstyle lobe posteriorly
bearing moderately long, yellow, uneven pubescence,
its lateral face with several ridges and furrows. Cercus
rather large, well protruding, showing long, dense,
slightly wavy, yellow pubescence. Aedeagus normally
wide in ventral view, extremely slender in lateral view,
apical shaft part short, fringed plates on thecal apex
oblique.

Body length 9-10 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Lustrous midstripe
on frons occupying 0.6 of width; pubescence dark on
midstripe, grey laterally. Thorax: Pubescence shorter
than in male. Legs: Yellow pubescence on basitarsi 3
found in some 2. Abdomen: Strong crimson colora-
tion always present in @, restricted to T II or extend-
ed over most of abdomen; T II more tapering than in
male; T V acute, dark, scattered pale pubescence
present.

Body length 9-10 mm.

Diagnosis. — The extremely large antennal ratio
distinguishes /ongicornis from all other known species.

Period of flight and distribution (fig. 93). — The
species occurs in May through July in the
Mediterranean region, Turkey, Ukraine and southern
Russia. It appears to be rare in the western, common
in the eastern part of the range and in Turkey and
adjacent areas.

The vandergooti group

Apomorphies: Lengthening of anterior surstyle lo-
be; extreme swelling of femora 3; extensive yellow co-
loration on legs.

Merodon vandergooti sp. n.
(figs. 58 a-c, 88)

Type-material: 4 Holotype “Turkey, Hakkari,
Suvarihalil pass 1250 m W side near Habul Deresı,
13.vi.1984 leg. J. A. W. Lucas’ (ZMAN). Paratypes: 20d;
183 with same data as holotype (16 JLR, 2 WH); 1d
[Turkey] ‘Ankara 650 m, 1.vi.1971 leg. KI. Warncke’ (JLR);
16 [Turkey] ‘Ankara, South of Ankara, 24.vi.1984 leg. J. A.
W. Lucas (LR).

Description

Male. — Head: Antennae conspicuously orange, an-
tenna 3 with convex upper margin, apex acute, an-

tennal ratio 1.5 to 1.7; pubescence golden yellow, rat-
her dense, producing silky lustre on frons; face blue
metallic lustrous, moderately dense white pruinose;
vertex angle 40°, ocellar angle 40°, tl-v ratio 0.4.

Thorax: Dark, with vestigial pruinose bands on
dark dorsum; katepisternum, anepisternum and ane-
pimeron weakly blue lustrous; pubescence golden on
dorsum and scutellum, greyish-golden on sides; in
many specimens some dark hairs present between
wing insertions giving suggestion of vague intaralar
band; posterior fringe of scutellum bearing dense yel-
low pubescence. Wings clear; halteres, squamae and
antisquamae yellow.

Legs: Bases of femora 1 and 2 dark, femora 3 all
dark, otherwise legs fiery orange throughout, with
orange pubescence; trochanters 3 showing distinct
ridge ending laterally in a low but marked hump; fe-
mora 3 enormously swollen, rather strongly curved;
triangular processus large, bearing 10-13 delicate bris-
tles on hardly serrate distal margin.

Abdomen: Dark; T II relatively large, tapering,
bearing vague yellow lateral spots; T II-IV with
oblique slightly arcuate, yellow pruinose bands, pu-
bescence concolorous with surface; S IV moderately
vaulted, narrowly and deeply emarginate posteriorly.

Genitalia: Anterior surstyle lobe moderately wide,
much elongate, the anterior part showing dense yel-
low pubescence throughout, ventral margin similarly
pubescent; separated from posterior surstyle lobe by
deep wide sulcus; posterior surstyle lobe low, elon-
gate, bearing dense, long, yellow pubescence on ven-
tral and posterior margins; there is a secondary lateral
lobe, bearing moderately long yellow pubescence.
Cercus rounded, well protruding, with dense yellow
pubescence. Aedeagus moderately slender with short
apical shaft part and fringed plates on thecal apex re-
cumbent.

Body length 13-14 mm.
Female. — Unknown

Diagnosis. — M. vandergooti is easily separated from
other Merodon species by the structure of the genital-
ia and the enormously swollen femora 3; the colour
pattern of the legs is very unusual as nearly all species
with partially yellow legs have dark bands on the tib-
iae. The colour pattern on the legs and orange anten-
nae are also found in aureotibia females. In vandergoo-
ti the antenna 3 is acute, in aureotibia obtuse; the
abdominal pruinose bands are much more arcuate in
aureotibia, where the triangular processus has a
strongly serrate distal margin with 6-7 robust bristles;
in vandergooti the triangular processus bears 10-13
fine bristles on a hardly serrate distal margin. These
differences cannot be accounted for by explaining
them as sexual dimorphism and are thus considered

Hurkmans: Monograph of Merodon

valid distinctions on the species level.

Period of flight and distribution (fig. 88). — M.
vandergooti occurs in June and has been found in cen-
tral and south-eastern Turkey.

Etymology. — It is a pleasure to dedicate this species
to Mr Volkert van der Goot who, over the last
decades, has largely stimulated research work on
Syrphidae in the Netherlands. A noun in genitive
case.

The nigritarsis group

Apomorphies: Aedeagus with apical shaft part
lengthened; two pairs of spines on outer face of aede-
agus.

Description. — Head: Antennae dark, antennal ra-
tio over 1.5, tl-v ratio about 0.4 in male; face bluish
lustrous, strongest in male; in female, frons lustrous
on median 0.25 of width.Thorax: Inconspicuous
pruinose bands present; pubescence rather dense,
short. Legs: Dark with yellow markings on tibiae, tar-
si 1 and 2 in normal (not melanistic) specimens; tro-
chanters 3 much variable between specimens, usually
humped; golden pubescence on sides of tarsi 3 con-
trasting dark upper face. Abdomen: Moderately slen-
der; T IV bulging in male; pruinose bands arcuate,
clear; S IV in male strongly vaulted, deeply emargi-
nate posteriorly. Genitalia: Anterior surstyle lobe el-
ongate, stalked, diamond-shaped, posterior surstyle
lobe elongate; aedeagus with 2 pairs of chitinous
spines on outer face and lengthened apical shaft part.

Merodon femoratoides stat. nov.

(figs. 59 a-d, 95)

Merodon spinipes var. femoratoides Paramonov, 1925: 158.
Lectotype d (here designated): ‘Karadagh bei Theodosia,
Krym, 3.vi.1919° (ZMHB) [examined].

Merodon spinipes var. femoratoides, Liepa 1969: 20; Peck
1988: 174. i

Lampetia spinipes var. femoratoides, Sack 1931: 328.

Material examined. — Algeria: 16 (BMNH); Greece: 2d,
12 (LR); Turkey: 126 (LR, BMNH, NHMW, WH); U. S. S.
R.: 2d Paralectotypes of spinipes var. femoratoides,
‘Warnutka, Sevastopol, Krym 11.vi.1923’ (ZMHB); 1d,
Krym, Karadagh 24.vi.25 [1925] Paramonov leg./ Typus
Merodon spinipes var. femoratoides 4 / B. Kurodke Pinob
[in Russian script] (ZMHB) [This specimen cannot be consid-
ered as a type specimen since it is not mentioned in the orig-
inal publication].

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin concex, apex subacute, antennal ratio 1.5;
pubescence yellow, face strongly white pruinose; ver-

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

tex angle 30°, ocellar angle 45°, tl-v ratio 0.4.

Thorax: Dark, with four clear pruinose bands and
a fifth anteromedian fainter one; pubescence dark yel-
low, rather short, even; katepisternum, anepisternum
and anepimeron blue lustrous. Wings slightly evenly
troubled; halteres, squamae and antisquamae yellow.

Legs: The tibiae basally and femora apically orange,
otherwise dark; trochanters 3 smooth, femora 3 mod-
erately swollen, triangular processus bearing 6-10
strong bristles on moderately serrate distal margin;
tibiae 3 bearing dense golden pubescence, tarsi 3 with
same pubescence on sides, contrasting dark upper
face.

Abdomen: Moderately slender; T II much taper-
ing; T II-IV with arcuate off-white pruinose bands; T
II bearing clear to almost absent yellow lateral spots
which may extend over part of T III; pubescence con-
colorous with background; S IV vaulted, acutely, nar-
rowly emarginate posteriorly up to half its length.

Genitalia: Anterior surstyle lobe elongate, clearly
stalked, the apical portion triangular, anteriorly
showing dense yellow pubescence; posterior surstyle
lobe with small anterior, accessory lobe bearing dense,
long, yellow pubescence; itself low and elongate,
showing dense yellow pubescence posteriorly. Cercus
inconspicuous and hidden between projecting poste-
rior surstyle lobes. Aedeagus rather slender, bearing
two pairs of spines on outer face, apical shaft part
lengthened, fringed plates on thecal apex oblique.

Body length 13-15 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons lustrous on
median one-third, whitish pruinose laterally. Thorax:
Pubescence as if shaven, golden; four well marked
pruinose bands present. Abdomen: T II tapering, lat-
eral spots conspicuous, extended over part of T III;
pruinose bands arcuate, widely interrupted and con-
spicuous.

Body length 12.5 mm.

Diagnosis. — M. femoratoides may be easily distin-
guished from avidus by the very different genitalia in
the & and the contrast of hair colour on tarsi 3 in
both sexes. The distinction with nigritarsis is in the
differently shaped anterior surstyle lobe in the males
and the less dark abdomen with larger lateral spots in
the females.

Period of flight and distribution (fig. 95). — The
species occurs in the Mediterranean, Asia Minor and
southern Russia and Ukraine from May through July;
its distribution range is large, but the species is un-
common.

190

Merodon nigritarsis Rondani

(figs. 60 a-g, 97)

Merodon nigritarsis Rondani, 1845: 258, 264. Lectotype d
(here designated): ‘68/ Torino D. Ghiliani’ in coll.
Rondani (LSF) [the number 68 refers to the description of
this species in Rondani’s catalogue] [examined].

Merodon nigritarsis, Rondani 1857: 55, 65; Schiner 1857:
415; Rondani 1873: 295; Strobl 1893: 76; Bezzi 1895:
16; Strobl 1900: 91; Sack 1913a: 447; Paramonov 1925:
157, 1926a: 15; Gil Collado 1930: 254; Peck 1988: 174.

Lampetia nigritarsis Sack 1931: 327; Séguy 1961: 177.

Material examined. — Austria: 46 (BMNH, NHMW,
ZMUC); France: 966, 322 (MNHN, KBIN, RMNH, BMNH);
Greece: 296, 192 (NHMW, ZMUC, den Hollander, KBIN,
Bonn, ZMAN, RMNH, Thessaloniki, BMNH); Hungary: 2d,
42 (RMNH, Bonn, ZMUC); Italy: 38, 39 paralectotypes,
‘68’ in coll. Rondani (LSF); one 9 paralectotype, ‘68/ Susa,
D. Ghiliani’ in coll. Rondani (LsF) and 136, 5® (LSF, KBIN,
ZMAN, RMNH); Poland: 1 2 (BMNH); Spain: 98,52 (MNHN,
BMNH, JLR, NHMW); Turkey: 96, 62 (LR, BMNH, NHMW,
ZMAN, RMNH); Yugoslavia: 496, 199 (BMNH, NHMW,
Bonn, ZMAN, RMNH).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apex subacute, antennal ratio 1.6;
pubescence yellow with dark tuft in ocellar region;
vertex angle 30°, ocellar angle 40°, tl-v ratio 0.4.

Thorax: Dark, lustrous on lateral dorsum, scutel-
lum and sides, bearing rather dense yellow pubes-
cence. Wings clear; halteres, squamae and antisqua-
mae yellow.

Legs: All femora dark, paler apically; tibiae yellow
with broad dark distal band; trochanters 3 with much
variable hump, usually well marked, femora 3 slight-
ly swollen, with normal triangular processus bearing
6-9 bristles on serrate distal margin; tibiae 3 and tarsi
3 normally shaped; the basitarsi 3 with strong con-
trast between dark pubescence on upper face and
golden pubescence on sides; this is a characteristic
that can be of use in the field as it is conspicuous to
the naked eye at short range.

Abdomen: Dark; T II with orange to yellow lateral
spots of variable size and clearness; T II] in many spec-
imens with yellow anterior lunules; T II-IV bearing ar-
cuate, narrowly interrupted pruinose bands; S IV very
deeply, narrowly emarginate, strongly vaulted.

Genitalia: Anterior surstyle lobe elongate, widened
subapically, more or less rounded apically, with dense
yellow even short pubescence there; posterior surstyle
lobe separated by small sulcus; rather elongate, mod-
erately high, posteriorly bearing dense, long, yellow
pubescence. Cercus rather inconspicuous, rounded,
pubescence as on posterior surstyle lobe. Aedeagus
bearing two pairs of spines on outer face; apical shaft
part clearly lengthened; fringed plates on thecal apex
recumbent.

Body length 12-16 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons lustrous on
median quarter, sides white pruinose, pubescence
concolorous; dark tuft in ocellar region less clear than
in male; ocellar angle 45 to 50°. Thorax: Shorter pu-
bescence; pruinose bands clearer. Legs: Trochanters 3
always smooth.

Body length: 12-17 mm.

Variation. — Merodon nigritarsis is one of the more
variable species of the genus; the hump on the tro-
chanters 3 and the shape of the anterior surstyle lobe
are notoriously variable between specimens, even
those collected together at the same site. There are
many local forms, each with their own peculiar aber-
rations, notably in the shape of the anterior surstyle
lobe. Since a fluent, continuous series can be arranged
using these specimens, I have refrained from describ-
ing local subspecies.

Diagnosis. — M. nigritarsis can be distinguished
from other Merodon except femoratoides in the males
by the general morphology of the genitalia and the
spines on the aedeagus and the contrast on the basi-
tarsi 3 in both sexes; from femoratoides it may be dis-
tinguished in the males by the differently shaped an-
terior surstyle lobe and basally convex aedeagus,
smaller lateral spots and less wide interrupted prui-
nose bands; in the females by the darker abdomen.

Period of flight and distribution (fig. 97). — M. nig-
ritarsis occurs in the Mediterranean and the southern
parts of central Europe from May through August;
possibly the species is bivoltine.

Discussion. — M. nigritarsis has been considered a
variety or subspecies of avidus. In view of the com-
pletely different genitalia a close relation between av-
idus and nigritarsis is out of the question. M. nigritar-
sis was named for the dark basitarsi 3; however, this
feature also occurs in other species externally resem-
bling nigritarsis, such as alagoezicus and its allies; the
lengthened apical shaft part of the aedeagus is another
shared feature.

The avidus group

Apomorphies: Anterior surstyle lobe anteriorly
much widened; T II extremely tapering.

Merodon avidus (Rossi)
(figs. 61 a-g, 96)

Syrphus avidus Rossi, 1790: 292. Syntypes: ‘in provinciis

Hurkmans: Monograph of Merodon

Florentina et Pisana’ [Toscana, Italy] [not examined].

Syrphus spinipes Fabricius, 1794: 296. Lectotype d (here
designated): ‘spinipes in coll. J. C. Fabricius (ZMUC) [seri-
ously damaged] [examined]

Merodon serrulatus Wiedemann in Meigen, 1822: 360.
Lectotype d : (here designated) ‘Portugall coll. Winthem/
serrulatus Wied.’ (NHMW) [examined]

Merodon italicus Rondani, 1845: 257, 259. Lectotype 9
(here designated): ‘67b’ in coll. Rondani (LSF); the ‘67b’
number refers to the description of ttalicus in Rondani’s
catalogue. [examined] syn. n.

Merodon rufitibius Rondani, 1845: 259, 265. Lectotype d
(here designated): ‘72’ in coll. Rondani (LSF) [examined];
the ‘72’ number refers to the description of rufitibius in
Rondani’s catalogue.

Merodon graecus Walker, 1852: 243. Lectotype ? (here des-
ignated): ‘Albania/ graecus Walker’ (BMNH) [examined].

Merodon aurifer Loew, 1862a: 83. Lectotype d (here desig-
nated): ‘Melatia [Malatya, Turkey] Erber/ coll. H. Loew/
Zool. Mus. Berlin’; on separate pin: ‘... Merodon aurifer
n. Berl. Zeit. vi.1862 pg 82 ...’ (ZMHB) [examined].

Merodon quadrilineatus Lioy, 1864: 473. [Type-material
probably never existed. Lioy published only theoretical
works and had no intention to describe new species.]

Merodon avidus, Rondani 1845: 259; Schiner 1857: 415;
Strobl 1893: 76; Bezzi 1895: 16, 1900: 91; Strobl 1900:
593; Villeneuve 1903: 115; Gil Collado 1930: 254; Van
der Goot 1971: 106, 1981: 216, 218; Barendregt 1982:
80; Marcos Garcia 1985: 196; Bradescu 1986: 123; Peck
1988: 167; Hurkmans 1988: 107.

Lampetia avida; Séguy 1961: 176; Van der Goot 1964a:
431.

Milesia spinipes, Latreille 1804: 331.

Merodon spinipes, Fabricius 1805: 197; Meigen 1822: 361;
Macquart 1828: 295, 1834: 516, 1842: 70; Rondani
1845: 258, 264, 1857: 55, 66; Schiner 1857: 414; Strobl
1893: 76; Bezzi 1895: 16, 1900: 91; Strobl 1900: 593;
Villeneuve 1903: 115; Czerny & Strobl 1909: 204; Sack
1913a: 444; Paramonov 1925: 146, 1926a: 15; Gil
Collado 1929: 409; Gil Collado 1930: 243; Bankowska
1980: 38, 40, 83; Marcos Garcia 1985: 137; Peck 1988:
174.

Lampetia spinipes, Oldenberg 1919: 388; Sack 1930: 96,
1931: 327, 1938: 21; Suster 1959: 243; Séguy 1961: 176.

Merodon serrulatus; Schiner 1857: 413, 1862: 348; Strobl
1893: 76; Bezzi 1895: 16; Strobl 1900: 593; Czerny &
Strobl 1909: 203, 204; Gil Collado 1930: 254.

Lampetia serrulata; Sack 1931: 322.

Merodon italicus, Rondani 1857: 54, 62; Schiner 1857: 413,
1862, 347.

Lampetia italica; Sack 1931: 322.

Merodon rufitibius, Rondani 1857: 55, 66; Palma 1863: 46;
Becker 1907: 253.

Lampetia rufitibia; Sack 1931: 327.

Merodon graecus, Schiner 1857; Schiner 1862: 347; Van der
Goot 1964b: 218; Peck 1988: 167.

Merodon aurifer, Peck 1988: 174.

Lampetia aurifera; Sack 1931: 327.

Merodon quadrilineatus, Peck 1988: 174.

Lampetia quadrilineata; Sack 1931: 327.

Material examined. — Albania: 1d, 19 (NHMw); Algeria:
26, 242 (MNHN, KBIN); Austria: 534, 632 (NHMW,
MNHN, BMNH, RMNH); Belgium: 3d (MNHN, KBIN); Cyprus:
136, 49 (KBIN, BMNH); Czechoslovakia: 16, 19 (BMNH,

191

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

JLR); France: 2666, 2109 (MNHN, BMNH, NHMW, RMNH,
JLR, ZMAN, WH); Germany: 36 (MENS, JLR); Greece: 966,
622 (BMNH, RMNH, NHMW, KBIN, MNHN, JLR, den
Hollander, ZMUC, Bonn, Coll. Thessaloniki, WH); Hungary:
293, 172 (Bonn, ZMUC, NHMW, BTM); Italy: 29
Paralectotypes of ztalicus ‘67b’ in coll. Rondani (LSF); 2d,
7% Paralectotypes of rufitibius ‘72’ in coll. Rondani (LSE);
3196, 1229 (ZMUC, KBIN, NHMW, JLR, RMNH, ZMAN,
BMNH, LSF, Coll. Marogna (Verona, Italy), WH); Lebanon:
19 (KBIN); Lybia: 48, 22 (BMNH); Morocco: 56, 89
(BMNH, MNHN, JLR, NHMW); Netherlands: 6d, 22 (RMNH,
JLR); Poland: 1d (ZMUC); Romania: 26, 12 (BMNH);
Spain: 708, 209 (ZMUC, ZMAN, RMNH, JLR, WH);
Switzerland: 56, 62 (RMNH, ZMAN, JLR); Turkey: 576,
302 (LR, WH, BMNH,NHMW); U. S. S. R.: 4d, 82 (MNHN);
Yugoslavia: 706, 409 (NHMW, BMNH, RMNH, JLR, MNHN,
Bonn, ZMAN).

Description

Male. — Head: Antennae with antenna 3 showing
subacute apex and convex upper margin, antennal ra-
tio 1.5, pubescence yellow, dark hair-tuft often pre-
sent in ocellar region; face rather densely white prui-
nose, partially obliterating blue lustrous surface;
occiput concolorous; vertex angle 35°, ocellar angle
40°-45°, tl-v ratio 0.5.

Thorax: Moderately wide, dark, rather dulled by
diffuse yellowish pruinosity, dorsum bearing longitu-
dinal pruinose bands of most variable clearness, kate-
pisternum, anepisternum and anepimeron very weak-
ly to conspicuously blue lustrous; pubescence erect,
yellowish, moderately dense. Wings clear, veins pale
to dark brown; halteres, squamae and antisquamae
yellow.

Legs: Dark on trochanters and femora, pale on tib-
iae and tarsi, but both ends of femora paler and dark
band present on tibiae is the usual pattern; there is an
immense variation resulting in relatively pale or all
dark legs; the femora always mainly dark; trochanters
3 may be acuminate; femora 3 moderately swollen,
bearing 6-9 bristles on serrate distal margin of trian-
gular processus.

Abdomen: Slender; dark; T II much tapering, usu-
ally with clear yellow lateral spots; pruinose bands on
T II-IV clear, arcuate, interrupted, of variable width
and colour; pubescence often concolorous with sur-
face, erect, longest anterolaterally; S IV strongly
vaulted, deeply acutely emarginate posteriorly.

Genitalia: Anterior surstyle lobe stalked, widening
to semicircular apical margin with short even yellow
pubescence; posterior surstyle lobe moderately high,
slightly elongate, with moderately dense, long, yellow
pubescence. Cercus rather large, bearing dense, long,
yellow, often wavy pubescence. Aedeagus moderately
slender, apical shaft part short, pipes plates on the-
cal apex recumbent.

Body length 11.5-18 mm.

192

Female. — Except for sexual dimorphism, differing
ferom the males as follows. Head: Lustrous midstripe
on frons occupying 1/4 to 3/4 of width, the lateral
pruinose strips usually well defined; face and occiput
less lustrous. Thorax: Pruinose bands clearer, slightly
wider. Abdomen: Lateral spots often reddish, con-
spicuous and extending over part of T III; in some
cases however virtually absent; pruinose bands less
clear, less wide. T V dark.

Body length 12-17 mm.

Variation. — There is much variation in the colora-
tion in both sexes. Head: The antennae may range
from black to bright yellow. Thorax: Lustre most var-
iable, density and colour of pubescence more con-
stant. Legs: The colour is extremely variable; all
joints, the tarsi and both ends of the tibiae are paler
than the rest of the legs, but the colour may be all
black or all yellow throughout. The pubescence is
whitish to yellow even in dark specimens. Abdomen:
The extent of the yellow colour is variable; the fe-
males often show a deeper colour than the males, and
are less variable in this respect. Genitalia: The male
genitalia are rather constant. The main variation oc-
curs in the length and density of the pubescence on
the posterior surstyle lobe and the cercus.

The large degree of variation found in avidus is not
randomly distributed over the specimens. Many series
collected at one site on one date, show uniform char-
acteristics. Size and colour are probably correlated
with quantity and quality of food ingested during the
larval stages. There is no clear trend in size, morphol-
ogy or colour of the various local forms over the en-
tire range. Therefore, to present descriptions of local
forms, varieties or subspecies is considered inappro-
priate.

Diagnosis. — M. avidus males could possibly be
confused with toscanus from which they are distin-
guished by the longer stalk of the anterior surstyle
lobe (which in zoscanus is bifid), the shorter apical
shaft part of the aedeagus and the usually much dark-
er abdomen. The unique genitalia are sufficient to
distinguish avidus males from any other Merodon
males at first sight. The females are posing many
problems since they cannot always be separated from
females of the species in the alagoezicus group. The
best separating characteristic is the pruinosity on the
thorax where avidus females always show four longi-
tudinal bands, whereas in the females of the alagoezi-
cus group there usually is a fifth, anteromedian band
as well. Moreover, the abdomen of females in the a/-

agoezicus group is usually narrower overall.

Period of flight and distribution (fig. 96). — M. av-
idus occurs in Europe, except the northern parts, in

north Africa, the Middle East and Asia Minor from

March through October; the species is very likely
multivoltine in the southern parts of its range (cf.
Marcos Garcia, 1985). The distribution does not ap-
pear to be associated with human activity. As avidus is
not particularly easy to collect and seems to occur in
rather the same habitats as other Merodon the relative
abundance in collections probably reflects abundance

in the field.

Biology. — Few details are known (if the great dis-
tribution and abundance is borne in mind). This sec-
ond most common Merodon occurs both in generally
wet, and in dry areas, such as the Vosges mountains of
France and the inland steppe region of Turkey. There
are indications that in the driest parts of its range av-
idus prefers relatively wet biotopes such as lush strips
along streams or marshes. Moreover imagines occur
early in the season in the hot, dry parts of its range,
probably to avoid water shortage later on. Water
shortage influences the vegetation and therefore the
possibilities for the larvae which are probably phy-
tophagous (all known Merodon larvae are phytopha-
gous). There seems to be a preference for flowers of
Umbelliferae, as recorded by Suster (1959) and con-
firmed by Mr. Lucas’ s (Pers. Comm.) and the
author’s observations in e.g. France, Italy, Greece and
Turkey. Some notes on the behaviour of the species
are provided by Hurkmans (1988).

Discussion. — No type-material of avidus has been
found; it might be supposed to be in the Rossi collec-
tion at ZMHB (Horn & Kahle 1936), but is not cur-
rently present there (dr. H. Schumann, in litt.). The
types of spinipes are mentioned purely formally as
they are too much decayed to be of any value what-
soever. There are considerable difficulties in distin-
guishing avidus from externally similar species, if the
genitalia are not used for identification. The existence
of the type-specimens in their present state does re-
grettably forestall designation of a neotype, which
would seem to be necessary, given the identification
problems mentioned. The original description by
Rossi (1790) and following emendations are detailed
enough to define avidus with respect to the external
characteristics. No examination of the genitalia had
been made until comparatively recently; especially in
avidus the use of genital characters for separation
from externally resembling species is essential. It is
however considered fair to rule that as the over-
whelming majority of museum specimens identified
as avidus examined fulfil the requirements of the orig-
inal description these specimens must be regarded as
the true avidus. Moreover most other species external-
ly resembling avidus can be (and have been) separated
by external characteristics as well, e.g. nigritarsis
Rondani and alagoezicus Paramonov.

Hurkmans: Monograph of Merodon

The crassifemoris group

Apomorphies: Posterior surstyle lobe very high,

with large anteriorly projecting lobe; cercus reduced.

Merodon crassifemoris Paramonov stat. n.

(figs. 62 a-c, 95)

Merodon spinipes (Fabricius) var. crassifemoris Paramonov,
1925: 158. Holotype d: ‘Theodosia, Krym, 13.vi.1923
[Crimea, U. S. S. R.] [not examined].

Merodon spinipes var. crassifemoris, Liepa 1969: 20; Peck
1988: 174.

Lampetia spinipes var. crassifemoris, Sack 1931: 328.

Material examined. — France: 16 (ZMAN); Greece: 26
(ZMAN); Turkey: 88 (WH, JLR); U. S. S. R.: 16, labelled
‘Karadagh Krym 16 vii 1929 S. Paramonov. leg. / Typus
Merodon spinipes Fbr. var crassifemoris var. nov. d | v.
Karodke Pinab’ (wrongly labelled as a type!) (ZMHB);
Yugoslavia: 16 (ZMAN).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin convex, apex very obtuse, antennal ratio
1.5, vertex angle = 30°, ocellar angle = 40°, tl-v ratio
0.5; pubescence creamy white with dark tuft in ocel-
lar region, face dense whitish pruinose, compound
eyes bearing dense white pubescence.

Thorax: Dark, metallic blue lustrous on lateral dor-
sum and scutellum, katepisternum, anepisternum
and anepimeron, longitudinal pruinose bands vesti-
gial to absent; pubescence dark yellow to brownish
with vague dark interalar band. Wings clear; halteres,
squamae and antisquamae yellow.

Legs: All dark except joints; trochanter 3 angular,
femora 3 markedly curved and swollen, with smallish
rather steeply rising triangular processus bearing 6-10
bristles on weakly serrate distal margin; tibiae 3 with
distal outer angle drawn out into small spur; tarsi 3
with golden pubescence on sides contrasting dark pu-
bescence on upper face, as in nigritarsis.

Abdomen: Dark; T II with vague brownish lateral
spots, distinctly tapering; T II-IV bearing arcuate
conspicuous greyish interrupted pruinose bands; pu-
bescence yellowish grey on bands, dark elsewhere; S
IV keeled throughout, posteriorly deeply emarginate.

Genitalia: The anterior surstyle lobe relatively
small, reniform, showing dense, short, yellow, even
pubescence apically and on margins; coalescent with
enormously elongate posterior surstyle lobe which
bears rather dense, yellow, uneven pubescence ven-
trally; posterior part of posterior surstyle lobe with
free anteriorly projecting cuneiform part, with dense
yellow pubescence laterally and postero-ventrally.
Cercus very small, bearing dense, long, yellow pubes-
cence. Aedeagus rather stout, apical shaft part short,
fringed plates on thecal apex oblique to erect.

193

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Body length 12-14 mm.
Female. — Unknown.

Diagnosis. — M. crassifemoris can be easily separat-
ed from other Merodon by the anteriorly projecting
posterior surstyle lobe; externally the species resem-
bles avidus, nigritarsis, femoratoides and the species in
the alagoezicus group.

Period of flight and distribution (fig. 95). — The
species occurs in the Mediterranean and Black Sea re-
gions in June and July.

Discussion. — Because type-material of crassifemor-
is was not available, identification of this species is
largely based on the original description and the spe-
cimen in ZMHB, identified as crassifemoris by Pa-
ramonov himself. Paramonov described var. crassif-
emoris as a variety of spinipes Fabricius (= avidus
Rossi) in view of several external similarities. The
genitalia however are rather different and do not indi-
cate a close relationship between avidus and crassifem-
oris.

The elegans group

Apomorphy: Thin chitinous flanges running along
outer face of aedeagus.

Description. — Head: Antennae orange to dark, an-
tennal ratio 1.5-1.8, vertex angle 25-35°, ocellar angle
35-55°, tl-v ratio 0.35-0.6; pubescence white to yel-
low. Thorax: Pubescence rather short, even; whitish
to pale brown. Legs: Dark, usually with yellow mark-
ings on apices of femora and at both ends of tibiae;
trochanter 3 often edged; femora 3 swollen, strongly
curved. Abdomen: Slender to rather stout, T II
strongly to moderately tapering; T II-IV bearing ar-
cuate pruinose bands, lateral spots variable. Genitalia:
The anterior surstyle lobelarge, elongate with concave
dorsal margin except manicatus, anterior and posteri-
or surstyle lobe separated by sulcus; aedeagus with
chitinous flanges on outer face.

Merodon bequaerti sp. n.
(figs. 63 a-c, 98)

Type-material: & Holotype ‘Noiseux Oran Algeria Dr J
Bequaert, 23.1V.10’ (written on lower face) / Merodon parie-
tum Mg, 3” (MNHN). 19 paratype, with label as holotype
(MNHN).

Description

Male. — Head: Antennae dark, antenna 3 somew-
hat paler, densely yellowish pruinose, upper margin
slightly concave, apex acute, antennal ratio 1.8; vertex

194

angle 30°, ocellar angle 40°, tl-v ratio 0.6; pubescen-
ce pale yellow with strong dark tuft in ocellar region,
occiput deep yellow pubescent; face rather bald,
black-bluish lustrous; compound eyes with rather
sparse, pale, yellow, short pubescence.

Thorax: Rather dull, some bluish lustre on sides;
pubescence even, short, pale brown, erect. Wings
tinged evenly, halteres, squamae and antisquamae yel-
low.

Legs: All dark; the tarsi, especially tarsi 3 with
dense recumbent golden pubescence; trochanters 3
with ridge on lower face, femora 3 quite swollen,
rather curved, with triangular processus bearing 10
bristles; tibiae 3 rather slender, tarsi 3 normally
shaped.

Abdomen: Dark, moderately slender, with hardly
conspicuous interrupted arcuate whitish pruinose
bands; T II somewhat tapering, with orange lateral
spots; pubescence rather dense, orangish brown (par-
tially worn off on holotype); T III-IV laterally with
cupreous lustre; all S dark, bearing scattered pale erect
pubescence; S IV slightly vaulted, its posterior margin
entire.

Genitalia: The anterior surstyle lobe ovoid, bearing
yellowish pubescence anteriorly and along ventral
margin and showing ventro-posterior angle; deep sul-
cus present between surstyle lobes, connected by a
stalk, visible as a broad chitinous plate in top view;
the posterior surstyle lobes square, with strong cen-
tro-posterior and lesser ventral accessory lobe; pubes-
cence golden, longest, erect on ventral margin and
lobes; the centro-posterior lobe bears a single black
bristle. Cercus short, wide, showing pale, yellow,
long, erect pubescence. Aedeagus with outer face ex-
cavate subbasally on outer face, there with paired free
chitinous plates, and bearing paired flanges running
along all of the outer face; apical shaft part slightly
lengthened, fringed plates on thecal apex suberect to
recumbent.

Body length 13.5 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Antenna 3 rounded
apically, though clearly concave on upper margin;
frons dark, slightly greenish metallic lustrous, sparse-
ly pubescent (probably artifact; head has been glued
on to the body at some time) ocellar angle 50°.
Thorax: Pubescence shorter than in male. Legs:
Trochanters 3 smooth. Abdomen: Less dense pubes-
cent except on lateral spots; T II less tapering; T V
dark, tapering.

Body length 13 mm.

Diagnosis. — M. bequaerti can be easily recognized
by the stout build of both sexes in comparison with
other species of the e/egans group. The females may

pose problems but the combination of swollen femo-
ra 3 and arcuate pruinose bands on the abdomen will
separate this species from e.g. ruficornis and its allies
which have more conspicuous, less arcuate pruinose

bands.

Period of flight and distribution (fig. 98). — This
species is known only from the types, collected in

Algeria in April.

Etymology. — The epitheton bequaerti has been de-
rived from the collector s name on the labels of both
type specimens. À noun in genitive case.

Merodon elegans sp. n.
(figs. 64 a-c, 99)

Type-material: 4 Holotype ‘Italia Sicilia V. S. v. d. Goot/
Etna rif. Filiciusa 1400-1500 m 22-28.vii.1961 / Lampetia
spinipes det. V. S. v.d. Goot 1963 / Type A’ (ZMAN). —
Paratypes: 52d, 249; 198, 10? with same data as holo-
type but collectors V. S. v.d. Goot & J. A. W. Lucas (14d
JLR, 58, 102 RMNH); 16 ‘Italia, S. Castaldo vi.1937, A.
Loken leg.’ (ZMUC); 38 ‘Sicilia Schiodte’ (ZMUC); 16
‘Italia, Calabria 18.v.1954 F. F. Tippmann’ (RMNH); 1d
‘France Aveyron 19.vii.1980 D. Prins’ (Coll. D. Prins,
Rhenen, The Netherlands); 38 ‘France Vaucluse Cheval
Blanc 7.vii.1973 H. I. Meuffels (JLR); 19, same data, col-
lected 9.vii.1973 LR); 28 ‘France Estagel 22.vi.1982 exc.
LH Wageningen’ (LHW); 2d ‘France, Chateau de
Peypertuse 25.vi.1982 exc. LH Wageningen (LHW); 1d
‘France, Mts. de Tauch, exc. LH Wageningen 23.vi.1982’
(LHW); 1d ‘S. F. Chaines des Alpilles, Les Baux 21.vii.1980
Barkemeyer (Coll. C. Claussen, Flensburg); 3d ‘Espana
Torla Huesca 1035 m, 8/26 vii.1974 J. Wolschrijn’ (JLR);
54 Espana Valencia Moraira 16/30.v.1981 van Aartsen (3
JLR, 1 RMNH, 1 ZMAN); 42 ‘Espana, Guadalajara, Molina
1100 m 11.vii.1977 W. Schacht (LR); 19 ‘Espana
Tragacete 1300 m, 17.vii.1974 W. Schacht’ (LR); 2d
‘Espana Catalonia, Espluga de F., 3.vii.1920 Codina’
(RMNH); 1d ‘Espana Catalonia Carretera [=road] de
Espluga de F. a Massies’ (RMNH); 1d, 19 ‘Espana Teruel
M. J. & J. P. Duffels / Rafales 35 km ZO v. Alcaniz 600 m
12.vii.1972° (ZMAN); 38 ‘Espana, Lebena (Santander)
5.vii.1986 Ma. A. Marcos Garcia’ (SAL); 16, 19 “Algerie,
Alger Dr. Bequaert, Foret de Buinen (MNHN); 1d ‘Algerie,
Alger Dr. Bequaert Benandreit’ (MNHN); 1d ‘Algeria, Le
Tarf 27.vii.1896 A. E. Eaton’ (BMNH); 38 ‘Maroc Meh-
dia (Ft. Lyautey) 31.v 1.vi.1950 P. M. F. Verhoeff 3d (-
RMNH).

Description

Male. — Head: The antennae orange to brown, an-
tenna | darkest, antenna 3 with upper margin slight-
ly convex, apex obtuse, antennal ratio 1.6; pubescen-
ce dull yellow, erect, even, with rather large dark tuft
in ocellar region; vertex angle 30°, ocellar angle 50°,
tl-v ratio 0,6; face and occiput diffusely whitish prui-
nose, blue metallic lustrous.

Thorax: Dark, with inconspicuous longitudinal

Hurkmans: Monograph of Merodon

pruinose bands; lateral dorsum and scutellum as well
as sides blue metallic lustrous; pubescence brownish
to yellow, erect, short, rather even, a dark interalar
band present in some specimens; pubescence on pos-
terior margin of scutellum longer; some diffuse prui-
nosity on dorsum and sides. Wings clear to evenly
slightly tinged throughout; halteres, squamae and
antisquamae yellow.

Legs: Dark with yellow markings apically on femo-
ra, both ends of tibiae and most of tarsi; trochanter 3
smooth, femora 3 quite swollen, metallic lustrous,
with normally shaped triangular processus bearing 6-
9 bristles on moderately serrate distal margin.

Abdomen: Dark; T II strongly tapering, with large
pale yellow to deep orange vaguely bordered lateral
spots; pruinose bands white, interrupted, moderately
wide, arcuate; pubescence yellowish, densest laterally;
S IV rather vaulted, narrowly emarginate posteriorly.

Genitalia: The anterior surstyle lobe large, reni-
form, with short, dense, even, yellow pubescence;
separated from posterior surstyle lobe by deep sulcus;
posterior surstyle lobe high, large, bearing rather
long, dense, yellow pubescence laterally and on ven-
tral-posterior margin, with lateral accessory lobe bear-
ing dense concolorous pubescence. Cercus angular,
well protruding, densely long yellow pubescent.
Aedeagus rather slender, with paired high chitinous
flanges on outer face and bearing paired basal humps;
apical shaft part short, fringed plates on thecal apex
oblique to recumbent.

Body length 13-15.5 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons with lustrous
midstripe occupying less than one-third of width;
clear frontal depression of ovoid outline present.
Thorax: Pruinose bands clearer, in some specimens a
fifth anteromedial band can be found. Legs: Orange
markings on basal femora more extensive, femora 1
and 2 often apically widely yellow. Abdomen: T II
conspicuously bright orange; pruinose bands on T III
IV continuous; pubescence sparser than in male.

Body length 13-15 mm.

Diagnosis. — The conspicuous flanges on the ae-
deagus distinguish e/egans from species outside the
elegans group. M. elegans may be separated from be-
quaerti by the longer posterior surstyle lobe, the slen-
derer habitus and greater size; from manicatus by the
different shape of the surstyle and possession of an ac-
cessory lobe on the posterior surstyle lobe in elegans,
from testaceus also in the surstyle shape, the height of
the aedeagal flanges, the overall size and the absence
of typical red abdominal coloration as in testaceus.
The females of elegans are similar to the females of the
alagoezicus group, avidus and many other species,

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

from which they can be distinguished by their frontal
depression.

Period of flight and distribution (fig. 99). — M. ele-
gans occurs from May to July in the western part of
the Mediterranean.

Discussion. — Van der Goot (1964a) noticed differ-
ences among sets of specimens then considered to be-
long to avidus, and distinguished between types A
and B. Examination of the genitalia showed that type
B corresponded to avidus whereas type A was shown
to be a new species, described here. The flight period
coincides with that of avidus in the mediterranean re-
gion; the distribution of e/egans is much more restrict-
ed than that of avidus and seems to be Mediterranean.
As the species are very similar externally (elegans has
longer, rougher pubescence on the thorax and more
metallic lustre on face and femora 3, which is more
swollen than in avidus) and their flight period over-
laps, some of the avidus records in literature probably
pertain to elegans.

Etymology. — The adjective elegans is latin, mean-
ing well-formed, and refers to the slender habitus of
the species.

Merodon manicatus (Sack)

(figs. 65 a-c, 98)

Lampetia manicata Sack, 1938: 21. Holotype 6: ‘Stip,
v.1937, leg. R. Meyer/ Lampetia manicata Sack det. Sack’
(HLMD) [examined].

Lampetia manicata; Sack 1938: 21.

Merodon manicatus, Peck 1988: 171.

Material examined. — Macedonia: 9 Paratype with same
labels as holotype (HLMD).

Description

Male. — Head: Antennae dark, antenna 3 ovoid,
upper margin strongly convex, apex obtuse, antennal
ratio 1.6; pubescence pale yellow with some dark pu-
bescence in ocellar region; vertex angle 30°, ocellar
angle 55°, tl-v ratio 0.35; face rather densely white
pruinose; compound eyes bearing rather short, dense,
white pubescence.

Thorax: The dorsum and scutellum showing weak
olivaceous lustre laterally; same lustre stronger on kat-
episternum, anepisternum and anepimeron; longitu-
dinal pruinose bands weak; pubescence golden, erect,
moderately dense, even, slightly longer and paler lat-
erally. Wings tinged overall; halteres, squamae and
antisquamae pale yellow.

Legs: Dark, with some olivaceous lustre on femora
3; tibiae with yellow markings at both ends; tarsi
dark, but with recumbent dense golden pubescence

196

on lower face, strongest on tarsi 3; trochanter 3 with
distinct sharp ridge, femora 3 strongly swollen, much
curved, bearing normally shaped triangular processus
bearing 7 bristles on moderately serrate distal margin.

Abdomen: Dark; T II with large orange lateral
spots, continued on T III; T II-IV showing rather
faint white pruinose arcuate bands best observed from
behind, hardly interrupted medially; all S orange,
bearing erect white pubescence; S IV darker, slightly
keeled, deeply narrowly emarginate posteriorly; ab-
dominal pubescence in general very sparse compared
to other Merodon, concolorous with surface and very
short medially.

Genitalia: The anterior surstyle lobe reniform,
bearing dense short even yellowish pubescence anteri-
orly and ventrally, separated from the posterior sur-
style lobe by a moderately deep sulcus; the posterior
surstyle lobe with yellow pubescence, bearing distinct
anteromedial accessory lobe with pubescence erect
and long; main disc of posterior surstyle lobe bearing
shorter pubescence, somewhat diamond-shaped.
Cercus angular, with same pubescence as posterior
surstyle lobe. Aedeagus moderately slender, bearing
paired translucent, twice interrupted (artifact?) chi-
tinous flange on outer face, apical shaft part slightly
lengthened, fringed plates on thecal apex oblique;
paired humps present basally on outer face.

Body length 11.5 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons with lustrous
midstripe occupying rather less than a quarter of the
frontal width just above the antennae, but wider pos-
teriorly; ocellar angle 60°. Thorax: Pruinose bands
more clear due to shorter, less dense pubescence; sides
much less lustrous than in male. Legs: The femora 3
less swollen, less curved, less lustrous. Abdomen:
Predominantly orange; T V with dark median zone;
pruinose bands widely interrupted; pubescence very
sparse, even shorter than in male.

Body length 11 mm.

Diagnosis. — The males manicatus are distinguished
from those of bequaerti by the much slenderer habit-
us and more rounded anterior surstyle lobe as well as
the sparse pubescence; from those of elegans also by
the rounded anterior surstyle lobe, and much smaller
size; from those of testaceus by the less slender habitus
of manicatus and its less tapering T II, and the more
rounded anterior surstyle lobe; from all other species
by the aedeagal flanges. The females of manicatus are
slenderer than those of elegans and bequaerti, but are
hard to separate from those of testaceus, they can how-
ever be distinguished by the more acute ocellar trian-
gle with ocellar angle 60°, (70° in testaceus). From all
other species they differ in the combination of ex-

tremely slender build, moderate size and red abdomi-
nal colour, which is of a quite different type than
found in the longicornis group where the colour is
deep crimson.

Period of flight and distribution (fig. 98). — The
species occurs in Macedonia in May, and is currently
known only from the types.

Merodon testaceus Sack

(figs. 66 a-c, 98)
Merodon testaceus Sack, 1913a: 445. Lectotype d (here des-

ignated): “Asia minor/ Alte Sammlung/ Merodon testaceus

type Sack/ testaceus det. Sack’ (NHMW) [examined].
Merodon testaceus, Paramonov 1925: 154; Peck 1988: 175.
Lampetia testacea; Sack 1931: 329, 1938: 21.

Material examined. — 3d, 19 Paralectotypes, ‘Asia
Minor/ idae det. Schiner/ Alte Sammlung/ testaceus det.
Sack’ (NHMw); 29 Paralectotypes, ‘Mann, Brussa [=Bursa,
Turkey], 1863/ idae det. Schiner/ Alte Sammlung/ testaceus
det. Sack’ (NHMW).

Description

Male. — Head: Antennae orange to brown, antenna
3 with upper margin convex, apex obtuse, antennal
ratio 1.8; vertex angle 25°, ocellar angle 35°, pubes-
cence pale yellow, tl-v ratio 0.35.

Thorax: Dark, with pruinose bands inconspicuous;
dorsum and scutellum dark, katepisternum, anepis-
ternum and anepimeron blue metallic lustrous, pu-
bescence rather sparse, short, whitish. Wings clear,
halteres, squamae and antisquamae yellow.

Legs: The femora dark, tibiae dark with yellow
markings at both ends, tarsi brown to yellow, in some
specimens with faint dark band; trochanters 3 with
conspicuous edge, femora 3 strongly swollen, curved,
with normally shaped triangular processus bearing 6
stout bristles on rather weakly serrate distal margin
with relatively large apical spurlet.

Abdomen: Slender; T II strongly tapering; pre-
dominantly orange to reddish brown, with clear arcu-
ate narrowly interrupted white pruinose bands on T
II-IV; S IV slightly keeled, deeply emarginate posteri-
orly. Genitalia: The anterior surstyle lobe bearing
dense, short, yellow, even pubescence apically and on
margins, with incision on dorsal margin, ridged later-
ally, separated from the posterior surstyle lobe; the
posterior surstyle lobe with anterior accessory lobe
bearing dense long yellow pubescence; main disc
rather elongate, protruding far posteriorly, bearing
rather dense yellow pubescence posteriorly. Cercus
angular, with dense long yellow pubescence.
Aedeagus moderately slender, bearing large paired
humps subbasally on outer face as well as paired chi-
tinous flanges; apical shaft part short, fringed plates

Hurkmans: Monograph of Merodon

on thecal apex recumbent.

Body length 10 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Frons black, lustrous
on median one-third, sides diffusely whitish prui-
nose, pubescence concolorous, ocellar angle 70°.
Thorax: Pubescence notably short, pruinose bands
more conspicuous.Legs: Femora 3 less swollen.
Abdomen: Slightly stouter than in male, orange to
brown throughout, pruinose bands slightly less con-
spicuous.

Body length 10.5 mm.

Diagnosis. — The males of testaceus can be distin-
guished from those of bequaerti by the less stout hab-
itus, from males of elegans and manicatus by the low-
er aedeagal flanges, and from those of elegans by
smaller size as well. Moreover, they differ from males
of manicatus also by slenderer abdomen, less rounder
antenna 3 and shorter thoracic pubescence. From all
other Merodon they differ by the aedeagal flanges.
The females can be distinguished by their red abdom-
inal coloration, size and slender habitus from all spe-
cies. The differences with females of manicatus are:
the larger ocellar angle (70° here, 60° in manicatus)
and the slightly greater antennal ratio (1.8 versus 1.6
in manicatus).

Period of flight and distribution (fig. 98). — M. tes-
taceus occurs in Turkey; the flight period is unknown.

The alagoezicus group

Apomorphies: Apical shaft part lengthened in ae-
deagus of the d; fringed plates on thecal apex of ae-

deagus oblique to erect.

Description

Head: Antennae variably coloured but antenna 3
often paler than rest; face with often strong blue me-
tallic lustre; pubescence yellow to whitish; vertex ang-
le in the males 25-35°, ocellar angle in both sexes ap-
proximately 45°.

Thorax: Dark, with longitudinal pruinose bands
on dorsum more or less clear; katepisternum, anepis-
ternum and anepimeron often with very strong blue
lustre. The females show 5 longitudinal pruinose
bands.

Legs: The usual pattern of coloration is: femora
dark, tibiae yellow with wide dark band and some-
times with distal spur, tarsi dark; the tarsi 3, especial-
ly basitarsi 3 often with dense golden recumbent pu-
bescence on sides, contrasting dark pubescence on
upper face; this contrast is less strong in the females.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Abdomen: Slender, T II strongly tapering, T IV of-
ten bulging to hold the relatively large genitalia in the
males; usually slender, triangular in the females; prui-
nose bands strongly arcuate, the one on T IV in the
males often recurving laterally; lateral portions of ter-
gites often metallic lustrous.

Genitalia: The anterior surstyle lobe lengthened,
extremely lengthened in some species; accessory lobes
may be present on both main surstyle lobes. Cercus
usually large, always well protruding. Aedeagus slen-
der, invariably with extremely lengthened apical shaft
part, outer face often with humps or plates; fringed
plates on thecal apex suberect to erect.

Identification of females in the alagoezicus group. —
Apart from the fentales of satdagensis which have been
so assigned on the grounds of associated capture, the
females in this group cannot be assigned to any spe-
cies yet. As a group these females can be distinguished
from those of other groups by the presence of 5 prui-
nose bands on the thorax (4 bands in other groups),
relatively large size and slender, usually triangular ab-
domen.

Merodon alagoezicus Paramonov

(figs. 67 a, 68 a-d, 99)

Merodon alagoezicus Paramonov, 1925: 151. Syntypes d:
‘Inaklu, Etschmiadzin, Erivan district, Armenia,
Southern slope of Mount Alagéz’ [not examined].

Merodon alagoezicus, Paramonov 1926a: 15;1926b: 318;
1927: 74; Liepa 1969: 20; Peck 1988: 166.

Lampetia alagoécica; Sack 1931: 303 [orthographic error].

Material examined. — Greece: 38 (ZMUC); Turkey: 81 4
(LR, WH, BMNH).

Description

Male. — Head: Antennae brown to pale brown, an-
tenna 3 palest, with upper margin convex, apex obtu-
se, antennal ratio 1.7; pubescence pale yellow, conspi-
cuously dense on frons; face strongly blue metallic
lustrous, sparsely pruinose throughout; vertex angle
30°, ocellar angle 40°, tl-v ratio 0.65; compound
eyes bearing inconspicuous pale pubescence.

Thorax: Dark; blue lustrous on lateral dorsum and
scutellum, katepisternum anepisternum and anepim-
eron; pubescence rather even, moderately long, pale
yellow on dorsum, longer and less even on sides; in
many specimens vague dark interalar band present;
longitudinal pruinose bands vestigial to well-defined.
Wings clear; halteres, squamae and antisquamae yel-
low.

Legs: The femora dark with narrow paler apices;
tibiae dark with yellow apices and, in most speci-
mens, yellow bases; tarsi dark, golden pubescence on

198

sides and lower face, upper face dark; trochanter 3
with inconspicuous low ridge, femora 3 swollen,
slightly curved, with normally shaped triangular pro-
cessus bearing 6-9 bristles on serrate distal margin;
tibiae 3 with distal outer angle much projecting, spur-
like; tarsi 3 normally shaped.

Abdomen: Dark; T II tapering, bearing yellow to
orange lateral spots with deep lanceolate postero-lat-
eral dark indention; T II-IV with arcuate interrupted
moderately wide pale pruinose bands, on T IV re-
curving laterally; pubescence mainly pale, with some
dark pubescence on dark surface in most specimens; S
IV widely, rather acutely emarginate posteriorly up to
two-thirds of its length.

Genitalia: Anterior surstyle lobe ventro-posteriorly
recurved, enormously elongate, its apical portion
showing dense, even, yellow, short pubescence; the
posterior surstyle lobe coalescent, rather low, with
moderately dense, yellow, erect pubescence along en-
tire medio-apical margin and posteriorly. Cercus
large, rounded, showing dense, long, erect, yellow pu-
bescence. Aedeagus smooth, with paired recumbent
chitinous plates on outer face guarding the subapical
cavity; apical shaft part long, the fringed plates on
thecal apex erect.

Body length 12-15 mm.

Female. — Unknown (see preceding observation).

Diagnosis. — In view of the curious genitalia ala-
goezicus can be confused with /ucasi only; from that
species it is distinguished by the projection at the dis-
tal outer angle of tibiae 3 and the lack of a semicircu-
lar projection on the posterior surstyle lobe; alagoezi-
cus can be easily distinguished from the other
members of this group by the much longer anterior

surstyle lobe.

Period of flight and distribution (fig. 99). — The
species occurs from June through August in south-
eastern Europe, Turkey and Armenia.

Biology. — Specimens have been collected in wet
grassland on white- and yellow-flowering Umbelli-
ferae; the majority of the specimens was collected in
mountains at altitudes over 1500 metres.

Merodon lucasi sp. n.

(figs. 67 d, 69 a-c, 102)

Type-material: 8 Holotype ‘Turkey, Hakkari,
Suvarihalil pass 2300-2400 m, 11.viii.1983, leg. J. A. W.
Lucas’ (ZMAN). Paratypes: 22d: 8d with same data as holo-

_ type (LR); 48 ‘Turkey, Konya 1300 m, 5/6.viii.1981, leg.

H. Coene et al.’ (LR); 16 ‘Turkey, Hakkari, Suvarihalil pass
2.viii.1982 M. Kuhbandner (LR); 26 “Turkey Pr. Hakkari
Suvarihalil Pass SE Beytisebap 2300 m 2.viii.1982 and

27.vi.1985 W. Schacht (LR); 16 “Turkey, Kars, Handere
20 km W. Sarıkamıs 2100 2200 m, 1.viii.1983 leg. J. A. W.
Lucas’ (JLR); 28 “Turkey, Hakkari, Suvarihalil pass 2300 m
27.vi.1985 C. J. Zwakhals (LR); 16 ‘Turkey, Hakkari S of
Yiiksekova, Sat Daglari 1700 m 28.vi.1985 leg. W. Schacht
(LR); 16 “Turkey, Hakkari, Varegös Sat Daglari 1700 m
29.vi.1985 W. Schacht (LR); 16 ‘Turkey, Hakkari,
Habero Deresi valley 1200 m 25.vi.1985 W. Schacht (LR);
Id ‘Turkey, Erzurum, Palandöken Dagi 2300 m
11.vii.1986 leg. J. A. W. Lucas’ (JLR).

Description

Male. — Head: Antennae dark, antenna 3 with up-
per margin concex, apex obtuse to subacute, antennal
ratio 1.4; vertex angle 35°, ocellar angle 45°, tl-v ratio
0.6, pubescence pale yellow, denser on frons where
causing silky lustre; face bronze to blue lustrous, spar-
sely white pruinose.

Thorax: Dark; blue lustrous on katepisternum, an-
episternum, anepimeron and (less strong) on lateral
scutellum and dorsum; pubescence pale brown to yel-
low, moderately dense; pruinose bands inconspicu-
ous, the median bands clearest; a fifth anteromedian
band often present; these bands often cross-connect-
ed along dorsal suture. Wings clear; halteres, squamae
and antisquamae pale yellow.

Legs: Dark with yellow markings on femoral apices
and both ends of tibiae; trochanter 3 with low but
well marked sharp ridge; femora 3 swollen, curved,
triangular processus bearing 7-9 bristles on weakly
serrate distal margin with clear apical pedestal; tibiae
3 with apical outer angle projecting slightly, tarsi 3
normally shaped.

Abdomen: Dark; slender; T II strongly tapering, T
IV bulging; T II with conspicuous orange lateral
spots, T II-IV with arcuate interrupted moderately
wide, well bordered white pruinose bands; pubes-
cence concolorous but many pale yellow hairs mixed
in laterally and posteriorly on T III-IV, densest on
lateral spots; S IV widely acutely emarginate up to
half its length.

Genitalia: Anterior surstyle lobe ventro-posteriorly
recurved, enormously elongate, apically with dense
even short yellow pubescence; the posterior surstyle
lobe coalescent, anteriorly showing semicircular ven-
tral projection and bearing yellow moderately dense
erect pubescence. Cercus large, quadrate, with dense,
long, yellow pubescence. Aedeagus rather slender,
with paired humps on basal outer face and bearing
chitinous plates guarding subapical cavity, apical
shaft part long, fringed plates on thecal apex narrow,
long, erect.

Body length 11-15 mm.

Female. — Unknown (see preceding observation).

Diagnosis. — M. lucasi can be distinguished from

Hurkmans: Monograph of Merodon

the other members of the alagoezicus group by the
much longer anterior surstyle lobe, and moreover
from alagoezicus by the semicircular plate on the pos-
terior surstyle lobe and by having no large spur on tib-
iae 3.

Period of flight and distribution (fig. 102). — The
species is known from Turkey where it was collected
in June and August.

Etymology. — It is a pleasure to dedicate this species
to Jan Lucas who collected an enormous amount of
specimens which were generously placed at the
authors disposal. A noun in genitive case.

Merodon nitidifrons sp. n.
(figs. 70 a-d, 101)
Type-material: & Holotype “Turkey, Hakkari,

Suvarihalil pass 2300 m [37° 25’ N, 43° 03’ E], 13.vi.1984,
leg. J. A. W. Lucas’ (ZMAN).

Description

Male. — Head: Antennae orange, antenna 3 with
convex darkened upper margin, apex subacute, an-
tennal ratio 1.3; vertex angle 25°, ocellar angle 35°,
tl-v ratio 0.55; pubescence yellow, densest and re-
cumbent on frons where causing silky lustre; face
densely white pruinose over blue metallic surface;
some dark pubescence present in ocellar region.

Thorax: Dark, blue lustrous on katepisternum, an-
episternum and anepimeron, lateral dorsum and scu-
tellum; conspicuous longitudinal bands on dorsum in
two pairs and a fainter anteromedian band; pubes-
cence dull golden, moderately dense, most even.
Wings clear; halteres, squamae and antisquamae yel-
low.

Legs: Dark but femoral apices and both ends of tib-
iae orange; trochanters 3 smooth, femora 3 slightly
swollen, triangular processus rather low, bearing 4-6
rather weak bristles on hardly serrate distal margin;
tibiae 3 with distal outer angle somewhat projecting.

Abdomen: Slender, T II much tapering, T IV bulg-
ing; T II-III with pale orange lateral spots, T I-IV
with arcuate rather wide pale pruinose bands (inter-
rupted on II-III, continuous and laterally recurved on
IV), pubescence concolorous; on closer examination
T IV shows small orange anterolateral spots.

Genitalia: The anterior surstyle lobe much elon-
gate, narrow, evenly wide throughout, apically re-
curved medially and bearing dense, short, even, yel-
low pubescence; basally bearing accessory lobe on
medial face; this lobe is visible as a ridge in lateral
view and has same pubescence as apical anterior sur-
style lobe part; the posterior surstyle lobe coalescent,
moderately high, elliptical, bearing moderately dense

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

long yellow pubescence in strip along margins.
Cercus wide, with long, dense, somewhat wavy, yel-
low pubescence. Aedeagus very slender, bearing
paired chitinous plates guarding subapical cavity, ap-
ical shaft part long, fringed plates on thecal apex sub-
erect.

Body length 15.5 mm.
Female. — Unknown. (see preceding observation)

Diagnosis. — M. nitidifrons is easily distinguished
from other members of the alagoezicus group by the
ribbon-shaped, medially recurved anterior surstyle

lobe.

Period of flight and distribution (fig. 101). — This
species is known only from the holotype, collected in
eastern Turkey in June.

Etymology. The epitheton nitidifrons derives from
the latin nitere, to shine, and frons, a part of the head
so indicated, and refers to the silky lustrous frons. It is
to be treated as a noun in apposition.

Merodon satdagensis sp. n.
(figs. 67 c, 71 a-c, 101)

Type-material: & Holotype ‘Turkey, Hakkari, Suvarihalil
pass 2300 m, SE Beytisebap 2.viii.1982 leg. W. Schacht’
(ZMAN). Paratypes: 31d, 22; 2d with same data as holotype
(LR); 54 topotypic, 4.viii.1982, leg. J. A. W. Lucas (LR) 5d
topotypic, 1 1.viii.1983, leg. J. A. W. Lucas (LR); 36 “Turkey,
Hakkari, Varegôs S. of Yüksekova 1700 m 29.vi.1983 leg. W.
Schacht’ (LR); 12d from same locality, 5-8.viii.1983, leg.
Schacht (JLR), 16, 29 from same locality, 5-8.viii.1983, leg.
Lucas (JLR); 16 “Turkey, Hakkari, Sat Dag, North of Mt.
Gavaruk 2900 m, 7.viii.1983 leg. Kl. Warncke’ (LR); 1d
‘Turkey, Hakkari, Habur Deresi S. of Yüksekova, 26.vi.1985,
leg. Schacht’ (LR); 18 “Turkey, Hakkari, S of Yüksekova
1700 m, 28.vi.1985, leg. W. Schacht’ (LR).

Description

Male. — Head: Antennae dark orange to brown, an-
tenna 3 paler, upper margin convex, apex obtuse, an-
tennal ratio 1.3, vertex angle 35°, ocellar angle 45°;
pubescence yellowish white, rather dense on face and
frons; face and vertex blue metallic lustrous, modera-
tely dense white pruinose; tl-v ratio 0.3.

Thorax: Dark, katepisternum, anepisternum ane-
pimeron, lateral dorsum and scutellum blue to green
or slate grey lustrous; two pairs of narrow longitudi-
nal pruinose bands present; pubescence pale brown to
yellow, moderately dense, rather even. Wings clear;
halteres, squamae and antisquamae pale yellow.

Legs: Dark, but tibiae yellow with wide dark distal
band, femora apically yellow in most specimens; tro-
chanters 3 smooth, femora 3 swollen, curved, trian-
gular processus normally shaped, bearing 7-9 bristles

200

on serrate distal margin; tibiae 3 apically with spurlike
outer angle.

Abdomen: Dark, slender, T II strongly tapering,-
with large pale yellow lateral spots; T II-IV with mod-
erately wide interrupted (continuous on T IV) whit-
ish strongly arcuate well bordered pruinose bands,
sometimes combined with red surface coloration; T
IV bulging; pubescence concolorous but many pale
hairs prsent throughout; S IV deeply widely emargi-
nate, strongly vaulted; T IV posteriorly pruinose.

Genitalia: The anterior surstyle lobe much elon-
gate, apical part recurved, dorso-ventrally aligned,
with subbasal ventral accessory lobe; both apical part
and this lobe short dense erect even yellow pubes-
cence; the anterior and posterior surstyle lobes separ-
ated by shallow sulcus; posterior surstyle lobe elon-
gate, twisted, with long, yellow pubescence
posteriorly and along medial distal margin. Cercus
large, elongate, with denser pubescence than posteri-
or surstyle lobe. Aedeagus extremely long, slender,
bearing paired chitinous flanges guarding subapical
cavity which due to extreme length of apical shaft part
is situated halfway up the aedeagus; fringed plates on
thecal apex erect, rather long and narrow.

Body length 12-15 mm.

Female. — Except for sexual dimorphism, differing
from the male as follows. Head: Lustrous midstripe
on frons occupying 1/4 of width; ocellar angle 45°,
antennae with third article darkest. Thorax:
Pubescence shorter, more even, a fifth anteromedian
pruinose band present. Legs: the spurlet on tibiae 3
lacking. Abdomen: T II less tapering, lateral spots
larger, pruinose bands less wide; T IV posteriorly
pruinose.

Body length 13 mm.

Diagnosis. — M. satdagensis males will be easily dis-
tinguished by the long anterior surstyle lobe with sub-
basal accessory lobe and twisted posterior surstyle
lobe. The females are very similar to other females of
species in this group; they can preliminarily be distin-
guished from those of other species by the strongly
serrate triangular processus; see preceding observa-
tion.

Period of flight and distribution (fig. 101). — M.
satdagensis is known from south-eastern Turkey
where it occurs late in June and in August. The
known range of the species is restricted to the Sat
Mountains in the Turkish province of Hakkari near
the Iranian and Iraqi borders.

Discussion. — The females have been assigned to
satdagensis given their concurrent flight in the same
biotope at Varegös in the Sat mountains of eastern
Turkey; moreover the much serrate triangular proces-

sus in these females occurs in males of only two spe-
cies of the alagoezicus group, viz. satdagensis and tani-
niensis, of which the latter is stouter, much lighter
coloured and in the possession of much wider ab-
dominal pruinose bands; the females concerned being
relatively dark and slender were assigned to satdagen-
sis.

Etymology. — The epitheton is an adjective, and
named after the Sat Daglari, a mountain range in
south-eastern Turkey.

Merodon schachti sp. n.
(figs. 67 b, 72 a-c, 100)

Type-material: 8 Holotype ‘Turkey, Hakkari,
Suvarihalil pass 2300 m, 2.viii.1982 leg. W. Schacht’
(ZMAN). Paratypes: 9 ; 24 topotypic, 11.viii.1983 leg. J. A.
W. Lucas (JLR); 16 topotypic, 13.vi.1984 leg. Lucas (LR);
16 ‘Turkey, Hakkari, Tanin-Tanin pass 1700 m (W side
near Habul Deresi) 12.vi.1984 leg. Lucas’ (LR); 3d from
same locality, 25.vi.1985 leg. W. Schacht (LR), 1d from
same locality, 25.vi.1985 leg. C. J. Zwakhals (LR); 1d
‘Turkey, Hakkari, Habur Deresi valley 26.vi.1985 1700 m
leg. Schacht’ (JLR).

Description

Male. — Head: Antennae orange-brown, antenna 3
slightly paler, upper margin convex, apex obtuse, not
rounded, antennal ratio 1.3; pubescence whitish,
densest on frons; face blue metallic lustrous, diffusely
white pruinose (densest medially); vertex angle 25°,
ocellar angle 40°, tl-v ratio 0.4.

Thorax: Dark, blue lustrous on katepisternum, an-
episternum and anepimeron, lateral dorsum and scu-
tellum; pruinose bands on dorsum in anteriorly co-
alescent pairs, narrow; pubescence sparse, pale brown
to yellow. Wings clear; halteres, squamae and anti-
squamae pale yellow.

Legs: Dark with yellow markings on apices of fe-
mora and both ends of tibiae; trochanter 3 smooth,
femora 3 swollen, somewhat curved, triangular pro-
cessus bearing 7-9 bristles on slightly serrate distal
margin (less serrate than in satdagensis but more than
in alagoezicus); tibiae 3 with distal outer angle extend-
ed, tarsi 3 normally shaped.

Abdomen: Dark: slender; T II strongly tapering, T
IV bulging, T II-III with large and small orange later-
al spots respectively; T II with weak, T III-IV with
strong whitish interrupted arcuate pruinose bands,
the one on T IV recurved laterally; posterior margins
of T III-IV pruinose; pubescence pale, but strictly
dark on posterior, dark surface of T III; S IV acutely
emarginate posteriorly up to 3/4 its length or even
more.

Genitalia: Anterior surstyle lobe very elongate, re-
curved apically, there dorso-ventrally aligned, apically

Hurkmans: Monograph of Merodon

bearing dense, yellow, short, even pubescence; anteri-
or surstyle lobe coalescent with gradually rising poste-
rior surstyle lobe elongate, low, and bearing moder-
ately long, yellow pubescence along its distal margin
and posteriorly; posterior surstyle lobe laterally guard-
ed by a free chitinous plate basally attached to the sur-
style base; this plate has a distinct antero-ventral angle
and a lateral approximately dorso-ventral ridge and
posteriorly shows dense yellow pubescence. Cercus
large, with dense, long, erect, yellow pubescence.
Aedeagus quite robust, apical shaft part lengthened
much, fringed plates on thecal apex erect, long and
narrow, some scattered short pubescence present on
inner face.

Body length 13 mm.
Female — Unknown (see preceding observation).

Diagnosis. — M. schachti can hardly be confused
with any other species given the very conspicuous lat-
eral guarding plate alongside the posterior surstyle
lobe in the males; moreover, the robust aedeagus sep-
arates it from other species in the alagoezicus group.

Period of flight and distribution (fig. 100). — The
species is known exclusively from the south-eastern-
most region of Turkey where it has been collected in
June and August.

Etymology. — The species is named after the collec-
tor of the type, the German entomologist Dr.
Wolfgang Schacht, of Munich. A noun in genitive
case.

Merodon taniniensis sp. n.
(figs. 73 a-c, 100)

Type-material: d Holotype “Turkey, Hakkari, Tanin-
Tanin pass W. side 1700 m, 12.vi.1984 leg. J. A. W. Lucas’
(ZMAN). Paratypes: 68; 2d with same data as holotype
(LR); 26 ‘Turkey, Hakkari, Tanin-Tanin pass E Uludere
2300 m 25.vi.1985 leg. W. Schacht’ (LR); 18 “Turkey,
Hakkari, Suvarihalil pass near Habul Deresı 1250 m
13.vi.1984 leg. Lucas’ (LR); 1d “Turkey, Malatya, Sarıhacı
45 km W. Malatya 21.vi.1984 leg. Lucas’ (JLR).

Description

Male. — Head: Antennae brown, antenna 3 with
upper margin convex, apex obtuse, antennal ratio 1.6,
pubescence whitish pale yellow throughout; vertex
angle 25°, ocellar angle 45°, tl-v ratio 0.4; face blue
metallic lustrous, densely white pruinose.

Thorax: Dark, lustrous on katepisternum, anepis-
ternum, anepimeron and lateral dorsum and scutel-
lum; dorsum showing 5 pruinose bands; pubescence
yellow, very even, moderately dense. Wings clear; hal-
teres, squamae and antisquamae pale yellow.

201

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Legs: Dark but tibiae yellow with wide dark distal
band; trochanter 3 with low sharp ridge, femora 3
swollen, curved, triangular processus bearing 6-8 bris-
tles on somewhat serrate distal margin.

Abdomen: Slender; T II strongly tapering, T IV
bulging, predominantly orange, with connected ante-
rior and posterior dark lunules on T II and weaker
brown medial coloration on T III; posterior lunules
on T III-IV pale brown to orange; T II-IV bearing
rather wide strongly arcuate, interrupted to continu-
ous whitish pruinose bands (recurved on T IV); pu-
bescence pale yellow, moderately dense (also on dark
surface); all S orange, S IV widely, obtusely emargi-
nate, slightly vaulted.

Genitalia: The anterior surstyle lobe large, rather
quadrate, slightly bifid anteriorly, bearing dense short
erect yellow even pubescence; connected by a slender
stalk to the base of the posterior surstyle lobe, a deep
narrow sulcus separating from the apical part; the
posterior surstyle lobe twisted, bearing two apical
ridges, both bearing long, erect, yellow, sparse pubes-
cence, posterior part of posterior surstyle lobe with
dense pubescence. Cercus large, mainly hidden
between the elongate posterior parts of the posterior
surstyle lobes, with dense, long, yellow pubescence.
Aedeagus slender, apical shaft part lengthened, ring
paired chitinous plates guarding subapical cavity
(these plates do not resemble those found in the ele-
gans group); fringed plates on apex aedeagi erect, long
and rather narrow.

Body length 14-16 mm.
Female. — Unknown (see preceding observation).

Diagnosis. — M. taniniensis can be distinguished
from other species of the alagoezicus group by the rela-
tively very large, rather quadrate anterior surstyle lobe
and the double ridges on the posterior surstyle lobe.

Period of flight and distribution (fig. 100). —
Known from the central and southern parts of eastern
Turkey where it has been collected in June.

Etymology. — The epitheton taniniensis, an adjec-
tive, refers to the Tanin-Tanin pass, the type locality
of the species in south-eastern Turkey.

Merodon toscanus sp. n.
(figs. 74 a-c, 99)

Type-material: d Holotype ‘Italia Toscana, Careggi
(Firenze) 19-21.v.1986 W. Hurkmans’ (ZMAN).

Description
Male. — Head: Antennae brown, antennae 3 with
upper margin convex, apex obtuse, antennal ratio 1.6,

202

vertex angle 20°, ocellar angle 35°; pubescence pale
yellow, face metallic blue lustrous, densely whitish
pruinose; a dark hair tuft present in ocellar region.

Thorax: Dark, metallic lustrous on katepisternum,
anepisternum, anepimeron, lateral dorsum and scutel-
lum; pubescence dull yellow, even, rather sparse; two
pairs of rather faint pruinose bands present. Wings
clear, halteres, squamae and antisquamae pale yellow.

Legs: All femora dark brown, tibiae orange with
distal wide dark bands, tarsi deep orange to brownish;
trochanter 3 smooth, femora 3 rather strongly swol-
len, curved, triangular processus bearing 7 bristles on
weakly serrate distal margin.

Abdomen: Rather slender; T II tapering; T IV
bulging; colour mainly deep orange, T II bearing
connected anterior and posterior dark lunules, T III
with deep orange posterior and paler anterior part, T
IV deep orange throughout; T II-IV with arcuate
moderately wide pruinose bands, just interrupted on
T I-III, continuous and laterally recurved on T IV;
all S orange, S IV with drop-shaped emargination,
strongly vaulted.

Genitalia: Anterior surstyle lobe elongate, round-
ed, bifid anteriorly, bearing dense short erect yellow
even pubescence throughout (sparser posteriorly);
connected to base of posterior surstyle lobe by a short
stalk; the posterior surstyle lobe low, elongate, bear-
ing an accessory lobe with dense long erect pubes-
cence medially; its anterior part bald, lustrous, ap-
proximately ovoid, its posterior part with sparse
yellow pubescence, but denser on margins and poste-
riorly. Cercus large, with dense, erect, yellow, long
pubescence. Aedeagus very slender, with large paired
basal humps on outer face; apical shaft part much
lengthened; fringed plates on thecal apex suberect,
rather narrow.

Body length 12 mm.
Female. — Unknown (see preceding observation).

Diagnosis. — M. toscanus has an accessory lobe on
the posterior surstyle lobe and a bald anterior part of
the posterior surstyle lobe not found in the other spe-
cies of the alagoezicus group. The bifid anterior sur-
style lobe is much stronger than in taniniensts.

Period of flight and distribution (fig. 99). — The
unique holotype was collected in May in Toscana,
Italy.

Etymology. — The species has been named after
Toscana, the region where the holotype was collected.
An adjective.

REMAINING SPECIES

Two of the three species discussed under this head-
ing are known only after female specimens. Since
these species share character states considered apo-
morphic for some species treated in this paper, they
are discussed here. Of the third species, M. viaticus no
material was examined, but according to its descrip-
tion it might occupy a position close to some of the
species treated in this paper.

Merodon affinis Gil Collado

Merodon affinis Gil Collado, 1930: 243, 255. Holotype 2:
‘Escorial, Arias Encobet, vii.1905’ [probably El Escorial,
Madrid] (IEE) [examined].

Merodon affinis, Peck 1988: 166.

Description

Female. — In many respects the female affinis is si-
milar to avidus, with following differences. Head:
According to the original description, antenna 3
slightly curved, antennal ratio 2.0, apex acute (anten-
na 3 missing on the holotype on both sides). Legs:
Femora 3 with distal margin of triangular processus
much stronger serrate than in avidus. Abdomen:
Outline triangular, pruinose bands narrowly inter-
rupted, less arcuate than in avidus.

Body length 12 mm.

Male. — Unknown.

Discussion. — Because of several differences in the
females, affinis cannot simply be regarded a synonym
of avidus, especially not in absence of the males. The
status of this species remains uncertain. In view of the
mentioned similarities with the females of avidus, af-
finis could conceivably belong to the avidus group but
to e.g. the alagoezicus or nigritarsis group as well.

Period of flight and distribution. — Known only
from the holotype, collected in central Spain in July.

Merodon aureotibia sp. n.

(fig. 106)

Type-material: 2 Holotype ‘Turkey, Adiyaman, Nemrut
Daglari, 1.vi.1983, leg. M. Kuhbandner (ZMAN). Paratypes:
32 ‘Tk [Turkey] 5-6-1988 10 km S. Ankara leg. Warncke’
(JLR).

Description

Female. — Head: Antennae bright orange, antenna
3 rather broad in outline, apex subacute, upper mar-
gin straight, arista long, suddenly tapering, antennal
ratio 2.8; pubescence golden; lustrous midstripe oc-
cupying 1/3 of width, ocellar region showing dark

Hurkmans: Monograph of Merodon

pubescence; ocellar angle 70°, lateral ocelli larger than
central ocellus; pubescence on compound eyes rather
thin, whitish.

Thorax: Rather narrow, black with weak bluish
lustre on katepisternum, anepisternum, anepimeron,
lateral scutellum and dorsum, with four longitudinal
yellow pruinose bands on dorsum, coalescent anteri-
orly; pubescence rather short, erect, uneven, golden
yellow, but darker, brownish in interalar band. Wings
clear; halteres, squamae and antisquamae pale yellow.

Legs: All orange except coxae and trochanteres
brownish; femora 3 swollen, rather straight, bearing 6
bristles on normal triangular processus.

Abdomen: Rather slender, dark with micaceous
lustre; T II bearing large orange lateral spots, T II-IV
with slightly arcuate, moderately wide, interrupted,
golden pruinose bands combined with golden pubes-
cence; pubescence concolorous otherwise, but yellow
laterally and on posterior fringe of T III-IV; T V far
projecting, rounded posteriorly.

Body length 11 mm.

Male. — Unknown.

Diagnosis. — M. aureotibia can be distinguished
from all other females of Merodon by its extensively
orange leg coloration.

Period of flight and distribution (fig. 106). — This
species occurs in June in central and south-eastern

Turkey.

Discussion. — In view of the unique leg coloration,
the narrow thorax and striking golden pubescence on
especially head and abdomen aureotibia has been de-
scribed as a new species. It is reminiscent of females of
aberrans but differs from it by the leg colour, shape
and colour of the antennae, narrower thorax and con-
spicuous orange lateral spots on T II. Moreover the
femora 3 in aureotibia are more swollen.

In the coloration of the.legs the females of aureoti-
bia resemble vandergooti, known only after males.
Reasons for not considering aureotibia as the female
sex of vandergooti are: (1) aureotibia has obtuse anten-
nae, orange and slightly swollen hind femora, arcuate
pruinose bands not obliquely placed and a serrate dis-
tal triangular processus margin with 6-7 bristles, (2)
vandergooti has acute antennae, dark and enormously
swollen hind femora, oblique bands and a hardly ser-
rate distal triangular processus bearing 10-13 bristles.

These differences are far too great and of a wrong
type to be considered as sexual dimorphism.

In view of the slender habitus, relatively long an-
tenna 3 with suddenly tapering arista, aureotibia
might possibly be related to the longicornis group.
However, none of the species in that group has

203

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

orange legs and relatively wide pruinose bands,
whereas aureotibia lacks the typical crimson abdomi-
nal colour of the longicornis group.

Merodon viaticus Fabricius

Merodon viaticus Fabricius, 1805: 197. Lectotype, sex un-
known (here designated): ‘viaticus’, [the upper half of a
thorax with basal wing fragments attached] in Sehestedt
& Tonder Lund coll. (ZMUC) [examined].

Merodon viaticus Wiedemann in Meigen 1822: 364;
Macquart 1828: 296; Rondani 1845: 258; Walker 1849:
598; Schiner 1857: 414.

Lampetia viatica, Sack 1931: 327.

Material examined. — Paralectotype, sex unknown
‘viaticus [part of a thoracic dorsum] in Kieler coll. (ZMUC).

Description

The size of the thoracic dorsum indicates an overall
size of about 12 mm. The background coloration is
dark, pubescence largely rubbed off, but where remai-
ning, erect and whitish; pruinose bands rather clear.
Wing fragments clear, brown veined.

Discussion. — The original description by Fabricius
(1805) does not mention the sex of the fly described.
As however a dark median line on the frons is men-
tioned it must have been a female. The habitus
should be similar to that of annulatus which is a syno-
nym of natans. Stated differences with viaticus are the
all dark abdomen in natans (with a red base in wiati-
cus), the all dark legs and smaller size, averaging 9 mm
for natans.

Wiedemann (in Meigen 1822) gives an extensive
description of viaticus, in which the abdomen is said
to be red on T I II and anterior T III; size is given by
Wiedemann as 5 to 6 lines, i. e. 10.5 to 12.2 mm.
Wiedemann’s description is considered to include
both sexes as he comments on differences in the
colour of antennae and tarsi between sexes.

Some specimens from Sicily, leg. Schigdte, labelled
‘viaticus’, without any further data (ZMUC) have been
re-identified as elegans sp. n.; differences with the de-
scription of viaticus are the size (14 mm), much swol-
len femora 3 (not mentioned in either of the descrip-
tions); legs yellow, not reddish or rusty brown-yellow
as in the descriptions; abdomen dark brown with ex-
tensive yellow coloration, not red, orange or purplish
as in descriptions.

M. viaticus is assigned a place in the section of
Merodon containing the species with arcuate pruinose
bands since these are mentioned in the original de-
scription.

As there are no females of species in this section of
Merodon with a cupreous lustre on the abdomen and
at the same time partially dark, partially orange legs,

204

viaticus cannot be synonymized with any of the spe-
cies. It was listed by Sack (1931: 327) as a synonym of
spinipes (=avidus); no reasons for this synonymy were
given.

ANNOTATED LIST OF SPECIES DISCUSSED IN THIS
PAPER

Abbreviations used to describe status: H =
Holotype examined; ? = Type material not located,
not examined; S = syntype(s) examined; P = paraty-
pe(s) examined; L = lectotype examined (here de-
signated); PL = paralectotype(s) examined.

Deposition Status Ann
distinctus assemblage
biarcuatus Curran, 1939 AMNH H =
clunipes Sack, 1913 ZMHB =
distinctus Palma, 1863 ? ? 1
dimorphus (Szilady,

1940) syn. n. TMA [lost] 2
femoratus Sack, 1913 ? S? 3
maride sp. n. ZMAN H, P =
ottomanus sp. n. ZMAN H, P =
testaceoides sp. n. RMNH, ZMAN H,P =
alexeji group
alexeji Paramonov, 1925 PC [lost] 4
altinosus sp. n. ZMAN H
hirsutus Sack, 1913 NHMW L, PL =
hypochrysos sp. n. ZMAN H, P =
kawamurae
Matsumura, 1916 ? 5

micromegas (Hervé-

Bazin, 1929) syn. n. MNHN, BMNH L, PL 5

marginicornis sp. N. ZMAN H
rufitarsis Sack,

1913 stat. nov. ZMHB H =
sophron sp. n. BMNH H =
tener Sack, 1913 DEIC IL, JBL, 6
trizonus (Szilady, 1940) TMA [lost] 2
tarsatus group
ankylogaster sp. n. RMNH lal, IP
auronitens sp. n. ZMAN H -
caudatus Sack, 1913 FIS [lost] 3
oidipous sp. n. ZMAN H, P =
persicus sp. n. RMNH H =
tangerensis sp. N. MNHN H =
tarsatus Sack, 1913 ZMHB H =

smirnovi Paramonov,

1926 syn. n. PC [lost] 4
turkestanicus Paramonov,

1926 PC [lost] 4
xanthipous sp. n. ZMAN H, P =
clavipes group
aberrans aberrans Egger,

1860 NHMW IE IRL, =

knerii Mik, 1867 NHMW? S? =

obscuripennis Palma,
1863
aberrans ssp. flavitibius
Paramonoy, 1925
aberrans ssp. isperensis
ssp. n.

brevis Paramonov, 1925
clavipes (Fabricius, 1781)

canipilus Rondani,

1865

clauda (Villers, 1789)

curvipes (Gmelin,

1790)

gravipes (Rossi, 1790)

sacki Paramonov,
1937 syn. n.
senilis Meigen, 1822
clavipes var. albus
Paramonov, 1927
clavipes var. ater
Paramonov, 1927
clavipes var. niger
Paramonov, 1927
cupreus sp. n.
dzhalitae Paramonov,
1926
hamifer Sack, 1913
karadaghensis Zimina,
1989
lusitanicus sp. n.
quadrinotatus
(Sack, 1931)
splendens sp.n.
velox Loew, 1862
velox var. anathema
Paramonov, 1925
velox var. armeniacus
Paramonov, 1925
warnckei sp. n.

pruni group

pruni (Rossi, 1790)

fulvus Macquart, 1834

fuscinervis
Von Réder, 1887

pallida Macquart, 1842
sicanus Rondani, 1845

pruni var. obscurus

Gil Collado, 1929

longicornis group

erivanicus Paramonov,
1925

kaloceros sp. n.

longicornis Sack, 1913

vandergooti group
vandergooti sp. n.

nigritarsis group

femoratoides (Paramonov,

1925) comb. n.

nigritarsis Rondani, 1845

PC
ZMAN, JLR

PC
NHMW, DEIC

Moscow
ZMAN, JLR

PC

PC
ZMAN, JLR

PC
ZMAN, JLR
NHMW

ZMAN, JLR

ZMHB
LSF

Hurkmans: Monograph of Merodon

avidus group
avidus (Rossi, 1790) ZMHB [lost] 9
spinipes (Fabricius,

1794) ZMUC IL il

serrulatus Wiedemann

in Meigen, 1822 NHMW It =

italicus Rondani, 1845 LSE L, PL =

rufitibius Rondani,

1845 LSF L, PL =

graecus Walker, 1852 BMNH IL =

aurifer Loew, 1862 ZMHB IL -
quadrilineatus Lioy,

1864 nom. nud. = 12
crassifemoris group
crassifemoris Paramonov,

1925 comb. n. PC [lost] 4, 13
elegans group
bequaerti sp. n. MNHN H, P =
elegans sp. n. ZMAN e. a. IND 14
manicatus (Sack, 1938) | HLMD EIND, =
testaceus Sack, 1913 NHMW L, PL =
alagoezicus group
alagoezicus Paramonov,

1925 PC [lost] 4
lucasi sp. n. ZMAN, JLR EUD -
nitidifrons sp. n. ZMAN H
satdagensis sp. n. ZMAN, JLR lial, Je -
schachti sp. n. ZMAN, JLR H,P =
taniniensis sp. n. ZMAN, JLR lal, IP -
toscanus sp. n. ZMAN H =
Remaining species
affinis Gil Collado, 1930 IEE H =
aureotibia sp. n. ZMAN H =
viaticus (Fabricius, 1805) ZMUC L, PL 15

Annotations

1. The types from the Palma collection have not
been located. The descriptions and drawings provi-
ded by Palma (1863) of obscuripennis and distinctus
are extensive enough to be used for diagnostic purpo-
ses. Sack (1931) has been followed in considering 0b-
scuripennis synonymous with aberrans.

2. The type-material, formerly at TMA, is lost (Dr.
A. Dély-Draskovits, in litt.).

3. The holotypes of caudatus and quadrinotatus,
and the syntypes of M. femoratus, all formerly in
Sack’s private collection, were bequeathed to the
Forschungsinstitut Senckenberg, Frankfurt/ Main.
According to Dr. W. Tobias, curator of Syrphidae in
the said museum, no Sack types of any Merodon are
currently there. When Sack left his collection to SMF
the material was presumably still stored in his own
house which was plundered shortly after World War
II. All Sack types of Merodon left to the Senckenberg

Museum are therefore to be considered lost.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

4. Many type-specimens from the S. J. Paramonov
collection have not been located. According to Liepa
(1969) most of the types originally in that collection
are lost. Some material which probably originated
from that collection has been found in ZMHB, includ-
ing type-material of Merodon femoratoides and M.
crassifemoris. Dr O. G. Ovchinnikova of the USSR
Academy of Sciences (St. Petersburg) informed me
that some types from the Paramonov collection might
still be preserved at the Kiev Zoological Museum.
This is also the view of Dr. Y. Nekrutenko, Kiev
(pers. comm.). Regrettably this cannot be confirmed.
The type of M. smirnovi should be stored in the
Zoological Institute, Moscow University but this also
could not be confirmed.

Reliably identified material, in some cases det. S. J.
Paramonov, was present of the following species: ala-
goezicus (ZMAN); alexeji (KBIN); brevis (KBIN); crassif-
emoris (ZMHB); erivanicus (ZMAN); femoratoides
(ZMHB) and turkestanicus (ZMAN). No material which
can be reliably assigned to smirnovi is known; in view
of its description this species is here considered syn-
onymous with zarsatus.

5. The deposition of the type(s) is not mentioned
in Matsumura’s original publication. After Horn &
Kahle (1936) the Matsumura collection is at the
University of Sapporo. Following enquiry to the de-
partment of Entomology, Hokkaido University,
Sapporo, the presence of the types of M. kawamurae
there cannot be confirmed. The type(s) (sex and
number unknown) originate from Kumamoto,
Kyushu. However, this is the name of both a city and
an extensive district. The description of the species
presented here is based on 29, 2d from Harima [in-
land sea of Japan at 34° 30° N, 134° 30° E], in BMNH
and fully complying with the original description.

6. Only female type-specimens have been exam-
ined, no male types have been found. The male from
Saratow, formerly preserved at TMA, is lost (Dr. A.
Dély-Draskovits, in litt.). The male and female syn-
types from the Becker collection are not at ZMHB and
have not been located elsewhere. A male type from
Uralsk, stated in the original publication to be in
NHMW, was not found there during a visit in 1987.
The 2 female syntypes from which the lectotype has
been designated are stored in DEIC, Eberswalde.

7. This species has been described without mention
of material, although it is stated by Villers (1789) that
Musca clauda occurs in Europe, in Bressia [? =
Brescia].

8. Musca curvipes Gmelin is directly referred to
Fabricius’ original description of Syrphus clavipes by

206

Gmelin (1790). Therefore type-material of this taxon

does not exist.

9. No type-material of Syrphus gravipes, Syrphus
pruni or Syrphus avidus has been located. After Horn
& Kahle (1936) the Rossi collection was transferred
to ZMHB; after Dr. H. Schumann (in litt.) no Rossi
types of Merodon are currently present there.

10. No material was examined of fulvus Macquart
or fuscinervis Von Réder. Both original descriptions
suggest a great similarity to pruni. Both species were
listed by Peck (1988) as synonyms of pruni. This syn-
onymy has been followed.

11. The type-material of spinipes consists of a sin-
gle, quite completely decayed probable syntype in
ZMUC. This taxon is here synonymized with avidus on
the ground of complete compliance of the descrip-
tions and figures throughout the literature, starting
from the first publications. It must of course be borne
in mind that Rossi or Fabricius may have been de-
scribing any species of the large group externally re-
sembling avidus.

12. No type-material of this taxon was probably
ever in existence, as Lioy, after Horn & Kahle (1936)
did only theoretic systematic work.

13. The holotype has not been located (see annota-
tion 1). In ZMHB a specimen dating from 1929, with
a label attached marking it as a type, was found. As
this specimen was collected after the publication of
var. crassifemoris it is not available as a type specimen;
however the labels are in Paramonov’ s handwriting.
The specimen fully complies with the description of
the var. crassifemoris. It is therefore available as a
check on identifications.

14. A large number of paratypes spread over many
institutions was designated to facilitate genital study.
This was done since elegans externally resembles avi-
dus and could well be confused with it.

15. The two syntypes from ZMUC, one from the
Kieler coll. and one from the Sehestedt and Tonder
Lund coll., are so seriously damaged that no conclu-
sion can be drawn as to the status of this species. No
other material identified as viaticus and complying
with the description of that species has been found.
Therefore the descriptions given are the only base for
identification.

ANNOTATED LIST OF SPECIES EXCLUDED FROM
MERODON

The status of the species described by Walker
(1849, 1852, 1857a, 1860) is not clear in all cases. As
not all material was available, the specimens studied
have been referred to as type-specimens. The present
author does not intend to designate lectotypes of spe-
cies excluded from Merodon.

Merodon angustiventris Macquart, 1855: 90.
Lectotype d (designated by Thompson 1988)
(BMNH) [examined] not considered to belong to
Merodon in view of morphology of femur 3 and wing
venation. According to Thompson (1988) the lecto-
type belongs to Quichuana angustiventris (Macquart),
a species occurring in the Neotropical region.

Merodon balanus Walker, 1849: 600. A d type-
specimen (BMNH) [examined] belongs to the genus
Mallota in view of femur 3 and wing venation.
Specimen originates from New York [Nearctic re-
gion, U. S. A.] where a natural occurrence of
Merodon is unknown.

Merodon bardus Say (in Walker 1849: 598). The d
type-specimen [examined] does not belong to
Merodon. After Osten-Sacken (1878: 135) this speci-
men belongs in Mallota.

Merodon bautias Walker, 1849: 600. Two syn-
types, d and $ (BMNH) [examined]. Both syntypes
originate from Georgia, United States. The facial
knob and wing venation are not compatible with
Merodon. After Osten-Sacken (1878: 135) this spe-
cies belongs in Mallota.

Merodon bicolor Walker, 1852: 243 [type-material
not examined]. In the original description ‘feelers’
[interpreted as antennae] are stated to possess six
joints, therefore the species is considered not to be-
long to the Syrphidae. The type-material originated
from Brazil [Neotropical region] where Merodon spe-
cies do not naturally occur.

Merodon bipartitus Walker, 1849: 599. The @
type-specimen (BMNH) originates from Georgia,
United States [examined]. In view of the wing vena-
tion, this species probably belongs to Mallota. See
Osten-Sacken (1878: 135).

Merodon chiragra Fabricius, 1805: 198 [type-mate-
rial not examined]. Not included in a shipping of all
other available Merodon types from ZMUC; this species
belongs in Sphegina.

Merodon coerulea Becker, 1912: 602 [type-material
not examined]. On the basis of the original descrip-
tion this species belongs in Eumerus.

Merodon contrarius Walker, 1849: 599. Both d
and © type-specimens (BMNH) [examined]. This ma-
terial originated from New Holland and belongs to
Orthoprosopa grisea (Walker).

Merodon edentulus Macquart, 1855: 90. Holotype,

Hurkmans: Monograph of Merodon

? from Cape of Good Hope [not examined]; a 2
non type (BMNH) dating from XIX century identified
as edentulus [examined] belongs to Mallota. The orig-
inal description mentions a facial knob, unknown in
Merodon.

Merodon interveniens Walker, 1860: 120. 3 Type-
specimen (BMNH) [examined]. This specimen origi-
nated from Macassar [Sulawesi] and does not belong
in Merodon in view of wing venation, shape of leg 3
etc.

Merodon melota Seguy, 1941: 13. 2 Holotype
from Agadir, Morocco (MNHN) [examined]. This spe-
cies belongs to Eumerus.

Merodon morosus Walker, 1849: 599. A 2 type-
specimen (BMNH) [examined]. This does not belong
to Merodon but according to Osten-Sacken (1878:
135) 1s Polydontomyia curvipes O.-S.

Merodon ornatus Brunetti, 1915: 232. Holotype in
Indian Museum, Calcutta [not examined] could not
be shipped in view of its conservation state. After
original description the wings have a brown stigma,
there is no triangular processus on the femur 3 and
the face is projecting; therefore this species is consid-
ered not to belong to Merodon.

Merodon scutellaris Shiraki, 1968: 200. [Type-ma-
terial not examined]. From the description and pho-
tographed holotype it can be concluded that this spe-
cies belongs to the genus Azpeytia.

Merodon tenebricus Walker, 1849: 601. Lectotype
9 (designated by Thompson 1988) (BMNH) [exam-
ined]. In view of the shape of femur 3 and wing vena-
tion this species does not belong to Merodon. The
identification as Palpoda furcata was given by
Thompson (1988: 220).

Merodon torpidus Walker, 1857a: 152. Lectotype
d (designated by Thompson 1988) (BMNH) [exam-
ined]. This does not belong to Merodon given the
shape of femur 3 and wing venation. The identifica-
tion as Orthoprosopa grisea Walker was given by
Thompson (1988: 220).

Merodon tuberculatus Brunetti, 1923: 214. The
types, in the Indian Museum, Calcutta, [not exam-
ined] were too fragile to be shipped; the description
of this species was based on that of Helophilus tuber-
culatus. The description and the figures (Brunetti,
1908) make clear that this species does not belong to
Merodon.

Merodon umbrifer Walker, 1849: 601. The d type-
specimen from Sierra Leone does not belong to
Merodon in view of leg morphology and wing vena-
tion.

Merodon varicolor Walker, 1857b: 122. The &
type-specimen from Borneo, in BMNH, belongs to
Mallota in view of the shape of femur 3 and the wing
venation pattern.

207

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Proposal of Azpeytia shirakii nomen n. for
Merodon scutellaris Shiraki, 1968

Since Merodon scutellaris Shiraki is incorrectly as-
signed to Merodon (see above), it is here proposed to
transfer this taxon to Azpeytia, where it clearly be-
longs. As there already exists an Azpeytia scutellaris,
published by Walker (1857b: 113), the epitheton of
the species described by Shiraki, when transferred to
Azpeytia, must be changed; the nomen novum
Azpeytia shirakii is hereby proposed for this taxon.

ACKNOWLEDGEMENTS

I would like to thank Mr Volkert van der Goot for
stimulating my work on the genus Merodon, his con-
tinued interest in my studies, and the translation of
the Zimina paper. This work would have been hardly
possible without the generous co-operation of Mr Jan
A. W. Lucas who kindly placed his extensive collec-
tion of Merodon specimens at my disposal. My warm
thanks go to Dr. Pjotr Oosterbroek who posed
questions to increase my curiosity, was always willing
to discuss ideas, stimulated the general progress of my
studies, and critically read the various drafts of the
manuscript. The facilities and advice provided by Dr.
H. W. J. van Dijk were of great value in producing
the final manuscript. I wish to thank Prof. Dr. J.
Stock, Dr. P. Oosterbroek, Dr. A. C. Pont, London,
Dra. M. A. Marcos Garcia, Salamanca, Mr C.
Claussen, Flensburg, Mr. J. A. W. Lucas, Rotterdam,
Dr. H. W. J. van Dijk, Zwolle, Mr. G. den
Hollander, IJsselstein, Mr. P. L. H. Hanegraaf and
Mrs. R. L. G. J. van der Voort, Houten, and Mr. V.
S. van der Goot, Amsterdam for their valuable com-
ments at the various stages of the work on Merodon,
or discussion of my research in general. I would like
to thank the persons already mentioned, Mrss.
Marcos Garcia and Petanidou, and Messrs. Barke-
meyer, Bausenwein, Ellis, Hauser, Lyneborg, Toth,
Treiber, Van Ooijen, Vujic and Wyatt for the inte-
resting material provided for examination.

Financial support for museum and field work was
gratefully received from the Uyttenboogaart-Eliasen
Stichting, Amsterdam.

My sincere gratitude is due to the following per-
sons for permission to study preserved specimens
from the collections in their care, and assistance in
tracing the type-specimens of various species:

Dr. Luca Bartolozzi, Firenze; Dr. Vlad. Bradescu,
Bucuresti; Mr. Claus Claussen, Flensburg; Dr. Ruth
Contreras-Lichtenberg, NHMW, Wien; Dr. M. Datta,
Calcutta; Dr. A. Dély-Draskovits, TMA, Budapest;
Dr. H. Feustel, HLMD, Darmstadt; Dr. R. Gaedike,
DEIC, Eberswalde -Finow; Dr. D. Grimaldi, AMNH,
New York; Dr. Patrick Grootaert, KBIN, Brussel; Dr.
P. J. van Helsdingen, RMNH, Leiden; Mr. Gijs Den

208

Hollander, IJsselstein; Prior P. Bruno Hubl, Bsa,
Admont; Dr. I. Izquierdo, IEE, Madrid; Dr. F.
Kaplan, Tel Aviv; Dr. C. Leonardi, MCSN, Milano;
Dr. Leif Lyneborg, ZMUC, Kobenhavn; Dr. Maria-
Angeles Marcos Garcia, SAL, Salamanca; Dr. Carolina
Martin, IEE, Madrid; Mrs. Sarah Mascherini, LSF,
Firenze; Dr. L. Matile, MNHN, Paris; Dr. E. Mingo
Pérez, IEE, Madrid; Mr. Peter van Ooyen, Utrecht;
Dr. O. C. Ovchinnikova, St.Petersburg; Ir. Dirk
Prins, Rhenen; Dr. Wolfgang Schacht, ZSBM,
München; Dr. H. Schumann, ZMHB, Berlin; Dr. P.
Tschorschnig, MENS, Stuttgart; Mr. Nigel P. Wyatt,
BMNH, London. Also I wish to thank Dr. W. Tobias,
Frankfurt/ Main and Dr. ©. G. Ovchinnikova, St.
Petersburg for their help in tracing the Sack and
Paramonov type-specimens, and Dr. Dely-
Draskovits, Budapest for information on the TMA
specimens.

Last but not least I wish to express my gratitude to-
wards my wife Marijke whose unstinting interest in
my work greatly contributed to the result.

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210

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HURKMANS: Monograph of Merodon

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Westwood, J. O., 1840. An introduction to the modern
classification of insects: Synopsis of the genera of British
Insects, Order 13: Diptera Aristotle (Antliata Fabricius,
Halteriptera Clairv.). — pp. 125-158.

Zimina, L. V., 1989. Novie Sirfidy Poda Merodon (Diptera,
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nus Merodon (Diptera, Syrphidae) from the east Crimea]
— Vestnik Zoologia 1989 no. 1: 24-29. (in Russian).

Received: January 1992.
Revised version accepted: June 1993.

Figs. 1-10. General morphology of Merodon. — Fig. 1. Lateral view of head. a, face; b, fions; c, oral margin; d, vertex; e, occi-
put. — Fig. 2. Lateral view of antenna. a, antenna with 3rd article showing convex upper margin, apex rounded; b, 3rd article
with upper margin basally convex, apically concave, apex acute; c, 3rd article straight margined, apex subacute; d, 3rd article
strongly widened, upper margin convex, incised subapically. Lines above and below mark measured lengths of articles 2 and
3 respectively; the antennal ratio (length 3 : length 2) is often used in descriptions. — Fig. 3. Dorsal view of vertex and frons
in d. Line portion a marks the length of tl, portion b the vertex height; angle alpha is vertex angle, beta is ocellar angle. — Fig.
4. Dorsal view of vertex and frons. (left), tl complete; (middle), tl incomplete: a linear strip separates the eyes, this condition
might be termed subholoptic; (right), eyes separate (this occurs in some species in the d, and in all 9). The ocellar region is
hatched. — Fig. 5. Lateral view of thorax (wings and legs omitted). 1, humeral knob; 2, dorsum with suture; 3, scutellum; 4,
mesopleuron (katepisternum); 5, pteropleuron (anepisternum); 6, sternopleuron (anepimeron); 7, coxa I; 8, coxa II; 9, coxa
III; W, wing insertion area; R, respiratory orifice (trachaea). — Fig. 6. Dorsal view of right wing. 1, anal cell (a); 2, discal me-
dial cell (dm); 3, combined 4th+5th radial ce | (r4+5); 4, basal medial cell (bm); 5, basal radial cell (br); 6, 2nd+3rd radial cell
(r2+3); 7, 1st radial cell (rl); 8, subcostal cell (sc); 9, costal cell (c); A, Costa (C); B, Subcosta (Sc); C, Ist branch of Radius
(R1); D, main branch of Radius (R); E, 2nd & 3rd fused branches of Radius (R2+3); F, 4th & Sth fused branches of Radius
(R4+5); G, Media (M); H, anterior Cubital (Cual); I, discal-medial Cubital (dm-cu); J, apical part of Media (M); K, analis
(A); L, humeral (h); M, radial-medial (r-m); N, 2nd anterior Cubital (Cua2); O, basal part of Cual; P, basal-medial Cubital
(bm-cu). — Fig. 7. Lateral view of leg III. a, trochanter (spine shown is present in d of some species only); b, femur, with c,
triangular processus; d, tibia, with e, apical spur; f, metatarsus; g, tarsus. — Fig. 8. Ventral view of d genitalia (alagoezicus
group). A, cercus; B, posterior surstyle lobe; C, apical margin of surstyle; D, ridge on inner face (medial face) of surstyle; E,
epandrium; F, membraneous part of surstyle; G, apical extension of anterior surstyle lobe; H, anterior surstyle lobe; J, acces-
sory lobe. — Fig. 9. Lateral view of & genitalia (alagoezicus group). A through J as in fig. 8; L, aedeagus; M, genital cap. — Fig.
10. Ventral view of aedeagus. A, apical shaft part; B, apex aedeagi; C, fringed plates; D, chitinous plates; E, subapical cavity;
F, thecal ridge basally bordering same; G, basal humps.

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Figs. 11-13. Aedeagus in Merodon. — 11, lateral view, lettering as in fig. 10, H, lateral ridge; 12, slender aedeagus with length-
ened apical shaft part A; S, paired spines; 13, stout aedeagus with recumbent fringed plates. — Figs. 14-17. Lateral view of 3
genitalia of Merodon, lettering as in Fig. 9. — 14, M. apimimus; 15, M. distinctus, 16, M. nigritarsis, 17, M. longicornis. — Fig.
18. Lateral view of head in d of different Syrphidae. — a, Azpeytia scutellaris Walker; b, Merodon avidus, c, Eumerus strigatus

Fallen; d, Mallota takasagoensis (Matsumura); e, Platynochaetus armipes Bezzi.

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Hurkmans: Monograph of Merodon

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Fig. 19. Dorsal view of right wing in d of different Syrphidae. — a, Eumerus strigatus Fallen; b, Azpeytia scutellaris Walker; c,
Mallota takasagoensis (Matsumura); d, Merodon avidus e, Platynochaetus armipes Bezzi. — Fig. 20. Lateral view of leg III in d
of different Syrphidae. — a, Merodon equestris, b, M. armipes, c, M. minutus, d, M. aberrans, e, M. ruficornis, f, Eumerus sulci-
tibius Rondani; g, E. tuberculatus Rondani; h, E. olivaceus Loew and E. nudus Loew; j, Eumerus strigatus Fallen; k, Azpeytza
scutellaris Walker; |, Azpeytia shirakii nomen novum; m, Mallota cimbiciformis Fallen; n, Mallota takasagoensis (Matsumura);
o, Platynochaetus armipes Bezzi; p, Platynochaetus rufus (Macquart).

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Fig. 21. Merodon biarcuatus. a, b, variation in antennal shape; c, distal end of f3. — Fig. 22. M. clunipes. a, lateral view of d
genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 23. M. distinctus. a, lateral view of 3 genitalia; b, c, ventral and lat-
eral view of aedeagus; d, habitus of 3. — Fig. 24. M. femoratus. a, b, ventral and lateral view of aedeagus; for lateral view of
genitalia compare fig. 22a. — Fig. 25. M. mariae. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus.

216

Hurkmans: Monograph of Merodon

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Fig. 26. Merodon ottomanus. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 27. M. testaceoides.
a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 28. M. alexeji. a, lateral view of d genitalia; b,
ventral view of aedeagus. — Fig. 29. Posterior end of surstyle and cercus in M. alexeji (a) and M. altinosus (b). Pubescence de-
leted to show difference in outlines. — Fig. 30. M. hirsutus. a, lateral view of & genitalia; b, c, ventral and lateral view of ae-
deagus. — Fig. 31. M. hypochrysos. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Fig. 32. Merodon kawamurae. a, lateral view of d genitalia; b, ventral view of overlapping posterior surstyle lobes; c, d, ven-
tral and lateral view of aedeagus; e, size correlated shape variation in antennae. — Fig. 33. M. marginicornis. a, lateral view of
g genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 34. M. rufitarsis. a, lateral view of & genitalia; b, c, ventral and

lateral view of aedeagus. — Fig. 35. M. sophron. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus; over-
lap of surstyle lobes as in Fig. 32b.

218

Hurkmans: Monograph of Merodon

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Fig. 36. Merodon ankylogaster. a, lateral view of d genitalia; b, lateral view of posterior abdomen: T III, IV = tergites III and
IV, S III, IV = sternites III and IV, A = appendage to S IV, G = genitalia. Pubescence on genital cap deleted. — Fig. 37. M.
auronitens. a, lateral view of & genitalia; b, apex aedeagi; c, anterior view of apical end of tibia 3 and basitarsus 3. — Fig. 38.
M. caudatus. a, lateral view of ¢ genitalia; b, c, ventral and lateral view of aedeagus; d, posterior view of distal tibiae 3 and
metatarsi 3. — Fig. 39. M. oidipous. a, lateral view of d' genitalia; b, ventral view of aedeagus; c, lateral view of posterior abdo-
men (lettering as fig. 36b); d, posterior view of metatarsi 3 and other tarsi 3.

219

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Fig. 40. Merodon persicus. a, lateral view of 3 genitalia; b, lateral view of posterior abdomen; c, posterior view of distal tibiae
3 and metatarsi 3, showing specialized hair-tuft on hump near base of metatarsi 3; d, lateral view of same hump, surrounding
pubescence deleted. — Fig. 41. M. tangerensis. a, lateral view of d genitalia; b, ventral view of aedeagus; c, see 40c; d, lateral
view of posterior abdomen. Pubescence on genital cap deleted. — Fig. 42. M. tarsatus. a, lateral view of & genitalia; b, c, ven-
tral and lateral view of aedeagus; d, posterior view of distal tibiae 3 and metatarsi 3 showing specialized pubescent area; e, lat-
eral view of posterior abdomen.

220

Hurkmans: Monograph of Merodon

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Fig. 43. Merodon turkestanicus. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus; d, posterior view of dis-
tal tibiae 3 and metatarsi 3 (note extremely long floccus); e, lateral view of posterior abdomen. — Fig. 44. M. xanthipous. a, lat-
eral view of & genitalia; b, lateral view of posterior abdomen; c, posterior view of distal tibiae 3 and metatarsi 3 (note long
floccus). — Fig. 45. M. aberrans. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus; d, outline of abdo-
men, d of aberrans aberrans (specimen of average size); e, same in aberrans isperensis. — Fig. 46. M. brevis. a, lateral view of d
genitalia; b, c, ventral and lateral view of aedeagus.

22,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Fig. 47. Merodon clavipes, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus; d, M. splendens, lateral view of
d genitalia. — Fig. 48. M. cupreus. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 49. M. ha-
mifer. a, lateral view of distal end of femur 3 and base of tibiae 3 in d ; b, same in 2. Fig. 50. M. lusitanicus (2). a, outline
of femur 3; b, same in ® M. aberrans. — Fig. 51. M. quadrinotatus (® ). Dorsal view of abdomen, hatched area represents black
pubescence; dotted, grey pubescence; blank, white pubescence. — Fig. 52. M. warnckei. a, lateral view of d genitalia; d, detail
of same at different angle showing projection bearing pubescence on margin of posterior surstyle lobe; c, b, ventral and later-
al view of aedeagus. — Fig. 53. M. pruni. a, lateral view of 8 genitalia; b, ventral view of aedeagus; c, d, lateral view of same in
small and large specimen (note size correlated morphology).

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Hurxmans: Monograph of Merodon

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N
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Fig. 54. Colour patterns on abdomina of females: a, b, Merodon erivanicus (showing extremes of variation); c, M. longicornis,
d, M. kaloceros. — Fig. 55. Outlines of antennae of females: a, M. erivanicus, b, M. longicornis, c, M. kaloceros. — Fig. 56.
Merodon kaloceros. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus. - Fig. 57. M. longicornis. a, lateral
view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 58. M. vandergooti. a, lateral view of d genitalia; b, c,
ventral and lateral view of aedeagus. — Fig. 59. M. femoratoides. a, lateral view of 3 genitalia; b, c, ventral and lateral view of
aedeagus; d, outline of trochanter 3 in lateral view.

223

TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

po 4

Se

SE

NINA
U
À N DIR, Ah
EE TÉLÉ
An ES
Pa | 60
= mm A a Sz
peser it Tn x <

wat

AU Mi
fs Ta AN, seo

= PS
= = LS

Di TAN £
Party | ; ES 60b

n È
LEER SS

Fig. 60. Merodon nigritarsis. a, lateral view of d genitalia (lectotype); b, same from specimen from Handere, Turkey; c, d, ven-
tral and lateral view of aedeagus (lectotype); e, f, same (Handere); g, variation of shape in outlines of trochanter 3. — Fig. 61.
M. avidus. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus; d, ventral view of d genitalia; e, abdomi-
nal colour pattern in pale 3; f, same in dark d; g, same in intermediate 2. — Fig. 62. M. crassifemoris. a, lateral view of d
genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 63. M. bequaerti. a, lateral view of & genitalia; b, c, ventral and lat-
eral view of aedeagus.

224

Hurkmans: Monograph of Merodon

22 dl D
a
= =

ZA = BET Fe

Pf

Fig. 64. Merodon elegans. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 65. M. manicatus. a,
lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 66. M. testaceus. a, lateral view of 3 genitalia; b,
©, ventral and lateral view of aedeagus. Pubescence on genital cap deleted. — Fig. 67. Distal ends of tibia 3 in: a, M. alagoezi-
cus, b, M. schachti, c, M. satdagensis, d, M. lucasi. — Fig. 68. M. alagoezicus a, lateral view of 3 genitalia; b, c, ventral and lat-
eral view of aedeagus; d, lateral view of triangular processus of femur 3.

225

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

he
vi |

tr,

SN +
ET TIZIA N

Sy

€

I
(y
2

S
N
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TER
Dy i}

Fig. 69. Merodon lucasi. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus d, lateral view of triangular pro-
cessus on femur 3.. — Fig. 70. M. nitidifrons. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus; — Fig. 71.

8
M. satdagensis. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus.

226

HURKMANs: Monograph of Merodon

at

ig, ==

i @ |
Sat
— Al N

ZB il Ps x

N)

\

FM

net Im es 73a u

N
N

N

€ 7

RES a 75

fo +

Fig. 72. Merodon schachti. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 73. M. taniniensis. a,
lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 74. M. toscanus. a, lateral view of 3 genitalia; b, c,
ventral and lateral view of aedeagus. — Fig. 75. M. aureotibia. a, outline of antenna.

227

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Vi we ANN

Fig. 76-79. Distribution of Merodon in biogeographic regions, hatched area: known conjunct range, dot or triangle: known
localities. — 76, Ethiopian region; 77, western Palaearctic; 78, south-western Asia. — Fig. 79. Distribution of Merodon kaw-
amurae, the only species with a certain natural occurrence in eastern Asia. — Fig. 80. Localities of Merodon in Turkey men-
tioned in this paper: 1, Izmir Smyrna’). 2, Bursa [’Brussa’]; Karacabey lies slightly to the W, on the coast. 3, Ankara. 4,
South of Ankara, near Ankara airport. 5, Aksehir [Ak-chehir]. 6, Konya. 7, Irmesan gedigi N of Akseki. 8, Sarihaci, 45 km
W of Malatya. 9, Malatya [Melatia]. 10, Altı Haral Gölü NE of Celikhan. 11, Nemrut Dag near Adiyaman. 12, Buglan
gecidi

(pass) near Bingöl. 13, Palandöken Dagi (Kayak) SW of Erzurum. 14. Ovit pass, near Ispir. 15, near Karaurgan, 8 km W of
Sarıkamıg. 16, Soganli railroad stop. 17, Handere, 20 km W of Sarikamis. 18, 11 km E of Karakurt. 19, Kars Deresi valley
5 km E of Sarıkamıs. 20, 20 km W of Uludere. 21, Akdamar, 40 km SW of Van. 22, Suvarihalil pass (on main east-west road
through Hakkari province). 23, Mengene Dagi, N of Baskale. 24, Tanin-Tanin pass (on main east-west road through
Hakkari province), W side. 25, Habero deresi valley, north of same road. 26. Tanin-Tanin pass near top. 27, Sat Dagları N
of Mount Gavaruk. 28, Chilo Daglari. 29, Sat Dagları S of Sat Gölü. 30, Varegös, SW of Yüksekova. 31, Locality S of
Yüksekova.

228

HURKMANs: Monograph of Merodon

Fig. 81-88. Distribution of Merodon. — 81, M. alexeji (MM), M. sophron (©) and M. tener (4); 82, M. hirsutus (©), M. hypo-
chrysos (A) and M. marginicornis (EB); 83, M. rufitarsis (8) and M. tarsatus (A); 84, M. ankylogaster (@), M. auronitens (W),
M. caudatus (©) and M. oidipous (à); 85, M. persicus (©), M. quadrinotatus (4) and M. xanthipous (li); 86, M. brevis (©)
and M. turkestanicus (MB); 87, M. kaloceros (WI) and M. tangerensis (A); 88, M. cupreus (©), M. hamifer (W), M. vandergooti
(W) and M. warnckei (A).

229

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 89-93. Distribution of Merodon. — 89, M. aberrans (A) and M. clavipes (B); 90, M. aberrans isperensis (®); 91. M. velox
(©) and M. splendens (A); 92, M. erivanicus (A); 93. M. longicornis (©) and M. lusitanicus (A).

230

HurkMANS: Monograph of Merodon

Fig. 94-98. Distribution of Merodon. — 94, M. pruni (M); 95. M. crassifemoris (A) and M. femoratoides (©); 96, M. avidus
(M); 97, M. nigritarsis (A); 98, M. bequaerti (©), M. manicatus (W) and M. testaceus (MM).

231

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 99-106. Distribution of Merodon. — 99, M. alagoezicus (©), M. dzhalitae (A), M. elegans (B) and M. toscanus (W); 100,
M. schachti (©) and M. taniniensis (4); 101, M. nitidifrons (©) and M. satdagensis (A); 102, M. altinosus (A) and M. lucasi
(©); 103. M. biarcuatus (A) and M. distinctus (©); 104, M. clunipes (MM) and M. femoratus (©); 105, M. mariae (@) and M.
testaceoides (À); 106, M. aureotibia (A) and M. ottomanus (B).

292

INDEX

References to the main treatment are in bold type.

aberrans 147, 155, 156, 176-178, 181-183, 203-205, 215,
221, 222, 230

aberrans aberrans 176, 204, 221

aberrans flavitibius 176

aberrans isperensis 147, 177, 230

affinis 157, 203, 205

alagoezicus 156, 157, 185, 191, 193, 198, 200, 205, 206,
225,232

alagoezicus group 147, 150, 153, 157, 159, 192, 194, 196,
197-203, 205, 212

alexeji 154, 157, 163-169, 182, 204, 206, 217, 229

alexeji group 147, 153, 162-165, 167-169, 204

altinosus 147, 154, 163, 164-165, 204, 217, 232

amaryllidis 152

Amphoterus 148

angustiventris 207

ankylogaster 147, 154, 156, 169, 204, 219, 229

apimimus 214

armipes 214, 215

aureotibia 147, 157, 189, 203-205, 227, 232

aurifer 191, 205

auronitens 147, 154, 169, 170, 204, 219, 229

avidus 152, 153, 157, 180, 182, 185, 190, 191-194, 196,
203-206, 214, 215, 224, 231

avidus group 147, 150, 153, 191, 203, 205

Azpeytia 147, 150, 207, 208, 214, 215

balanus 207

bardus 207

bautias 207

bequaerti 147, 155, 194-197, 205, 224, 231

bessarabicus 153

biarcuatus 156, 157, 159, 204, 216, 232

bicolor 207

bipartitus 207

bombiformis 152

brevis 155, 176, 177, 178, 183, 205, 206, 221, 229

canipilus 178, 179, 205 x

caudatus 151,154, 157, 169, 170, 171, 204, 205, 219, 229

chiragra 207

cimbiciformis 215

clauda 178, 179, 205, 206

clavipes 149, 155, 157, 178-186, 204-206, 222, 230

clavipes group 147, 153, 175, 181, 182, 204

clavipes var. albus 178, 180, 205

clavipes var. ater 178, 205

clavipes var. niger 178, 205

clunipes 154, 156, 158-160, 204, 216, 232

coerulea 207

contrarius 207

Copestylum 150

crassifemoris 153, 193, 194, 205, 206, 224, 231

crassifemoris group 147, 150, 153, 193, 205

cupreus 147, 155, 156, 179-181, 183, 205, 222, 229

curvipes 149, 178, 179, 205-207

dimorphus 159, 204

distinctus 150, 154, 156-159, 182, 204, 205, 214, 216,
232

distinctus assemblage 147, 153, 157, 204

dobrogensis 153

dzhalitae 155, 180, 181, 183, 205, 232

HURKMANS: Monograph of Merodon

edentulus 207

elegans 147, 155, 157, 195-197, 204-206, 225, 232

elegans group 147, 153, 194, 195, 202, 205

equestris 149, 150, 152, 166, 215

Eristalis 149

erivanicus 157, 186-188, 205, 206, 223, 230

Eumerus 148, 150, 207, 214, 215

Exmerodon 149, 152, 167

femoratoides 156, 157, 189-191, 194, 205, 206, 223, 231

femoratus 154, 156, 158, 160, 161, 180, 204, 205, 216,
232

flavitibius 155, 156, 176, 178, 205

fulcratus 149, 152, 153, 167, 168

fulvus 185, 186, 205, 206

furcata 207

fuscinervis 185, 186, 205, 206

graecus 191, 205

gravipes 178, 179, 205, 206

grisea 207

hamifer 155, 156, 176, 181, 205, 222, 229

Helophilus 207

hirsutus 154, 156, 164, 165, 169, 204, 217, 229

hypochrysos 147, 154, 162, 165, 204, 217, 229

interveniens 207

isperensis 147, 155, 156, 177, 205, 230

italicus 191, 192, 205

kaloceros 147, 155, 157, 187, 188, 205, 223, 229

karadaghensis 155, 156, 181, 205

kawamurae 154, 157, 165, 166, 186, 204, 206, 218, 228

kiritshenkoi 153

knerii 176, 205

loewi 152 ‘

longicornis 155, 157, 186-188, 205, 214, 223, 230

longicornis group 147, 153, 162, 186, 188, 197, 203-205

lucasi 147, 156, 157, 198, 199, 205, 225, 226, 232

lusitanicus 147, 157, 181, 182, 205, 222, 230

Mallota 150, 207, 214, 215

manicatus 155, 157, 194-196, 205, 225, 231

marginicornis 147, 154, 162, 166, 167, 169, 204, 218, 229

mariae 147, 154, 157, 160, 161, 204, 216, 232

Megatrigon 148

melota 207

micromegas 165, 166, 204

minutus 215

morosus 207

nigritarsis 156, 157, 182, 190, 193, 194, 205, 214, 224,
25)

nigritarsis group 147, 150, 153, 189, 203, 205

nitidifrons 147, 155, 157, 199, 200, 205, 226, 232

nudus 215

obscuripennis 176, 205

oidipous 147, 155, 171, 204, 219, 229

olivaceus 215

ornatus 207

Orthoprosopa 207

ottomanus 147, 154, 161, 162, 204, 217, 232

pallida 185, 205

Palpoda 207

persicus 147, 154, 171, 172, 204, 220, 229

Platynochaetus 150, 214, 215

Polydontomyia 207

pruni 153, 157, 183, 185, 186, 205, 206, 222, 231

pruni group 147, 153, 185, 205

pruni var. obscurus 185, 186, 205

quadrilineatus 191, 205

233

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

quadrinotatus 156, 182, 205, 222, 229
Quichuana 207

ruficornis 171, 195, 215

rufitarsis 154, 162, 167, 168, 204, 218, 229
rufitibius 191, 192, 205

rufus 152, 215

sacki 178, 179, 205

satdagensis 147, 156, 157, 198, 200, 201, 205, 225, 226,

232
schachti 147, 156, 157, 201, 205, 225, 227, 232
scutellaris 147, 207, 208, 214, 215
senilis 178, 179, 205
serrulatus 191, 205
shirakii 147, 208, 215
sicanus 185, 205
smirnovi 171-174, 204, 206
sophron 147, 154, 162-164, 168, 204, 218, 229
Sphegina 207
spinipes 189, 191, 193-195, 204-206
splendens 147, 155, 176, 182, 183, 205, 222, 230
strigatus 214, 215
sulcitibius 215
takasagoensis 214, 215

234

tangerensis 147, 154, 172, 173, 204, 220, 229
taniniensis 147, 155, 157, 201, 202, 205, 227, 232
tarsatus 154, 156, 167, 173, 174, 204, 206, 220, 229
tarsatus group 147, 153, 169, 171, 204

tenebricus 207

tener 153, 157, 168, 204, 229

testaceoides 147, 154, 162, 204, 217, 232

testaceus 155, 157, 162, 195-197, 205, 225, 231
torpidus 207

toscanus 147, 155, 157, 192, 202, 205, 227, 232
trizonus 169, 204

tuberculatus 207, 215

turkestanicus 155, 156, 169, 174, 204, 206, 221, 229
umbrifer 207

vandergooti 147, 153, 188, 189, 203, 205, 223, 229
vandergooti group 147, 150, 153, 188, 205
varicolor 207

velox 155, 157, 183-186, 205, 230

velox var. anathemus 183, 184, 205

velox var. armeniacus 183, 184, 205

viaticus 203, 204-206

warnckei 147, 155, 156, 176, 184, 185, 205, 222, 229
xanthipous 147, 154, 175, 204, 221, 229

HERMAN DE JONG

Institute for Taxonomic Zoology, Amsterdam

THE PHYLOGENY OF THE NEPHROTOMA
FLAVESCENS SPECIES GROUP (DIPTERA:

TIPULIDAE)

De Jong, H., 1993. The phylogeny of the Nephrotoma flavescens species group (Diptera:
Tipulidae). — Tijdschrift voor Entomologie 136: 235-256, figs. 1-114, tabs. 1-2, appendix.
[ISSN 0040-7496]. Published 10 December 1993.

The phylogeny of the currently recognized 22 species and subspecies of the west Palaearctic
Nephrotoma flavescens species group (Diptera, Tipulidae) is analysed on the basis of 50 morpho-
logical characters. The results of the present study are compared with the phylogeny of the fla-
vescens group published by Oosterbroek (1980). A synopsis of the distribution of the species
and subspecies is given.

H. de Jong, Department of Entomology, Institute for Taxonomic Zoology (Zoological

Museum), Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands.
Key words. — Diptera, Tipulidae, Nephrotoma, phylogeny, west Palaearctic.

In this paper I will discuss the phylogenetic rela-
tionships of the 22 species and subspecies of the
Nephrotoma flavescens species group sensu Ooster-

broek (1980).

The concept of the flavescens group adopted here is
in accordance with the views of Oosterbroek (1980,
1982). A list of the 22 species and subspecies of the
flavescens group as recognized in the present paper is
given in table 1. A systematic revision of the twenty
species and subspecies of the flavescens group known
at that time was incorporated in Oosterbroek (1978).
Subsequently, Oosterbroek (1982) introduced N.
cretensis and N. guestfalica hartigiana as new members
of the group. Oosterbroek (1985a) furthermore in-
cluded the Japanese species N. subpallida Alexander
in the flavescens group. For reasons explained below
(section “Discussion of adopted phylogeny’), I do not
accept this combination. An as yet unnamed species,
known of female specimens originating from north-
ern Algeria and northern Tunisia, awaits description
until the male sex becomes available. It is not includ-
ed in the phylogenetic analysis of the present paper.

The phylogeny of the flavescens group is analysed as
part of a research project on the historical biogeogra-
phy of the western Mediterranean (see De Jong in
press). Previously, Oosterbroek (1980) presented a
phylogenetic analysis of this species group, but addi-
tional information on the distribution of certain char-
acters within Nephrotoma prompted a reconsideration
of Oosterbroek’s conclusions. In the section
‘Discussion of adopted phylogeny’, I will compare the

results of the present study with Oosterbroek’s phylo-
genetic analysis of the flavescens group.

The flavescens group has an essentially west
Palaearctic distribution, with most of its species and
subspecies being confined to the Mediterranean sub-

Table 1. The species and subspecies of the Nephrotoma fla-
vescens species group as recognized in this paper. + = exam-
ined; - = not examined. Females of an as yet undescribed
species are known from north Algeria and north Tunisia.

3} 2
appendiculata appendiculata Pierre, 1919 + +
appendiculata pertenua Oosterbroek, 1978 + +
astigma Pierre, 1925 + +
beckeri Mannheims, 1951 + +
cretensis Oosterbroek, 1982 + +
exastigma Oosterbroek, 1978 + +
flavescens Linnaeus, 1758 + +
fontana Oosterbroek, 1978 + +
guestfalica guestfalica Westhoff, 1879 + +
guestfalica hartigiana Oosterbroek, 1982 + +
guestfalica surcoufi Pierre, 1925 + 4
lempkei Oosterbroek, 1978 + +
minuscula Mannheims, 1951 + +
nasuta Oosterbroek, 1975 + +
quadrifaria quadrifaria Meigen, 1804 + +
quadrifaria farsidica Savchenko, 1957 - +
saccai Mannheims, 1951 + +
schaeuffele Mannheims, 1964 + +
spatha Oosterbroek, 1975 + +
submaculosa Edwards, 1928 + +
sullingtonensis Edwards, 1938 + +
theowaldi Oosterbroek, 1978 + +

235

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 1. Distribution of the Nephrotoma flavescens species group.

region (fig. 1). The distribution range of six represen-
tatives of the flavescens group extends towards north-
west Europe. A single male specimen of the species
flavescens Linnaeus has been reported from western

Greenland (cf. Mannheims & Theowald 1971).

Detailed information on the distribution of the
species and subspecies of the flavescens group can be
found in Oosterbroek (1978: distribution maps of
the species and subspecies discussed therein) and
Oosterbroek (1982, 1985b: additional distributional
data). Oosterbroek & Theowald (1992) summarized
the ranges of the Palaearctic Tipulidae, including the
members of the flavescens group. Distribution maps
of each of the species and subspecies of the flavescens
group will be included in a forthcoming paper deal-
ing with the historical biogeography of the western
Mediterranean. A synopsis of the distribution of the
members of the flavescens group is given at the end of
the present paper.

A key to the identification of the species and sub-
species of the flavescens group (as well as all other
western Palaearctic members of Nephrotoma known
at the time) can be found in Oosterbroek (1979c).
The more recently described N. cretensis and N. guest-
falica hartigiana will key out under couplet 13 and

236

couplet 3 respectively. Oosterbroek (1982) listed dis-
tinguishing characters for both taxa.

MATERIAL, METHODS AND TERMINOLOGY

The majority of specimens used for the present stu-
dy are preserved in the collection of the Department
of Entomology of the Institute for Taxonomic
Zoology (Zoological Museum), Amsterdam. Addi-
tional material was kindly lent by Dr. Eulalia Eiroa
(Department of Animal Biology, University of
Santiago de Compostela, Spain). Specimens were
usually dry pinned, occasionally material was preser-
ved in alcohol.

As most characters used for the phylogenetic analy-
sis pertain to structures of the terminalia of both sex-
es, preparations of these parts were made. To this end
the specimen was relaxed for a few hours in a humifi-
er, after which the (end of the) abdomen was de-
tached. The removed parts were macerated for about
three to five minutes in a nearly boiling 10% KOH
solution in a bain-marie. The macerated structures
were rinsed with water (several times) and 70% alco-
hol, upon which they were transferred to a watch-
glass filled with glycerol. Examination of the termi-
nalia was carried out using a Wild stereomicroscope

with a magnification up to 100 X. The drawings were
made with the aid of a drawing tube on the micro-
scope. For lasting preservation the terminalia were
stored in glycerol in a microvial pinned under the
pertaining specimen.

The programs HENNIG86, version 1.5 (Farris 1988)
and PAUP, version 3.1.1 (Swofford 1993) were used to
analyse the phylogeny. For details on the computa-
tional methods employed, see the section ‘Discussion
of adopted phylogeny’.

The terminology used is largely in accordance with
McAlpine (1981). I employ some additional terms to
denote details of the male and female terminalia.
Tangelder (1985) published a review of the morphol-
ogy of the terminalia of the Tipulidae, and for further
details I refer the reader to her paper. The terms used
in the present text are explained in figures 3-10, 13,

14, 30 and 50.

BIOLOGY

The species of the flavescens group occur in a varie-
ty of habitats, ranging from grassland, gardens,
scrubs, deciduous as well as coniferous forests to dry
sandy places. Adults are usually on the wing in spring
and early summer, although minuscula Mannheims

DE Jone: Phylogeny Nephrotoma flavescens group

(Cyprus and the Levant) flies as early as the middle of
January and records of flavescens Linnaeus (central
and western Europe) are known from as late as
September. The species of the flavescens group can be
found from sea-level up to 2400 m, the highest alti-
tude recorded for flavescens in the Alps.

Several authors described immature stages of mem-
bers of the group, viz. of appendiculata appendiculata
Pierre, flavescens, guestfalica guestfalica Westhoff,
quadrifaria quadrifaria Meigen, and submaculosa
Edwards (cf. Brindle 1960, Chiswell 1956,
Hemmingsen & Jensen 1972, Theowald 1957,
1967). Oosterbroek (1978) presented details on the
biology of each of the then recognized members of
the flavescens group.

SYSTEMATIC POSITION

Mannheims (1951) separated the west Palaearctic
species of Nephrotoma with acutely pointed cerci in
the female from the remaining species of the genus.
Savchenko (1973), largely adopting this major divi-
sion, included most of the species involved in his cor-
nicina species group. The cornicina group sensu
Savchenko contains the species of Nephrotoma that in
the male carry an extension on the posterior margin

Table 2. Character state matrix of the species and subspecies of the Nephrotoma flavescens species group.

(Sub)species Character

5 10 15 20 25 30 35 40 45 50
appendiculata 00010 11000 11011 00100 01000 11011 10111 00000 00011 00000
a. pertenua 00010 11000 11011 00100 01000 11011 10111 00000 00011 00000
astigma 10010 00010 10111 10100 01110 01111 11000 10000 00011 01000
beckeri 00110 00000 10001 01110 01000 01010 00010 00000 00011 00010
cretensis 00110 00000 10001 01110 01000 01010 00000 00000 00011 00010
exastigma 10010 00000 10000 00100 01000 01111 00000 00000 00011 01000
flavescens 10010 00010 10111 10100 01110 01011 11000 10000 00011 01000
fontana 00010 00010 11011 00100 01100 01011 10000 : 00000 00011 00000
guestfalica 00011 00000 10001 10101 00001 01111 00000 01010 00011 10000
£ hartigiana 00011 00000 10001 10101 10001 01111 00000 01010 00011 10000
£ surcouf? 00011 00000 10001 10101 10001 01111 00000 01010 00011 10000
lempkei 10010 00011 10111 10100 01110 01111 11000 10000 00011 01100
minuscula 10010 10100 10011 00100 01000 01011 10111 00000 11011 00001
nasuta 00000 00000 10101 00100 01000 01010 00000 00000 00000 00000
quadrifaria 01000 00000 11011 01100 01000 01111 10010 00101 00101 00000
q. farsidica 01000 00000 11011 01100 01000 01111 10010 00101 00101 00000
SACCAL 00000 00000 1-001 00100 01000 01010 00010 00000 00010 00000
schaeuffelei 00000 00000 10000 00000 01000 00010 00000 00000 00100 00000
spatha -0010 00000 10001 10000 01000 01011 00000 00000 00100 00000
submaculosa 10010 00011 10111 10100 01110 01011 11000 10000 00011 01100
sullingtonensis 00010 00010 11011 00100 01000 01011 10000 00000 00011 00000
theowaldi 00010 10100 10011 10100 01000 01011 10111 00000 11011 00001

257

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Fig. 2. Consensus tree of two equally parsimonious trees with length 73, consistency index 68 and retention index 84 for the
species and subspecies of the Nephrotoma flavescens species group. — synapomorphy; LJ: homoplasy; X : reversal.

of sternite eight and that are devoid of bristles on the
alar squama. Thus Savchenko excluded the species
guestfalica Westhoff from his cornicina group, mean-
while adding several central and east Palaearctic as
well as Oriental species to the group.

Based on three presumed synapomorphies, the fla-
vescens species group was first recognized by Ooster-
broek (1980) as a subgroup of the cornicina group.
The monophyly of the flavescens group will be further
discussed below.

The exact position of the flavescens group within
the cornicina group is not yet established. Ooster-
broek (1980) presented a phylogeny of the cornicina
group, but restricted his analysis to the western
Palaearctic species. This approach, for obvious rea-
sons, led to the conclusion that the flavescens group is
closely related to western Palaearctic species of the
cornicina group. I have made a preliminary analysis of
the phylogeny of the cornicina group with the inclu-
sion of some of its east Palaearctic and Oriental mem-
bers. Results of this investigation show that the flaves-
cens group is probably more closely related with east
Palaearctic species of the cornicina group. An exhaus-
tive analysis of the cornicina group, however, was be-

238

yond the scope of the present paper.

In the phylogenetic analysis of the flavescens group
I employed the non-flavescens group members of the
cornicina group as the first level outgroup, and the re-
maining species of Nephrotoma as the second level
outgroup.

PHYLOGENETIC ANALYSIS

In this section I will discuss the characters used in
the phylogenetic analysis of the flavescens group. Each
character dealt with contains a brief description of its
presumed plesiomorphous and apomorphous states,
accompanied by their respective codings (0: plesio-
morphous, 1: apomorphous). The character state ma-
trix is given in table 2. The resulting cladogram for
the flavescens group is shown in figure 2 and is discus-
sed in the next section. Autapomorphies of the species
and subspecies of the flavescens group are given in the
appendix to this paper.

Legs

1. — Tarsal claws of male: (0) medial tooth present (fig.

11, arrowed); (1) medial tooth absent (fig. 12) (cf.
Oosterbroek 1980: 337, character 24).

The male claws lack the medial tooth in the species
exastigma, minuscula and the members of the clade
astigma to submaculosa. In the species spatha the male
claws usually bear this tooth, but there are specimens
with some or all of the claws toothless. Absence of the
medial tooth on the male claw is due to reduction.
Females of Nephrotoma always have the tarsal claws
devoid of teeth.

Species with toothless claws in the male occur fre-
quently in other groups of Nephrotoma. According to
Oosterbroek (1980: 337) and Tangelder (1985: 160),
this character state is usually restricted to smaller
monophyletic groups. Referring, among other things,
to this character state, Oosterbroek (1980) united the
species exastigma and spatha with the members of the
clade astigma to submaculosa, an interpretation not
followed in the present paper.

Wing

2. — Wing-membrane: (0) unmarked; (1) dark crossband
running from pterostigma over base of discal cell, crossvein
m-cu and apical portion of vein Cu; wing-tip darkened (fig.
13).

The subspecies quadrifaria quadrifaria and quadri-
faria farsidica are distinguished among the members
of the flavescens group by dark shades below the pte-
rostigma and on the wing-tip. Other species of
Nephrotoma with similarly marked wings, e.g. atrosty-
la Alexander, bifusca Alexander, gaganboi Tangelder,
and neopratensis Alexander, are not closely related
with quadrifaria.

Male terminalia

3. — Aedeagal guide, apex: (0) gradually tapering towards
tip (figs. 5, 14-17, 19-26, 28); (1) with knob-like enlarge-
ment at tip (fig. 18).

The species beckeri and cretensis are sister species
that, among other things, are characterized by the
shape of the aedeagal guide that terminates in an en-
larged tip. Oosterbroek (1982) distinguished this
shared derived character state in his discussion on cre-
tensis (as ‘shape of central part of adminiculum’).

4. — Aedeagal guide, gonapophysis: (0) dorsal extension
elongate (figs. 14, 16, 17); (1) reduced in length (figs. 5, 15,
18-20, 22-26, 28).

The species spatha and the majority of species and
subspecies of the clade beckeri to submaculosa either
have a relatively short or completely reduced dorsal
extension of the gonapophysis. Contrary to this, both
subspecies of quadrifaria possess a relatively long dor-
sal extension (fig. 21). I interpret the character state
in quadrifaria as a reversal of character state 4(1).

The majority of species of the cornicina group have

DE Jone: Phylogeny Nephrotoma flavescens group

relatively long dorsal extensions of the gonapophyses.
Outside the cornicina group long dorsal extensions
are known in species of the analis, brevipennis, dorsal-
is, and pedunculata species groups of Nephrotoma (cf.
Oosterbroek 1979b, 1984, 1985a; Tangelder 1983,
1984).

5. — Aedeagal guide, gonapophysis: (0) posterior exten-
sion apically rounded (figs. 14-19, 21, 23-26, 28); (1) dor-
soventrally flattened and with upcurved acute tip (fig. 20).

The three subspecies of guestfalica show the apo-
morphous state of this character. Within the flaves-
cens group, the only other taxa with an acute dorsal
tip of the posterior extension of the gonapophysis are
both subspecies of appendiculata (fig. 22). In the lat-
ter two subspecies the posterior extension is mediolat-
erally flattened (see character 7).

6. — Aedeagal guide, gonapophysis: (0) posterior exten-
sion parallel sided or tapering towards tip (figs. 5, 14-17, 19-
21, 24-26, 28); (1) spatulate at tip (figs. 22, 23).

The members of the clade appendiculata to theo-
waldi have the tip of the posterior extension of the
gonapophysis dilated and laterally flattened, the pos-
terior extension thus appearing as a spatulate struc-
ture. In both subspecies of appendiculata the spatulate
part has an acute dorsal tip (fig. 22; see next charac-
ter), in minuscula and theowaldi it is rounded off and
somewhat downcurved (fig. 23; see character 8).

7. — Aedeagal guide, gonapophysis: (0) tip of posterior ex-
tension rounded off when seen in profile (figs. 14-17, 19,
21, 24-26, 28); (1) dorsally with acute tip (fig. 22).

The two subspecies of appendiculata show the apo-
morphous state of this character. The only other
members of the flavescens group with a dorsally acute
posterior extension of the gonapophysis are the three
subspecies of guestfalica (fig. 20). Here the extension
is dorsoventrally flattened and thus of an overall dif-
ferent structure (see character 5).

8. — Aedeagal guide, gonapophysis: (0) ventral margin of
posterior extension almost straight when seen in profile (figs.
5, 14-22, 24-26, 28); (1) downcurved at tip (fig. 23) (cf.
Oosterbroek 1980: 337, character 25).

The tip of the posterior extension of the gonapo-
physis is somewhat downcurved in the species minus-

culaand theowaldi.

9. — Aedeagal guide, gonapophysis: (0) variously shaped
(figs. 14-23); (1) consisting of a mediolaterally flattened and
relatively high anterodorsal part and an elongate posterior
projection which gradually tapers towards its tip (figs. 5, 24-
26, 28).

The species of the clade sullingtonensis to submacu-
losa all have a similarly built aedeagal guide plus ap-

239

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

tg7 tg8 tg9 tg10 ogonst igonst lat shell centr rdg

N
NIT

ANDER

gon

ventr app

comp apod ant pt

_ post apod ant lat pt =

È

aed ant apod 6 7

tg8 tg9 tg10 cerc

A

spmth genfk st9

240

pendages, which I interpret as a synapomorphy.

10. — Aedeagal guide, gonapophysis: (0) anterodorsal
margin rounded (figs. 26, 27); (1) anterodorsal margin in-
curved and acutely tipped (fig. 28, 29, arrowed) (cf.
Oosterbroek 1980: 337, character 28).

The apomorphous character state separates the spe-
cies lempkei and submaculosa from the remainder of

the flavescens group.

11. — Aedeagal guide, rod: (0) rod present ventral of
sperm pump (fig. 30); (1) absent (fig. 31) (cf. Oosterbroek
1980: 332, character 12, as ‘adminicular rods’).

The species of the flavescens group are devoid of
rods of the aedeagal guide, although slight vestiges of
these structures can be found in individual specimens
of the species beckeri and saccai. Rods of the aedeagal
guide are usually present in other species of
Nephrotoma, including the species of the cornicina
group outside the flavescens group. Reduction of the
rods occurs throughout Nephrotoma (cf. Oosterbroek
1980: 349, character 5).

12. — Aedeagus: (0) basal, tubular section about 0.50 of
total length or longer (figs. 32-35, 37, 39, junction of tubu-
lar and branched sections of aedeagus arrowed); (1) basal,
tubular section about 0.40 to 0.20 of total length (figs. 36,
38, junction of tubular and branched sections of aedeagus
arrowed).

Half of the species and subspecies of the clade
quadrifaria to submaculosa have the branching point
of the aedeagus situated before its midlength.
Reversal of this state seems to have occurred twice
within this clade, once in the ancestor of both minus-
cula and theowaldi (cf. fig. 37) and once in the ances-
tor of the clade astigma to submaculosa (cf. fig. 39). In
the latter clade reversal of character 12(1) coincided
with shortening of the aedeagus (see next character).

The presumed plesiomorphous situation of charac-
ter 12 can be found in non-flavescens group members
of Nephrotoma with a partly tripartite aedeagus, e.g.
alticrista Alexander, integra Alexander, medioproducta
Alexander, nigrohalterata Edwards, and subpallida

Alexander.

DE Jone: Phylogeny Nephrotoma flavescens group

13. — Aedeagus: (0) relatively long and slender (figs. 32,
33, 35-38); (1) short and thick (figs. 34, 39).

The species nasuta (fig. 34) and the members of the
clade astigma to submaculosa (cf. fig. 39) have a rela-
tively short aedeagus. Oosterbroek (1980: 336, char-
acter 23) also mentioned the species exastigma and
spatha as having a short aedeagus (as ‘intromittent
organ’). Oosterbroek (1980: 350-351, diagram 1;
391-393, appendix 2B) related the length of the ae-
deagus with the length of the second abdominal ter-
gite. Contrary to this, the relative proportions of the
aedeagus are perhaps better expressed by comparing
the length of the aedeagus with its diameter. When
the latter procedure is followed and the - arbitrary -
limit between relatively long and short is appointed to
a quotient of 100 for the total length of the aedeagus
and its diameter just after the sperm pump, the aedea-
gus of both exastigma (fig. 35) and spatha (fig. 33)
turns out to be relatively long (cf. measurements in
Oosterbroek 1980, appendix 2A).

In the species of the cornicina group outside the fla-
vescens group, the aedeagus is a relatively long structure.

14. — Gonocoxite: (0) dorsomedially fused with base of
aedeagal guide (fig. 40); (1) separate (fig. 41, gap arrowed).

The species of the clade quadrifaria to submaculosa
show a membranous gap between the base of the ae-
deagal guide and the adjacent dorsomedial corner of
the gonocoxite. In the remaining species of the flaves-
cens group both structures are fused, as they are in the
other members of the cornicina group. Oosterbroek
(1980: 337, character 26, as separation of ventral ap-
pendages of adminiculum and sternite nine) distin-
guished the same character state but recognized it in
the species of the clade astigma to submaculosa only.
Oosterbroek noted that the gap between both struc-
tures can be narrow in other species of Nephrotoma.
Several species of the flavescens group show this latter
state, e.g. fontana.

15. — Gonocoxite, medisternal sclerotization: (0) fused
with ventral appendage of aedeagal guide (fig. 42, connexion
arrowed); (1) structures separate from each other (fig. 43,
gap arrowed; figs. 8, 47-49).

Figs. 3-10. Nephrotoma flavescens, 3-8, male; 9, 10, female. — 3, terminalia, lateral view; 4, tergite 10, ventral view; 5, aedea-
gal guide, lateral view; 6, sperm pump, lateral view; 7, inner gonostylus, lateral view; 8, sternite 9 and gonocoxites, ventral
view; 9, terminalia, lateral view; 10, hypogynial valve, medial view.

Abbreviations: aed: aedeagus; aed gd: aedeagal guide; ant apod: anterior apodeme of sperm pump; ant lat pt: anterolateral part
of inner gonostylus; ant pt: anterior part of inner gonostylus; centr rdg: central ridge of tergite 10; cerc: cercus; comp apod:
compressor apodeme of sperm pump; crst: crest of inner gonostylus; ej duct: ejaculatory duct; gen fk: genital fork; gon: gon-
apophysis of aedeagal guide; goncx: gonocoxite; hyp vlv: hypogynial valve; i gonst: inner gonostylus; lat shell: lateral shell of
tergite 10; ma rdg: major ridge of hypogynial valve; med scl: medisternal sclerotization; mi rdg: minor ridge of hypogynial
valve; o gonst: outer gonostylus; post apod: posterior apodeme of sperm pump; rdg: ridge of inner gonostylus; sp2: lateral
remnant of genital bridge; spmth: spermatheca; st7 etc.: sternite 7 etc.; st8 ext: extension of sternite 8; tg7 etc.: tergite 7 etc.;

ventr app: ventral appendage of aedeagal guide.

241

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Within the flavescens group, schaeuffelei and exas-
tigma are the only species which have the ventral ap-
pendages of the aedeagal guide fused with the medis-
ternal sclerotizations. In all other members of the
group these structures are separated by a membranous
gap. When the medisternal sclerotizations are well de-
veloped, they are united with the appendages of the
aedeagal guide in the non-flavescens group members
of the cornicina group.

16. — Gonocoxite, medisternal sclerotization: (0) well de-
veloped, large (fig. 42, 44, 46-48); (1) reduced in size, small
or absent (figs. 8, 43, 45, 49).

It is assumed that the presence of medisternal scle-
rotizations is a plesiomorphy within the genus
Nephrotoma (Oosterbroek 1980: 321; Tangelder
1985: 144). Reduced medisternal sclerotizations oc-
cur within the flavescens group in the species spatha
(fig. 43), the three subspecies of guestfalica (cf. fig.
45), the species theowaldi, and the members of the
clade astigma to submaculosa (figs. 8, 49). The shape
of the reduced medisternal sclerotizations varies
among these taxa.

17. — Gonocoxite, medisternal sclerotization: (0) not in
contact with ventromedial margin of gonocoxite or only so
along a relatively short zone (figs. 8, 42, 43, 45, 47-49); (1) fu-
sed for more than half its length with gonocoxite (figs. 44, 46).

When in contact with the gonocoxites, the medi-
sternal sclerotizations are usually only fused with
these structures near their anterior margins (cf. fig.
47). The sister species beckeri (fig. 44) and cretensis,
and both subspecies of quadrifaria (cf. fig. 46) are
characterized by a more complete fusion of both
structures. I assume that fusion evolved independent-
ly in both pairs of taxa.

18. — Inner gonostylus: (0) a projecting posterolateral
part present (figs. 50-52, structure arrowed in figs. 51 and
52); (1) posterolateral part reduced (figs. 7, 53-66).

Within the flavescens group, the species schaeuffelei
and spatha have a projecting structure placed postero-
laterally on the inner gonostylus. I interpret this
structure as the homologue of the posterolateral part
of the inner gonostylus of other genera and subgene-
ra of Tipulidae. As such it is a plesiomorphy in
Nephrotoma. In several species of the flavescens group
its remnant is readily identifiable in the low protuber-
ance covered with long hairs on the posterolateral
margin of the inner gonostylus (e.g. nasuta, fig. 53;
exastigma, fig. 57; appendiculata, fig. 62; minuscula,
fig. 63; sullingtonensis, fig. 64).

Several members of the cornicina group outside the
flavescens group have a distinct projecting posterolat-
eral extension on the inner gonostylus (fig. 50; cf.
Oosterbroek 1980: 330, character 5). Oosterbroek
interpreted this character state as an apomorphy of

242

his cornicina subgroup, but considering its occurrence
outside and within Nephrotoma, it is probably better
regarded as a plesiomorphy on this level of analysis.

19. — Inner gonostylus: (0) general shape various, seen in
lateral view with relatively short and high anterior part (figs.
7, 50-53, 57-66); (1) general shape similar, in lateral view
anterior part and crest appearing in line, elongate and slen-

der (fig. 54).

The inner gonostyli of the species beckeri and cre-
tensis show a number of features that can be interpret-
ed as apomorphies: the anterior part is acutely bent
over laterally (fig. 55; less so in the other members of
the flavescens group), the anterolateral part is smooth-
ly curved when seen in dorsal aspect (fig. 56; acute in
the other species of the flavescens group) and the crest
is posteriorly produced into an acute tip (fig. 54; the
only other representative of the flavescens group with
an acute crest is the subspecies guestfalica guestfalica,
fig. 58). Oosterbroek (1982) mentioned the synapo-
morphous resemblance of the shape of the inner go-
nostyli of beckeri and cretensis in his discussion on the
latter species.

20. — Inner gonostylus: (0) general shape various, lateral
edge toothless (figs. 7, 50-57, 61-66); (1) general shape si-
milar, laterally with edge toothed (figs. 58-60).

The three subspecies of guestfalica share some apo-
morphous peculiarities in the structure of the inner
gonostylus: the anterior part is large and produced,
while the lateral margin is present as a rim carrying
blackish sclerotized points. The inner gonostyli of the
subspecies differ in degree of reduction of the dorsal
crest (see next character).

21. — Inner gonostylus: (0) crest on anterior part well de-
veloped, posteriorly extending (fig. 58); (1) crest reduced in
length (figs. 59, 60).

Oosterbroek (1980: 326, character 12) considered
a posteriorly produced crest a synapomorphy of the
species of the cornicina group. A posteriorly produced
crest can therefore be interpreted as a plesiomorphy
within the flavescens group. Reduction of the posteri-
or extension of the crest is found in the subspecies
guestfalica hartigiana and guestfalica surcoufi. In guest-
falica hartigiana the crest is shortened (fig. 59), in
guestfalica surcouf? it is completely absent (fig. 60).

The members of the clade fontana to submaculosa
also show reduction of the posterior extension of the
crest. Reduction in this clade resulted in a distinct sit-
uation and is treated under character 23.

22. — Inner gonostylus: (0) without ridge (fig. 50); (1)
with ridge on its lateral side (figs. 7, 51-57, 61-66) (cf.
Oosterbroek 1980: 332, character 11).

All members of the flavescens group, with the ex-

DE JONG: Phylogeny Nephrotoma flavescens group

Figs. 11-29.—11, 12, Nephrotoma spatha, male last tarsal segment; 13, N. quadrifaria, wing; 14, N. schaeuffelei, aedeagal guide
and gonapophysis, lateral view; 15, N. spatha, idem; 16, N. nasuta, idem; 17, N. saccai, idem; 18, N. beckeri, idem; 19, N. ex-
astigma, idem; 20, N. guestfalica guestfalica, idem; 21, N. quadrifaria quadrifaria, idem; 22, N. appendiculata appendiculata,
idem; 23, N. minuscula, idem; 24, N. sullingtonensis, idem; 25, N. fontana, idem; 26, 27, N. astigma, 26, idem; 27, aedeagal
guide and gonapophysis, dorsal view; 28, 29, N. lempkei, 28, aedeagal guide and gonapophysis, lateral view; 29, aedeagal
guide and gonapophysis, dorsal view.

Abbreviations: aed gd: aedeagal guide; Cu: cubital vein; d: discal cell; dors ext: dorsal extension of gonapophysis; m-cu: me-
dio-cubital crossvein; post ext: posterior extension of gonapophysis; sq: squama.

243

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

ception of the three subspecies of guestfalica, are char-
acterized by the presence of a ridge on the posterior
part of the inner gonostylus. (Contrary to
Oosterbroek’s assertion, nasuta does have a ridge
underneath the crest; cf. fig. 53, arrowed).

Most other species of the cornicina group, and all
other Nephrotoma examined, lack this feature. The
only representatives of the cornicina group with a
ridge-like structure are cornicina cornicina Linnaeus,
cornicina sardiniensis Oosterbroek, and moravica

Figs. 30-39. — 30, Nephrotoma ligulata, sperm pump and part of aedeagus, lateral view; 31, 32, N. schaeuffelei, 31, idem; 32,
sperm pump and aedeagus, lateral view; 33, N. spatha, idem; 34, N. nasuta, idem; 35, N. exastigma, idem; 36, N. quadrifar-
ia quadrifaria, idem; 37, N. minuscula, idem; 38, N. fontana, idem; 39, N. astigma, idem.

Abbreviation: rod: rod of aedeagal guide.

244

Martinovsky. These three taxa together form a
monophyletic group within the cornicina group (cf.
Oosterbroek 1980). Here the ridge-like structure
probably evolved independently of that of the flaves-

cens group.

23. — Inner gonostylus: (0) crest on anterior part well de-
veloped, posteriorly extending (figs. 52-58, 61-64); (1) crest
reduced in length, posteriorly laterally downcurved (figs. 7,

65, 66).

In the clade fontana to submaculosa, the apomor-
phous state of this character can be found.
Oosterbroek (1980: 337, character 27) recognized
this apomorphy for the species of the clade astigma to
submaculosa. I claim that it also applies to the species

fontana (fig. 65).

24. — Inner gonostylus: (0) ridge without pubescence
(figs. 51-57, 61-65); (1) ridge covered with dense pubescen-
ce (fig. 7, 66) (cf. Oosterbroek 1980: 332, character 11X).

The species of the clade astigma to submaculosa are
distinguished from the remainder of species and sub-
species of the flavescens group, by the presence of pu-
bescence on the ridge of the inner gonostylus.

25. — Outer gonostylus: (0) at base with anteromedially
directed extension (figs. 67, 69); (1) with elongate ventrola-
teral extension (fig. 68).

The outer gonostylus of the majority of species of
Nephrotoma is provided with a more or less well de-
veloped anteromedially directed extension at its base.
In the three subspecies of guestfalica the outer gono-
stylus basally bears a long ventrolaterally directed ex-
tension.

26. — Outer gonostylus: (0) elongate, without strikingly
long hairs (figs. 67, 68); (1) broad, long haired along poste-
rior margin, with longest hairs about as long as width of ou-
ter gonostylus (fig. 69).

The outer gonostylus of the majority of species of
Nephrotoma is a rather slender structure when seen in
lateral view, covered with moderately long hairs. In
the two subspecies of appendiculata, the outer gono-
stylus is relatively broad and bears long hairs along its
posterior margin.

27. — Sperm pump: (0) sclerotized part of ejaculatory
duct attached to sperm pump (figs. 30, 31, 70, ejaculatory
duct arrowed); (1) membranous gap between sperm pump
and sclerotized part of ejaculatory duct (figs. 6, 71-73, 75)
(cf. Oosterbroek 1980: 332, character 14).

In the flavescens group the sclerotized part of the
ejaculatory duct is attached to the sperm pump in the
species schaeuffelei only (figs. 31, 70). In the other
species and subspecies of the group there either is a
membranous gap between the body of the sperm

DE Jone: Phylogeny Nephrotoma flavescens group

pump and the sclerotized part of the ejaculatory duct
or the ejaculatory duct is not sclerotized at all (see
next character). The gap between the sperm pump
and the sclerotized part of the duct is narrow in the
species beckeri (fig. 73), in the other relevant species
and subspecies of the flavescens group it is wider (figs.
1972245)

Oosterbroek (1980: 326, character 11) recognized
the presence of a midposterior appendage to the
sperm pump (as ‘aedeagus’) as a synapomorphy for
the cornicina group. I interpret the midposterior ap-
pendage as a sclerotization of the wall of the ejacula-
tory duct (seminal duct of Byers 1961). The species of
the cornicina group outside the flavescens group have
this sclerotized structure well developed, being at-

tached to the sperm pump (fig. 30).

28. — Sperm pump: (0) vestige of sclerotized portion of
ejaculatory duct present (figs. 6, 71-73, 75); (1) absent (fig.
74).

As mentioned under the preceding character, some
members of the flavescens group do not show a trace
of sclerotization of the ejaculatory duct. The species
and subspecies involved are exastigma (fig. 74), the
subspecies of guestfalica and quadrifaria, and astigma
and lempkei. It seems that reduction of the sclerotiza-
tion of the ejaculatory duct evolved along several lin-
eages and to various degrees within the flavescens

group.

29. — Sperm pump, anterior apodemes: (0) without long
membranous extensions at tip (fig. 30); (1) with long mem-
branous extensions at tip (fig. 6, 31, 71).

All members of the flavescens group are character-
ized by the possession of long translucent extensions
at the tips of the anterior apodemes of the sperm
pump. I did not find such extensions in species of
Nephrotoma outside the flavescens group.

30. — Sperm pump, anterior apodeme: (0) broad, bulging
in dorsal view (figs. 70, 73); (1) reduced to narrow strip
(figs. 72, 74, 75).

The anterior apodemes of the sperm pump are usu-
ally well developed in Nephrotoma. Within the flaves-
cens group, reduced anterior apodemes are present in
the species spatha (fig. 72) and the members of the
clade exastigma to submaculosa (figs. 74, 75).

Oosterbroek (1980: 333, character 15) also recog-
nized a reduction of the anterior apodemes (as ‘later-
al appendages of the aedeagus’) as an apomorphy in
the flavescens group. The largest anterior apodemes in
the flavescens group are found in the species nasuta
and saccai. Oosterbroek distinguished a monophylet-
ic group composed of the remainder of species and
subspecies of the flavescens group on account of this
feature. As relatively large anterior apodemes are also

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

7 post lat pt

Figs. 40-56. — 40, Nephrotoma guestfalica guestfalica, base of aedeagal guide in part plus right gonocoxite, posterolateral view;
41, N. quadrifaria quadrifaria, idem; 42, N. schaeuffelei, male sternite 9, gonocoxites and medisternal sclerotizations, ventral
view; 43, N. spatha, idem; 44, N. beckeri, idem; 45, N. guestfalica guestfalica, idem; 46, N. quadrifaria quadrifaria, idem; 47,
N. appendiculata appendiculata, idem; 48, N. minuscula, idem; 49, N. astigma, idem; 50, N. ligulata, inner gonostylus, later-
al view; 51, N. schaeuffelei, idem; 52, N. spatha, idem; 53, N. nasuta, idem; 54-56, N. beckeri, inner gonostylus, 54, lateral
view; 55, anterior view; 56, dorsal view.

Abbreviation: post lat pt: posterolateral part of inner gonostylus.

246

present in schaeuffelei, beckeri, and cretensis, I do not
adopt this interpretation.

31. — Sperm pump, compressor apodeme: (0) deeply in-
cised (fig. 76); (1) almost entire, dorsal margin either slight-
ly concave or entirely convex (fig. 77).

Within the flavescens group, the species and sub-
species of the clade quadrifaria to submaculosa have a
compressor apodeme which consists of an undivided
structure.

32. — Sperm pump, compressor apodeme: (0) entire, with
midposterior keel; (1) without midposterior keel.

The more basal species and subspecies of the clade
quadrifaria to submaculosa have the compressor apo-
deme provided with a well developed midposterior
keel. The keel is completely reduced in the species of
the clade astigma to submaculosa.

33. — Sperm pump, posterior apodeme: (0) yellowish co-
loured, elongate (figs. 6, 30, 31, 70-72, 74); (1) dark colou-
red and short (fig. 75).

Dark sclerotized, short posterior apodemes occur
in the species of the clade appendiculata to theowaldi
of the flavescens group. The remainder of species of
the group have the posterior apodemes less heavily
sclerotized.

In the clade appendiculata to theowaldi sclero-
tization of the posterior apodemes coincides with re-
duction of these structures. Oosterbroek (1980: 336,
character 22) recognized short posterior apodemes (as
‘posterior appendages of the aedeagus’) as an apo-
morphy of the clade appendiculata to theowaldi.
However, reduced posterior apodemes are also
known in the species nasuta and beckeri (fig. 73). The
apodomes in the two latter species are not blackish
sclerotized.

34. — Sp2: (0) haired (figs. 78, 83, arrowed); (1) bare
(figs. 79-82).

As in the majority of other species of the cornicina
group, most members of the flavescens group have the
posteromedial margin of sp2 near midlength provid-
ed with a concentration of hairs. Within the flavescens
group, these hairs are absent in the species saccai (fig.
79), beckeri (fig. 80) and in the clade quadrifaria to
theowaldi (cf. figs. 81, 82).

35. — Sp2: (0) variously shaped (figs. 78-81, 83); (1) si-
milar (fig. 82).

The species of the clade appendiculata to theowaldi
possess a similarly built sp2. The bare structure has
the dorsal part well developed and anteriorly protrud-
ing, while the ventral part is present as a straight ven-
tromedially directed appendix.

DE Jone: Phylogeny Nephrotoma flavescens group

36. — Sp2: (0) variously shaped (figs. 78-82); (1) similar
(fig. 83).

In the clade astigma to submaculosa the sp2 struc-
ture has a small and protruding dorsal part, while it is
slightly produced posteriorly near midlength at the
point of insertion of the hairs. The ventral part is
straight and gradually tapers towards its tip.

37. — Sternite 8: (0) with posterior extension (figs. 3, 84,
86); (1) without (fig. 85).

The only members of the flavescens group without
an extension on the posterior margin of sternite 8 are
the three subspecies of guestfalica. All other species of
the cornicina group have a well-developed posterior
extension of sternite 8.

38. — Sternite 8: (0) posterior extension dorsoventrally
compressed (fig. 84); (1) laterally compressed (fig. 86).

The extension of sternite 8 is short and laterally
compressed in both subspecies of quadrifaria. The
majority of species of the cornicina group have the ex-
tension either dorsoventrally flattened or rounded in
cross-section. Only cornicina cornicina, cornicina sar-
diniensis and moravica have the extreme apex of the
extension laterally compressed. However, in these
taxa the posterior extension of sternite 8 is dorsoven-
trally flattened at its base and elongate.

39. — Tergite 10: (0) variously shaped (figs. 3, 4, 89-94);
(1) consisting of two cup-shaped structures provided with
black spines along their anteromedial and posterior margins,
each carrying an acute lateral extension (figs. 87, 88).

The three subspecies of guestfalica are characterized
by their uniquely shaped tergite 10. Oosterbroek
(1980: 336, character 17P) considered guestfalica and
quadrifaria sister taxa, a relationship presumed,
among other things, to be supported by similarities in
the structure of tergite 10 (as ‘extension of tergite 9’).
In my opinion the shape of tergite 10 in quadrifaria
(figs. 89, 90) has no apomorphy in common with
that of guestfalica (figs. 87, 88).

40. — Tergite 10: (0) variously shaped, with relatively nar-
row medial incision, caudally directed (figs. 3, 4, 87, 88, 91-
94); (1) posteriorly widely emarginate, caudodorsally direc-
ted (figs. 89, 90).

The two subspecies of quadrifaria are characterized
by their peculiarly shaped tergite 10.

41. — Tergite 10: (0) central ridges posteriorly not or only
slightly produced (fig. 91); (1) central ridges posteriorly pro-
duced (fig. 93) (cf. Oosterbroek 1980: 336, character 17S;

tergite 10 as ‘extension of tergite 9’).

The central part of tergite 10 protrudes only mod-
erately in most species of the flavescens group. Only

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TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

minuscula and theowaldi show a posterior elonga-
tion of this part of tergite 10.

42. — Tergite 10: (0) lateral shells rounded along ventral
margin when seen in lateral view (fig. 92); (1) ventral mar-
gin truncate (fig. 94).

The species minuscula and theowaldi are the only
members of the flavescens group which have the ven-
tral margin of the lateral shells of tergite 10 straight.
In the other members of the group, as well as in most
other species of Nephrotoma, this margin is rounded.

Female terminalia

43. — Cercus: (0) long, well-developed, gradually tapering
towards tip (fig. 99); (1) reduced in length, tip rounded off
(figs. 95, 97, 106).

Within the flavescens group a shortened, blunt-
tipped cercus is known in the species schaeuffelei (fig.
95), spatha (fig. 97), and the two subspecies of quad-
rifaria (fig. 106). Besides being reduced in length, the
cercus of both subspecies of quadrifaria is also re-
duced in height. In the remainder of species and sub-
species of the flavescens group the cercus is long and
either gradually tapers towards the tip (fig. 99), or is
produced into a nipple-like extension (see next char-
acter).

Oosterbroek (1980: 326, character 14 and page
328, character 1) listed the presence of pointed cerci
in the female as one of the synapomorphies of the cor-
nicina group. The cerci in this group are also more ro-
bust than is usual in Nephrotoma (cf. Savchenko
1973: 35, 119). The plesiomorphous form of the cer-
cus in the cornicina group can be found in supposed-
ly basal species like forcipata Pierre and ligulata
Alexander. Here the well-developed cercus gradually
tapers towards its tip. In the flavescens group a similar-
ly shaped cercus is found in the species nasuta (fig.
99) and in the as yet undescribed species from Algeria
and Tunisia. The cercus of nasuta and the north-west
African species probably represents the plesiomor-
phous form in the flavescens group.

44, — Cercus: (0) gradually tapering towards tip (fig. 99);
(1) tip produced into a nipple-like extension (figs. 9, 101).

With the exception of both subspecies of quadri-
faria, the species and subspecies of the clade saccai to
submaculosa are characterized by the possession of a
nipple-like projection of the tip of the cerci. (The
nipple-like projection is frequently worn off in older
specimens).

As stated under the preceding character, the plesio-
morphous condition of the female cercus in the fla-
vescens group is probably represented by a moderately
pointed structure as found in nasuta (fig. 99).

45. — Hypogynial valve: (0) tip entire, with ventral mar-

248

gin evenly convex (figs. 96, 98, 100, 102); (1) tip irregular,
appearing to be broken off (figs. 9, 10, 103-105, 108-110)
(cf. Oosterbroek 1980: 333, character 16).

The species of the clade beckeri to submaculosa
show the apomorphous condition of this character.
Within the cornicina group a similar situation is
known in the non-flavescens group species saghaliensis
Alexander, where it apparently evolved independent-
ly (cf. fig. 6 of Oosterbroek 1985a. Fig. 13 of
Oosterbroek 1985a, depicts a damaged specimen of
esakii Alexander in which the tip of the hypogynial
valve is actually broken off; in intact specimens of this
species the hypogynial valve terminates in an acute
tip).

The basal species of the flavescens group, as well as
the other members of the cornicina group, have slen-
der, reduced hypogynial valves of which the ventral
margin of the tip is entire (cf. figs. 96, 98, 100, 102).

46. — Hypogynial valve: (0) dorsal margin continuous
(figs. 9, 10, 96, 98, 100, 102-104, 107-110); (1) dorsal mar-
gin with posteriorly directed point at about two-thirds of its
length (fig. 105).

The three subspecies of guestfalica are, among oth-
er things, characterized by their apomorphously
shaped hypogynial valves.

47. — Hypogynial valve: (0) major and minor ridges ba-
sally in contact with each other (figs. 102, 103, 108); (1) se-
parate (figs. 10, 109, 110).

In the Tipulidae there are usually two ridges on the
medial side of the hypogynial valve, a major (dorsal)
and a minor (ventral) one (cf. fig. 10). As this situa-
tion is found also in the majority of species of
Nephrotoma, I agree with Oosterbroek (1980: 326),
who interpreted the presence of two ridges as a plesio-
morphy for the genus. Tangelder (1985: 151), how-
ever, considered the same character state an apomor-
phy within Nephrotoma.

When present, both ridges are usually separate, like
they are in the species schaeuffelei of the flavescens
group (fig. 96). A partly or entirely reduced minor
ridge is frequently found in the flavescens group, e.g.
in schaeuffelei (fig. 96), spatha (fig. 98), nasuta (fig.
100) and the subspecies of guestfalica (fig. 105). In
other species of the group the minor ridge is well de-
veloped and both major and minor ridges are fused at
their base, e.g. in saccai (fig. 102), beckeri (fig. 103),
cretensis, the subspecies of appendiculata and in fonta-
na (fig. 108).

In the sistergroup of fontana, viz. the clade astigma
to submaculosa, both ridges are separate. Judged by
the situation in the nearest outgroups of this clade,
this trait can be interpreted as an apomorphy (cf. figs.
109, 110). The species exastigma (fig. 104) also has

the ridges separate, probably as the result of a secon-

DE Jone: Phylogeny Nephrotoma flavescens group

Figs. 57-77. — 57, Nephrotoma exastigma, inner gonostylus, lateral view; 58, N. guestfalica guestfalica, idem; 59, N. guestfalica
hartigiana, idem; 60, N. guestfalica surcoufi, idem; 61, N. quadrifaria quadrifaria, idem; 62, N. appendiculata appendiculata,
idem; 63, N. minuscula, idem; 64, N. sullingtonensis, idem; 65, N. fontana, idem; 66, N. astigma, idem; 67, N. exastigma, out-
er gonostylus, lateral view; 68, N. guestfalica guestfalica, idem; 69, N. appendiculata appendiculata, idem; 70, N. schaeuffelei,
sperm pump, dorsal view, compressor apodeme omitted; 71, 72, N. spatha, 71, sperm pump, lateral view; 72, sperm pump,
dorsal view, compressor apodeme omitted; 73, N. beckeri, idem; 74, N. exastigma, idem; 75, N. appendiculata appendiculata,
idem; 76, N. guestfalica guestfalica, compressor apodeme, anterior view; 77, N. quadrifaria quadrifaria, idem.

249

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 78-100. — 78, Nephrotoma nasuta, sp2, medial view; 79, N. saccai, idem; 80, N. beckeri, idem; 81, N. quadrifaria quad-
rifaria, idem; 82, N. appendiculata appendiculata, idem; 83, N. astigma, idem; 84, N. exastigma, posterior margin of male ster-
nite 8, posterolateral view; 85, N. guestfalica guestfalica, idem; 86, N. quadrifaria quadrifaria, idem; 87, 88, N. guestfalica
guestfalica. 87, male tergite 10, ventral view; 88, male tergite 10, lateral view; 89, 90, N. quadrifaria quadrifaria, 89, male ter-
gite 10, ventral view; 90, male tergite 10, lateral view; 91, 92, N. appendiculata appendiculata, 91, male tergite 10, dorsal view;
92, male tergite 10, lateral view; 93, 94, N. minuscula, 93, male tergite 10, dorsal view; 94, male tergite 10, lateral view; 95,
96, N. schaeuffelei, 95, cercus, lateral view; 96, hypogynial valve, medial view; 97, 98, N. spatha, 97, cercus, lateral view; 98,
hypogynial valve, medial view; 99, 100, N. nasuta, 99, cercus, lateral view; 100, hypogynial valve, medial view.

dary development. (There is in my opinion no com-
pelling reason to consider the dorsal ridge in exastig-
ma and in the clade astigma to submaculosa a new
structure, non-homologous with the major ridge of
the other members of the flavescens group; cf.
Oosterbroek 1980: 336, character 19A and B).

250

48. — Hypogynial valve: (0) minor ridge short (fig. 109);
(1) posteriorly extended (fig. 110).

The minor ridge is elongate in the species lempkei
and submaculosa, reaching towards the tip of the hy-
pogynial valve.

49. — Sternite 9: (0) entire (fig. 112); (1) with break be-

tween its midposterior and anterolateral parts (fig. 111, ar-
rowed).

The species beckeri and cretensis are the only repre-
sentatives of the flavescens group that have a membra-
nous interruption in between the broad anterolateral
portion of sternite 9 and its slender posterior projec-
tion.

50. — Sternite 9: (0) yellowish coloured (fig. 111); (1) an-
teriorly blackish coloured, i.e. heavily sclerotized (fig. 112,
arrowed).

Contrary to the other members of the group, the
anterior margin of sternite 9 is blackish coloured in
the species minuscula and theowaldi.

DISCUSSION OF ADOPTED PHYLOGENY

Phylogenetic analysis of the character state matrix
given in table 2, with a zero-vector as outgroup, resul-
ted in two most parsimonious trees with length 73,
consistency index 68, and retention index 84. I used
exact tree-calculating methods with both computer
programs employed, viz. the ‘ie’ (implicit enumera-
tion) command of HENNIG86 and the branch-and-
bound algorithm of PAUP. The binary coded charac-
ters had the default weight 1. No a priori or a
posteriori adaptations of the data were introduced to
adjust the results of the analysis. Both programs pro-
duced the same pair of equally parsimonious trees.

The two trees obtained differ in the position of the
species flavescens. It either is the sister species of a
clade (astigma (lempkei, submaculosa)) or constitutes a
trichotomy with astigma and the clade (lempkei, sub-
maculosa). In the first tree character state 28(1) is con-
sidered an apomorphy for the clade (astigma (lempkei,
submaculosa)), with subsequent reversal to the plesio-
morphous state in submaculosa. The second tree de-
picts character state 28(1) as a homoplasy in the spe-
cies astigma and lempkei. The second tree is
topologically identical with the strict consensus tree
which is shown in fig. 2.

The four most basal species in the cladogram split
off consecutively. Next in the transition series follows
the pair of sister species beckeri and cretensis.

The remainder of the cladogram contains several
lineages of which the monophyly is only weakly cor-
roborated.

The clade exastigma to submaculosa is substantiated
by a single homoplasy, viz. the narrow anterior apo-
demes of the sperm pump, which are also found in
the species spatha (character 30). The clade contain-
ing the species exastigma and the three subspecies of
guestfalica is also relatively weakly supported by one
homoplasy only (character 28). The character state
involved, viz. the complete reduction of the sclero-
tized part of the ejaculatory duct, also occurs in the

DE Jone: Phylogeny Nephrotoma flavescens group

two subspecies of quadrifaria, as well as in astigma
and lempkei.

The clade quadrifaria to theowaldi is the third line-
age within the cladogram of which the monophyly is
demonstrated by a single homoplasy, in this case the
bare sp2 (character 34). Within the flavescens group
the same character state is known in the species saccai
and beckeri.

In certain aspects the results of the phylogenetic
analysis of the present paper differ considerably from
those presented by Oosterbroek (1980: 333, clado-
gram 2B; see also Oosterbroek & Arntzen 1992). Fig.
113 shows the topology of Oosterbroek’s cladogram
of the flavescens group. For ease of comparison, the
topology of the consensus tree of the present paper is
given in fig. 114.

Oosterbroek (1980: 332) listed three synapomor-
phies in support of the monophyly of the flavescens
group, two of which he considered weak and one
strong. According to his views, the members of the
flavescens group should be distinguished by a straight-
ened dorsal margin of the gonocoxite (‘dorsal margin
of the male sternite nine is flattened’; Oosterbroek
1980: 332, character 10; considered weak). Ooster-
broek noted that this character state is absent in the
species nasuta and spatha. My observations demon-
strate that the posterodorsal edge of the gonocoxite is
curved also in exastigma, the three subspecies of guest-
falica (cf. fig. 38) and in schaeuffelei. I therefore do
not accept the straightened dorsal margin of the gon-
ocoxite as a synapomorphy of the flavescens group.

The other two synapomorphies of the flavescens
group distinguished by Oosterbroek are also used in
the present paper. They are the presence of the mem-
branous ridge underneath the free part of the crest
(Oosterbroek 1980: 332, character 11; considered
weak because of its presumed absence in nasuta and
guestfalica, but see character 22 of the present paper)
and the absence of the rods of the aedeagal guide
(Oosterbroek 1980: 332, character 12; considered
strong; cf. character 11 of the present paper).

Oosterbroek (1980: 332, characters 13 and 14;
346, character 2) distinguished saccai as the sister spe-
cies of a clade containing the remainder of species and
subspecies of the flavescens group. The monophyly of
this latter clade was substantiated by the presence of a
partly trifid aedeagus and the reduction of the sclero-
tized part of the ejaculatory duct.

However, contrary to Oosterbroek’s (1980, 1985a)
assertion, non-flavescens group species with a partly
trifid aedeagus are known within Nephrotoma. East
Palaearctic and Oriental Nephrotoma species showing
this feature include alticrista Alexander, dodabettae
Alexander, inorata Alexander, integra Alexander, me-
dioproducta Alexander, nigrohalterata Edwards, rajah
Alexander, and subpallida Alexander. These species

251

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 101-112. — 101, 102, N. saccai, 101, cercus, lateral view; 102, hypogynial valve, medial view; 103, N. beckeri, hypogy-
nial valve, medial view; 104, N. exastigma, idem; 105, N. guestfalica guestfalica, idem; 106, 107, N. quadrifaria quadrifaria,
106, cercus, lateral view; 107, hypogynial valve, medial view; 108, N. fontana, idem; 109, N. astigma, idem; 110, N. lempkei,
idem; 111, N. beckeri, female sternite 9, dorsal view; 112, N. minuscula, idem.

do not belong to the cornicina group in its current
conception. It should be noted, moreover, that the
majority of the Afrotropical species of Nephrotoma
have the apical section of the aedeagus modified. The
apical portion in these species looks like a ribbon or a
flattened tube with sclerotized lateral margins, much
as in the east Palaearctic and Oriental species with a
trifid aedeagus, but then without the medial filament.
A partly trifid aedeagus, which is also known in spe-
cies of the tipulid subgenera Tipula (Acutipula
Alexander), 7. (Lunatipula Edwards), T. (Odonatisca
Savchenko), 7. (Papuatipula Alexander), and 7.
(Pterelachisus Rondani) could well represent a plesio-
morphy in Nephrotoma. I therefore assume that the
structure of the aedeagus in saccai, which is tubular
throughout, represents an autapomorphy of the spe-
cies (cf. appendix). Likewise, the reduction of the
sclerotized part of the ejaculatory apodeme is evaluat-

252

ed differently in the present study (cf. characters 27
and 28).

In this context it should be noted that Oosterbroek
(1985a: 247) assigned the Japanese species subpallida
Alexander to the flavescens group on account of its
partly trifid aedeagus. When a trifid aedeagus is re-
garded a plesiomorphy on this level of analysis, the
feature - of course - advances no argument to incor-
porate any species in the flavescens group. Moreover,
the male of subpallida has well developed rods of the
aedeagal guide, a plesiomorphous character state not
found in the flavescens group (cf. character 11 of the
present paper). The ovipositor of subpallida has
blunt-tipped, well-developed cerci and large hypogy-
nial valves, both structures thus being of a different,
presumably plesiomorphous, type compared with the
cerci and hypogynial valves of the members of the fla-
vescens group (cf. characters 43 and 45 of the present

text). For these reasons I do not include subpallida in
the flavescens group.

Next in Oosterbroek’s transition series is the spe-
cies nasuta, which was considered the sister species of
the remainder of the flavescens group on account of
the plesiomorphously large anterior apodemes of the
sperm pump (cf. Oosterbroek 1980: 333, character
15). Reduction of the size of the anterior apodemes is
employed in a slightly different manner and at an-
other branching point in the present paper (cf. char-
acter 30).

The sister group of schaeuffelei in Oosterbroek’s
phylogeny is characterized by the serrate hypogynial
valves (cf. Oosterbroek 1980: 333, character 16).
However, incorporated in this clade is the species spa-
tha, a species of which the female sex was unknown to
Oosterbroek in 1980. The female of spatha was de-
scribed by Eiroa & Novoa (1987) based on material
collected in Galicia, Spain. It has the ventral margin
of the tip of the hypogynial valve smoothly curved
(fig. 98) and can therefore not be placed in the clade
distinguished by the serrate hypogynial valve (cf.
character 45 of the present paper). [Oosterbroek &
Artzen (1992) excluded spatha from their cladogram
of the flavescens group].

Figures 113 and 114 both depict the species pair
beckeri and cretensis as the sister group of the remain-
der of species and subspecies of the flavescens group.

According to Oosterbroek’s views, guestfalica and
quadrifaria should be considered sister taxa. This re-
lationship was postulated on account of the reduction
of the hypogynial valves (Oosterbroek 1980: 334,
character 16B), the shape of tergite 10 in the male
(loc. cit.: 336, character 17P) and the presence of
bristles on the alar squama (as ‘neala’; loc. cit.: 336,
character 18). The shape of the hypogynial valves of
both species is very different (cf. figs. 105, 107). The
same holds for the shape of male tergite 10 (cf. figs.
87-90, and characters 37 and 38 of the present
paper). Although probably a plesiomorphy in the
family Tipulidae, the presence of bristles on the squa-
ma is quite unique in the genus Nephrotoma. I have to
assume that these bristles developed independently in
both guestfalica and quadrifaria.

The position of the species fontana in the clado-
gram of Oosterbroek differs noticably from its allot-
ted place in the consensus tree of the present text. The
sister group of fontana is, according to Oosterbroek,
characterized by the absence of the downcurved part
of the minor ridge of the hypogynial valve
(Oosterbroek 1980: 336, character 19). However,
not all members of this ‘sister group’ lack the down-
curved part of the minor ridge, viz. both subspecies of
appendiculata and flavescens (fig. 10). My interpreta-
tion of the position and state of development of the
ridges of the hypogynial valve, which deviates from

DE Jong: Phylogeny Nephrotoma flavescens group

Oosterbroek’s views, is explained under characters 47
and 48 of the present text.

The clade appendiculata to theowaldi of
Oosterbroek fully concords with my ideas on the re-
lationships of the species involved.

The exact phylogenetic relationship of the species
sullingtonensis was left unresolved by Oosterbroek.
On account of the shape of the gonapophysis of the
aedeagal guide it can be considered to be more close-
ly related with the species of the clade astigma to sub-
maculosa (cf. character 9 of the present paper).

According to Oosterbroek, exastigma is closely re-
lated with the species of the clade astigma to submac-
ulosa. He corroborated this postulate by the presence
of both minor and major ridges of the hypogynial
valves (Oosterbroek 1980: 336, character 19A, with
the major ridge assumed to represent a newly devel-
oped structure), the short aedeagus (loc. cit.: 336,
character 23), and the toothless claws in the male (loc.
cit.: 337, character 24). The three characters are dis-
cussed under the respective characters 47, 13 and 1 of
the present text. My argumentation results in a more
basal position of exastigma in the flavescens group.

Oosterbroek’s clade astigma to submaculosa is iden-
tical to the clade astigma to submaculosa presented
here, inclusive of the trichotomy I was unable to re-
solve.

In summary, both our cladograms (figs. 113, 114)
recognize the same basal species in the flavescens
group, with the exception of the species spatha. In
Oosterbroek’s solution spatha occupies a more de-
rived position. The same applies to exastigma, which
is placed next to spatha in Oosterbroek’s phylogeny.
A sister group relationship of guestfalica and quadri-
faria, as suggested by Oosterbroek, was not substan-
tiated in the present study. The species fontana has a
more derived position in my consensus tree. The tri-
chotomy of which sullingtonensis forms part in fig.
113 is resolved in fig. 114. Here sw/ingtonensis is de-
picted as the sister species of the clade fontana.to sub-
maculosa. Both Oosterbroek’s and my solutions agree
on the composition of the clades appendiculata to
theowaldi and astigma to submaculosa. Differences
between figs. 113 and 114 are the result of selection
and interpretation of characters. They do not repre-
sent artefacts of the analytical methods employed, viz.
the ‘manual’ approach of Oosterbroek versus the
computational procedure of the present study.

DISTRIBUTION

In this section I will give a synopsis of the distribu-
tion of the species and subspecies of the flavescens
group. For the time being, more detailed information
can be found in Oosterbroek (1978, 1982, 1985b)
and in Oosterbroek & Theowald (1992).

253

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

saccai
nasuta
schaeuffelei
becker
cretensis
guestfalica
g. hartigiana
g. surcoufi
quadrifaria
q. farsidica
fontana
appendiculata
a. pertenua
minuscula
theowaldi
sullingtonensis
exastigma
spatha
astigma
flavescens
lempkei
submaculosa

Fig. 113. (Left) Topology of the phylogeny of the
Nephrotoma flavescens species group according to
Oosterbroek (1980) and Oosterbroek & Arntzen (1992).

The most basal species in the phylogeny of the fla-
vescens group, schaeuffelei, is known from the Elburz
mountains (Iran) and the Talysch mountains
(Azerbaydzhan) to the south-west of the Caspian Sea.

The species spatha is endemic to north-west Spain.
The species nasuta is known from a few localites in
western, central and eastern Turkey, while saccai oc-
curs in south-west Italy and northern Sicily.

The species pair beckeri and cretensis has an eastern
Mediterranean distribution, with beckeri being dis-
tributed in Cyprus and the adjacent mainland of
Turkey, as well as Lebanon and Israel. Its sister spe-
cies is endemic to Crete.

The species exastigma is confined to northern
Algeria and Tunesia.

The three subspecies of guestfalica are essentially
west Mediterranean, with the nominate subspecies
having the largest range of the three, extending from
England and south Sweden in the northwest, Spain in
the south-west to western Turkey in the east. The
subspecies guestfalica hartigiana is endemic to

254

schaeuffelei
spatha
nasuta
saccai
beckeri
cretensis
exastigma
guestfalica
g. hartigiana
g. Surcoufi
quadrifaria
q. farsidica
appendiculata
a. pertenua
minuscula
theowaldi
sullingtonensis
fontana
astigma
flavescens
lempkei
submaculosa

Fig. 114. (Right) Topology of the consensus tree of the
Nephrotoma flavescens species group of the present paper.

Sardinia, while the subspecies guestfalica surcoufi is
distributed in Morocco, Algeria and Tunesia.

The nominate subspecies of quadrifaria is distrib-
uted over west and central Europe with its range ex-
tending towards the east over the Crimea peninsula
and the Caucasus mountain range. The subspecies
quadrifaria farsidica is known from north-west Iran
in the area adjacent to the Caspian Sea.

The nominate subspecies of appendiculata is large-
ly distributed over west and central Europe, eastwards
reaching towards the westcoast of the Caspian Sea
and southwards to Anatolia. It is absent from south-
west Europe (Spain, Italy) where it is replaced by the
subspecies appendiculata pertenua. In this context the
occurrence of appendiculata sensu stricto on Sardinia
is remarkable. The subspecies appendiculata pertenua
is known from north-west Morocco and south-west
Europe, including Corsica, Sicily and Malta. In its
turn, this subspecies is absent from Sardinia. The spe-
cies minuscula is known from Cyprus and the Levant,
theowaldi occurs in south and south-west Turkey and

in the Greek island Rhodos.

The species sullingtonensis is distributed from
northern Morocco through Spain and France to
southernmost England. The species fontana is en-
demic to Algeria, while astigma is confined to the
north of Algeria and Tunesia. Both flavescens and sub-
maculosa are widespread throughout western and cen-
tral Europe, with submaculosa occurring in Corsica,
Sardinia, Sicily and the mainland of Italy. The species
flavescens is absent in the four last mentioned areas.
The species /empkei is at present known only from the
islands Mallorca and Menorca.

ACKNOWLEDGEMENTS

For the loan of material of both sexes of the species
Nephrotoma spatha\ would like to thank Eulalia Eiroa
(Santiago de Compostella, Spain).

Pjotr Oosterbroek and Frederick Schram, and an
anonymous referee, read and commented upon an
earlier draft of this paper. Although I did not follow
all of their suggestions for adjustment of the text, I
greatly appreciate their willingness to review the
paper. The investigations were supported by the
Foundation for Biological Research (BION), which is
subsidized by the Netherlands Organization for
Scientific Research (NWO).

REFERENCES

Brindle, A., 1960. The larvae and pupae of the British
Tipulinae (Diptera: Tipulidae). — Transactions of the
Society for British Entomology 14: 64-114.

Byers, G. W., 1961. The crane fly genus Dolichopeza in
North America. — University of Kansas Science Bulletin
42: 665-924.

Chiswell, J. R., 1956. A taxonomic account of the last instar
larvae of some British Tipulinae (Diptera: Tipulidae). —
Transactions of the Royal Entomological Society of
London 108: 409-484.

Eiroa, M. E. & F. Novoa, 1987. Descripcién de la hembra
de Nephrotoma spatha (Diptera, Tipulidae). — Eos 63: 19-
20.

Farris, J. S., 1988. Hennig86, version 1.5 + reference. — Port
Jefferson Station, New York.

Hemmingsen, A. M. & B. Jensen, 1972. Egg characteristics
and body size in crane-flies (Diptera: Tipulidae) with
comparative notes on birds and other organisms. —
Videnskabelige Meddelelser fra Dansk Naturhistorisk
Forening 135: 85-127.

Jong, H. de, in press. The phylogeny of the Tipula
(Acutipula) maxima species group, with notes on its dis-
tribution (Diptera, Tipulidae). — Entomologica
Scandinavica 24.

McAlpine, J. F., 1981. Morphology and terminology -
adults. — In: McAlpine, J. F. et al. Manual of Nearctic
Diptera. Volume 1. Research Branch, Agriculture
Canada, Monograph 27: 9-63.

Mannheims, B., 1951. Tipulidae. — Fliegen der palaearktis-
chen Region 15, Lieferung 167: 1-64, pls. I-VII.

DE JONG: Phylogeny Nephrotoma flavescens group

Mannheims, B. & B. Theowald, 1971. Die Tipuliden von
Grönland (Diptera, Tipulidae). — Bonner zoologische
Beiträge 22: 332-342.

Oosterbroek, P., 1978. The western palaearctic species of
Nephrotoma Meigen, 1803, (Diptera, Tipulidae) Part 1. —
Beaufortia 27: 1-137.

Oosterbroek, P., 1979a. The western palaearctic species of
Nephrotoma Meigen, 1803, (Diptera, Tipulidae) Part 2. —
Beaufortia 28: 57-111.

Oosterbroek, P., 1979b. The western palaearctic species of
Nephrotoma Meigen, 1803 (Diptera, Tipulidae) Part 3. —
Beaufortia 28: 157-203.

Oosterbroek, P., 1979c. The western palaearctic species of
Nephrotoma Meigen, 1803 (Diptera, Tipulidae) Part 4,
including a key to the species. — Beaufortia 29: 129-197.

Oosterbroek, P., 1980. The western palaearctic species of
Nephrotoma Meigen, 1803 (Diptera, Tipulidae), Part 5,
Phylogeny and Biogeography. — Beaufortia 29: 311-393.

Oosterbroek, P., 1982. New taxa and data of western
Palaearctic Nephrotoma (Diptera: Tipulidae). -
Entomologische Berichten, Amsterdam 42: 41-44.

Oosterbroek, 1984. A revision of the crane-fly genus
Nephrotoma Meigen, 1803, in North America (Diptera,
Tipulidae). Part Il: non-dorsalis species-groups. —
Beaufortia 34: 117-180.

Oosterbroek, P., 1985a. The Nephrotoma species of Japan
(Diptera, Tipulidae). — Tijdschrift voor Entomologie
127: 235-278.

Oosterbroek, P., 1985b. Some Tipulidae new for Corsica
and Sardinia (Diptera). — Entomologische Berichten,
Amsterdam 45: 121-122.

Oosterbroek, P. & J. W. Arntzen, 1992. Area-cladograms of
Circum-Mediterranean taxa in relation to Mediterranean
palaeogeography. — Journal of Biogeography 19: 3-20.

Oosterbroek, P. & B. Theowald, 1992. Family Tipulidae. —
Catalogue of Palaearctic Diptera 1: 56-178.

Savchenko, E. N., 1973. Crane flies (fam. Tipulidae), sub-
fam. Tipulinae (conclusion) and Flabelliferinae. — Fauna
SSSR (N.S. 105) Two-winged insects II (5): 1-281. (In
Russian).

Swofford, D. L., 1993. PAUP, Phylogenetic Analysis Using
Parsimony, version 3.1. — Illinois Natural History Survey,
Champaign, Illinois.

Tangelder, I. R. M., 1983. A revision of the crane fly genus
Nephrotoma Meigen, 1803, in North America (Diptera,
Tipulidae). Part I: the dorsalis species-group. — Beaufortia
33: 111-205. _

Tangelder, I. R. M., 1984. The species of the Nephrotoma
dorsalis-group in the Palaearctic (Diptera, Tipulidae). —
Beaufortia 34: 15-92.

Tangelder, I. R. M., 1985. Phylogeny of the Nephrotoma
dorsalis species-group (Diptera, Tipulidae), mainly based
on genital characters. — Beaufortia 35: 135-174.

Theowald, B., 1957. Die Entwicklungsstadien der
Tipuliden (Diptera, Nematocera), insbesondere der west-
palaearktischen Arten. — Tijdschrift voor Entomologie
100: 195-308.

Theowald, B., 1967. Familie Tipulidae. (Diptera,
Nematocera). Larven und Puppen. =
Bestimmungsbücher zur Bodenfauna Europas 7: 1-100.

Received: 12 May 1993
Accepted: 27 August 1993

295)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

APPENDIX

In this section the recognized autapomorphies of
the species and subspecies of the Nephrotoma flaves-

cens group are listed.

appendiculata appendiculata: no autapomorphies recog-
nized.

appendiculata pertenua: differing from nominal subspecies in
details of male tergite 10 only.

astigma: shape of lateral shells of male tergite 10 in lateral
view.

beckeri: shape of male tergite 10; shape of inner gonostylus.

cretensis. shape of compressor apodeme of sperm pump.

exastigma: lateral prescutal stripes straight; inner gonostylus
with sclerotized ridge from anterior to lateral part.

flavescens medial prescutal stripes with dull lateral margins,
downcurved part of lateral prescutal stripes dull.

fontana: shape of extension of male sternite eight; shape of
gonapophysis of aedeagal guide.

guestfalica guestfalica: position and shape of crest of inner go-
nostylus.

guestfalica hartigiana: crest of inner gonostylus partly re-

duced.

256

guestfalica surcoufi: crest of inner gonostylus entirely re-
duced.

lempkei: medial projection on male sternite eight slender,
somewhat tapering towards tip.

minuscula: aedeagal guide with dorsolaterally extending
ridges; form and length of central ridges of male tergite
10.

nasuta: nasus long; shape of male tergite 10; shape of aedea-
gus, particularly at branching point.

quadrifaria quadrifaria: shape of inner gonostylus; shape of
male tergite 19.

quadrifaria farsidica: medial projection of male sternite eight
directed dorsally.

saccat: shape of male tergite 10; shape of gonapophysis of ae-
deagal guide; sperm pump posteriorly with upcurved
plate; aedeagus single throughout; genital fork reduced.

schaeuffelei: shape of inner gonostylus; outer gonostylus an-
teriorly with pubescent incurved swelling; shape of sp2.

spatha: hairs on vertex short; shape of male tergite 10.

submaculosa: no autapomorphies recognized.

sullingtonensis. prescutal stripes partly confluent.

theowaldi: aedeagus near branching point with pair of later-
al extensions; crest of inner gonostylus twisted along its
longitudinal axis.

A. NEBOISS

Museum of Victoria, Melbourne, Australia

NEW SPECIES OF THE GENUS MOLANNA CURTIS
FROM SULAWESI (TRICHOPTERA: MOLANNIDAE)

Neboiss, A., 1993. New species of the genus Molanna Curtis from Sulawesi (Trichoptera:
Molannidae). — Tijdschrift voor Entomologie 136: 257-258, figs. 1-4 [ISSN 0040-7496].

Published 10 December 1993.

The description and figures of Molanna jolandae sp. n. are given. This is the second species re-
corded from Sulawesi, and extends the range of generic distribution to southern Sulawesi

Tenggara.

A. Neboiss, Department of Entomology, Museum of Victoria, 71 Victoria Crescent,
Abbotsford 3067, Victoria, Australia.
Key words. — Trichoptera; Molannidae; Molanna; Sulawesi; taxonomy.

The present paper is based on material recently
collected in Sulawesi. Among the numerous speci-
mens from a number of localities, members of the
family Molannidae are scarce and have been collected
at only one location. There is only one previous
Sulawesi record, a single specimen on which the de-
scription of Molanna cupripennis Ulmer (1906) is
based. The type locality ‘Bonthain Peak’ is in south-
western Sulawesi (Sulawesi Selatan). The present ma-
terial is from the southeastern part of the island
(Sulawesi Tenggara) near Moramo (4°09’S
122°38’E). It may be noted, that the extensive collec-
tion from Dumoga Bone National Park in the north
of the island (Sulawesi Utara), accumulated by the
Wallace Expedition 1985, did not contain any speci-
mens of Molanna.

MATERIAL AND METHODS

Specimens were prepared by clearing the abdomen
in cold KOH solution, then transferring to glycerol
for drawing and later storage in microvial. Drawings
of genitalia were made with the assistance of a camera
lucida on a Wild M20 compound microscope. The
wings, prepared as temporary microscope mounts in
glycerol, were examined and drawn using a Wild M8
dissecting microscope also equipped with camera lu-
cida. All dissected and figured specimens are identi-
fied by the author’s notebook number with prefix
‘PT-.... All specimens have been deposited in the
National Museum of Natural History, Leiden, The
Netherlands (RMNH).

‘TAXONOMIC PART

Molanna jolandae sp.n.
(figs. 1-3)

Type material. — Holotype d: Sulawesi Tenggara,
Moramo (4°09’S 122°38’E), Sungai Sena, 50 m asl, 15 Nov
1989, R. de Jong and J. Huisman (site S 8946: mixed forest,
dense undergrowth) in RMNH [Genitalia prep. PT-1968 fig-
ured, specimen dry mounted]. — Paratypes: Sulawesi
Tenggara, Moramo, Sungai Moramo, 200 m asl, 16 Nov.
1989, at light, R. de Jong and J. Huisman, 26 (site S 8947:
mixed forest, dense undergrowth) in RMNH [wing prep. PT-
1969 figured; both specimens in alcohol].

Description

Wings slender, rounded apically (fig. 3), length d
8.6-9.0 mm, densely covered with short, dark brown
pubescence, hind wing anal margin with long, dark
brown fringe, venation similar to that of Molanna
cupripennis Ulmer.

Male. — Dark, blackish brown, antennae, maxillary
and labial palpi as well as fore legs densely covered
with coarse blackish brown hairs, mid- and hind-tarsi
also covered with dark hairs, whereas femora and tib-
iae are pale yellow with scattered small, dark blackish
semi-erect spines, apical spurs covered with dark
hairs, preapical spurs with paler hairs.

Genitalia (figs. 1, 2) with abdominal segment IX in
lateral view broadly rounded ventrally, mid-dorsal
margin broad with slight mesal depression, anterolat-
eral angles may be more or less produced. Superior
appendages elongate ovoid, flattened, lower basal
margin produced into an incurved lobe. Segment X
large, extended and strongly curved ventrad, dorsom-
esally separated almost to the base, usually with three
strong spines on dorsal margin on either side. Inferior

257

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-4. Molanna species. — 1, M. jolandae sp. n., male genitalia (holotype), lateral; 2, ventral; 3, wing venation (paratype);
4, M. cupripennis Ulmer, male genitalia lateral (from Ulmer 1951).

appendages short, two branched, lower branch with
terminal half curved dorsad, a distinct spine apically,
upper branch directed posteriorly, slightly concave,
apex rounded. Phallus short, rather robust, strongly
curved, membranous, small chitinous spine internal-
ly.

Female unknown.

Remarks

This species is very similar to Molanna cupripennis
Ulmer, but, besides being smaller (length of forewing
9 mm instead of 14 mm), differs by the presence of
spines on segment X, and the shape of the inferior ap-
pendages, the branches of which are not incised api-
cally. The figures by Ulmer (1906, 1951) (here repro-
duced as fig. 4) also show notable differences in shape
and position of segment IX and the superior appen-
dages.

Etymology. — The species is named after Jolanda
Huisman in recognition of her efforts gathering ex-

258

tensive material of Trichoptera from many localities
in Sulawesi.

ACKNOWLEDGEMENT

I wish to express my sincere thanks to Jan van Tol
of RMNH Leiden for access to this most interesting
material.

REFERENCES

Ulmer, G., 1906. Neuer Beitrag zur Kenntnis aussereu-
ropaische Trichopteren. — Notes from the Leyden
Museum 28: 1-116.

Ulmer, G., 1951. Kôcherfliegen (Trichopteren) von den
Sunda-Inseln (Teil 1). — Archiv fiir Hydrobiologie
(Supplement) 19: 1-528 + plate 1-28.

Received: 16 March 1993
Accepted: 15 April 1993

NICO NIESER' & PING PING CHEN’

1 Tiel, The Netherlands

2 Beijing Academy of Agriculture & Forestry Sciences, Beijing, P. R. China

THE RHAGOVELIA (HETEROPTERA: VELIIDAE) OF

SULAWESI (INDONESIA)

Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), IV.

Nieser, N. & P. P. Chen, 1993. The Rhagovelia (Heteroptera: Veliidae) of Sulawesi
(Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), IV. — Tijdschrift
voor Entomologie 136:259-281, figs. 1-84 [issn 0040-7496]. Published 10 December 1993.
Apart from additional records for several species for Sulawesi, ten new species are described:
Rhagovelia blagiokommena sp. n., Rhagovelia chrysomalla sp. n., R. daktylophora sp. n., R. horaia
sp. n., À. pseudocelebensis sp. n., R. robina sp. n., R. trichota sp. n., R. tropidata sp. n. from
Sulawesi, À. kastanoparuphe sp. n. from Sulawesi and Buton and R. kalami sp. n. from Buton.
A revised key to species from Sulawesi and Buton is given. Additional records for Rhagovelia
from Sabah are given in an appendix.

Correspondence: Dr. N. Nieser, Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands.

Key words. — Sulawesi (= Celebes); Buton; Sabah; Veliidae; Rhagovelia, key; new species.

As our contribution to the Fauna Malesiana pro-
gram, we are working on the Nepomorpha and
Gerromorpha mainly collected by staff members of
the Zoological Museum Amsterdam and the
National Museum of Natural History Leiden. The
present one is the fourth in the series, the first dealt
with Naucoridae, Nepidae and Notonectidae (Nieser
& Chen 1991), while the second and third treated
the Gerridae (Nieser & Chen 1992, Chen & Nieser
1992)

Rhagovelia is a large tropicopolitan genus of small,
stream inhabiting, Veliidae. About 150 species have
been described so far, but the number of extant spe-
cies is probably twice as high. The subfamily
Rhagoveliinae is characterized by a large, deeply cleft,
apical segment of the middle tarsus with a fan-like
structure (fig. 1). This is considered an adaption to li-
ving on waters with a strong current. Two Oriental
genera are included in the subfamily, viz. Tetraripis,
with both middle and hind tarsi with cleft apical seg-
ment provided with a fan-like structure, and
Rhagovelia, with only middle tarsi modified in this
way. Tetraripis is only known from Sri Lanka and
western Malaysia.

Formerly, it was thought that tropical America
would be much richer in species of Rhagovelia than
Asia (Bacon 1956, Hungerford & Matsuda 1961,
Lundblad 1936, 1937). In a recent revision, however,
Polhemus & Polhemus (1988) described 26 new spe-
cies from Borneo, Sulawesi and Maluku (the Moluc-
cas). Only two species were previously known from

this area. They stated, moreover, that they had many
more undescribed Indo-Australian species in their
collection. Thus the apparent paucity of species in
tropical Asia is at least partly due to insufficient
knowledge of the fauna.

Although a complete survey of the Rhagovelia of
Sulawesi was included in the revision by Polhemus &
Polhemus (1988), we recognised again ten new spe-
cies in the collections we studied, thus doubling the
number of known species. As very probably more spe-
cies are still to be discovered, the key to species from
Sulawesi and Buton presented here is to be used with
care. New records of this genus for Sabah are presen-
ted in appendix 1.

Measurements are in millimetres, length has been
measured in dorsal view from the anterior margin of
the head along the central axis to the posterior margin
of the last visible abdominal tergite in apterous, and
to the apex of the hemielytra in macropterous speci-
mens. ‘Width’ has been measured at the base of the
connexiva in dorsal view. The first two segments of
the fore tarsi have not been measured, as they are
small (0.02-0.03), usually partly hidden in an excava-
tion of the fore tibia and thus difficult to measure pre-
cisely. Moreover, they do not provide specific charac-
ters. For the same reason the measurements of the
first two segments of the hind tarsi have been omit-
ted.

The seventh abdominal segment, on which the
connexiva end, is the last before the modified genital
segments. In females the eighth tergite is sometimes

259)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 1-16. — 1, diagrammatical drawing of the fan like structure in the cleft apical segment of intermediate tarsi of
Rhagoveliinae. — 2-6. R. blogiokommena paratypes, 2-4 male, 2 apex of abdomen, lateral view, 3 paramere, 4 hind femur; 5-6
female, 5 apex of abdomen, lateral view, 6 right half of thorax and abdomen in dorsal view. Scales 2,4-6 1 mm, 3 0.25 mm.
—7-11. R. chrysomalla paratypes; 7-9 male, 7 apex of abdomen, lateral view, 8 paramere, 9 hind femur; 10-11 female, 10 ap-
ex of abdomen, lateral view, ‘7’ and ‘8’ indicate 7th and 8th (gonocoxa) segment, 11 right half of thorax and abdomen in dor-
sal view, ‘7’ and ‘8’ indicate tergites 7 and 8. Scales 7,9-11 1 mm, 8 0.25 mm. — 12-16. R. daktylophora paratypes; 12-14 male,
12 apex of abdomen, lateral view, 13 paramere, 14 hind femur; 15-16 female, 15 thorax and abdomen, lateral view, 16 right
half of thorax and abdomen in dorsal view. Scales 12,14-16 1 mm, 13 0.25 mm.

260

more or less horizontal (fig. 11) in which cases the
eighth tergite is the last visible in dorsal view. In other
species the eighth tergite is pointing downward (fig.
34), in which case the seventh tergite is the last one
fully visible in dorsal view. Another character that
may give some trouble is the presence of minute den-
ticles on the body, especially in greyish black species.
Good optics with a magnification of at least 30 X and
a good light source is needed to study this character.
These denticles are more difficult to distinguish in
wet than in dry specimens. They are often most dis-
tinct on the proepisterna, which are the sclerites ante-
rodorsally of the anterior coxae, on both sides of the
rostrum.

The species dealt with in this paper were mainly
collected by J. P. Duffels (ZMA), N. Nieser (NC), J.
van Tol (RMNH) and G. Zimmermann (NC/ZC) on
several expeditions to Sulawesi, Buton and Sabah
(Eastern Malaysia). Unless otherwise indicated speci-
mens are deposited in the collections mentioned in
brackets after the collectors. Some reference speci-
mens have been sent to the additional collections
mentioned in following list: Museum Zoologi Bogor,
Bogor, Jawa (MBBJ); Bagian Pertanian, Universitas
Haluoleo, Kendari, Sulawesi (BPUH); J. T. Polhemus
collection (University of Colorado Museum),
Englewood, Co. U. S. A. (PC); Nieser collection,
Tiel, The Netherlands (NC); Chen collection, Beijing,
P. R. China (PCHC); National Museum of Natural
History, Leiden, The Netherlands (RMNH); G.
Zimmerman collection, Marburg, B. R. D. (zc);
Zoölogisch Museum, afd. Entomologie, Amsterdam,
The Netherlands (ZMA) and Zoological Museum,
Copenhagen, Denmark (ZMC). Some additional in-
formation on localities can be found in Nieser &

Chen (1991).

ACKNOWLEDGEMENTS

Thanks are due to Drs J. P. Duffels, J. van Tol and
G. Zimmermann for putting specimens in their care
at our disposal and to Dr. J. T. Polhemus for making
paratypes of most of the species described by
Polhemus & Polhemus (1988) available for study.

SYSTEMATIC PART

Key to Rhagovelia of Sulawesi (apterous specimens)
1. Pronotum short, its length much shorter than
SMT DES ce
ND aide os R. samarinda Polhemus & Polhemus
[Described from E Kalimantan and Sarawak, one
macropterous female has been reported from
Lore Lindu N. P. If a species with short pro-
notum is found, it may well represent a new spe-
cies]

12,

NIESER & CHEN, Rhagovelia of Sulawesi

Pronotum long, covering, all or nearly all of mes-
ONOMIME NE 2
Dorsum of pronotum and abdomen covered by
striking appressed golden pubescence. Transverse
yellowish-orange band anteriorly on pronotum
separated from yellowish prosternum by variable
and sometimes indistinct dark brown spots be-
hind the eyes. [Large species, length 3.8-4.3 mm]
CIR denke anne R. chrysomalla sp. n.
Dorsum of pronotum and abdomen not covered
by striking appressed golden pubescence .......... 3

. Transverse yellowish-orange band anteriorly on

pronotum narrow, laterally at most reaching half-

Waythelgyes®. onere na 4
Transverse yellowish-orange band anteriorly on
pronotum wide, reaching laterally of eyes … … 20
Males wann cata Seber seh tue >
Bemalessn cn PR na 12
Abdominal sternite 7 (sixth visible ventrally) not
longitudinally carinate (fig. 56) nnen 6

Abdominal sternite 7 longitudinally carinate ....7

. Dorsum of abdomen hirsute; apex of paramere

ENE (HE) een R. trichota
Dorsum of abdomen with normal pubescence,
not hirsute; apex of paramere pointed (fig. 75) ...
Silicone POOR: MN RON en RER R. hamjadi
[Only known by the type series from Lore Lindu
N. P. Polhemus & Polhemus 1988: 194-195,
figs. 134-140]

Abdominal sternite 7 with a sharply pointed tuft
of stiff setae, looking at first sight like a short
stout peg, at basal third of longitudinal median
carina (fig. 68). Hind femur inflated to a variable
extend, at least twice as wide as middle femur ...
7 Don haa en Deer wiles rate tas antes à R. pruinosa
Abdominal sternite 7 without peg like tuft of
setae, hind femur 1.5 times or less as wide as mid-

dlefemunka tee tn en 8
Abdominal sternites 4-6 with thick long golden
yellowish pilosity (fig. 35)... 9
Abdominal sternites 4-6 with normal inconspi-
cuous pubescence .......... R. kastanoparuphe sp. n.
Abdominal tergite 7 about as long as its basal
width Aten rh heh os een ite 10
Abdominal tergite 7 longer (about 1.4x) than its
bald ee th eee 10

. Paramere apically broad, leaf-like (fig. 36, 43-44)

ara ee aa R. pseudocelebensis sp. n.
Paramere apically less broad (fig. 41-42)
hte a NT, eretico mic. has agi R. celebensis

. Sternites 5-7 with ventrolateral impressions resul-

ting in a broad median keel longitudinally on the-

se,sternites (tie PD) a R. daktylophora
Ventrolateral impressions and keel restricted to
sterne Ale R. sulawesiana

Hind femur dorsoventrally flattened, lower mar-

261

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 17-29. — 17-20. R. horaia; 17-19 male, 17 apex of abdomen of holotype, lateral view, 18 paramere of paratype, 19 hind
trochanter, femur and tibia of holotype; 20 female paratype, apex of abdomen, lateral view. Scales 17, 19, 20 1 mm, 18 0.25
mm. — 21-23. Right half of proctiger of males; 21 R. horaia paratype, scale 0.25 mm, 22 R. grayi, 23 R. lorelinduana, 22 &
23 drawn after J. & D. Polhemus (1988). — 24-29. R. kalami paratypes; 24-26 male, 24 apex of abdomen, lateral view, 25 pa-
ramere, 26 hind trochanter, femur and tibia; 27-29 female, 27 hind trochanter, femur and tibia, 28 right half of thorax and
abdomen, dorsal view, 29 apex of abdomen, lateral view. Scales 24, 26-29 1 mm, 25 0.25 mm.

262

13.

17e

18.

2D

gin more or less straight (fig. 78) … R. hamjadi
Hind femur not flattened dorsoventrally, with a
convex lower margin (fig. 77)
Connexiva margined with yellowish brown and
set with long setae (longer than width of hind ti-
bia) along entire length, converging caudally but
Bon folded.over territe 7 .........e sen R. unica
[Known only by the unique holotype female
from Dumoga Bone N. P., Polhemus & Polhe-
mus 1988: 195, figs. 43-45]

Combination of characters not as above ......... 14

. Tergite 8 more or less horizontal, in the same pla-

me as GEE nen ne R. pruinosa
Abdominal tergite 8 distinctly directed ventrad...
RE oe eh RF ont AIRE A A 15

. Caudal part of connexiva folded over abdominal

tergite(s), at least covering most of tergite 7 ...16
Connexiva not folded over abdominal tergites . ..

. Connexiva strongly constricted on segments 4

and 5, more or less parallel and nearly meeting on
segments 6 and 7 (fig. 61). Tergite 7 with a me-
diocaudal finger-like process ........ R. sulawesiana
Connexiva constricted on segment 4, convergent
caudally and meeting dorsally over posterior part
of tergite 7 which lacks a mediocaudal finger-like

PEO GCESSM LTE OO) Linn R. trichota
Connexiva converging caudally in a straight or
aid Sa NES. re 18

Connexiva distinctly constricted at the suture be-
tween segment 6 and 7, tergite 7 with a distinct
mediocaudal upwardly directed finger-like pro-
CESS (asl OI) AES R. daktylophora
Upper margin of connexiva brownish, their cau-
dal apices long, tergite 7 with a more or less trian-
gular hind margin (figs. 33, 34)
o iena R. kastanoparuphe
Upper margin of connexiva blackish, caudal api-
ces short, hind margin of tergite 7 either truncate
or with a mediocaudal finger-like projection (figs.

EON des 19

. Hind margin of tergite 7 truncate, straight .........

PR e ne ee R. celebensis
Hind margin of tergite 7 with a mediocaudal fin-

ger-like projection . R. pseudocelebensis
„Melles benede Ilio AT 21
Reese RSR er re 29

. Posterior tibia straight or softly curved, without

larger subapical tooth in inner row (fig. 64) ....22
Posterior tibia curved, usually distinctly, with one
or more larger teeth subapically in inner row
(figs. 19, 26)
Metanotum and first abdominal tergite orange-li-
ke to light reddish brown. Posterior femur relati-
vely slender, about 3.5 times as long as wide
a ee I IRE SERRE à R. robina

DI

24.

29:

26.

27.

28

29)

30.

al.

NIESER & CHEN, Rhagovelia of Sulawesi

Metanotum and first abdominal tergite dark.
Posterior femur more inflated, three times or
slightly less as long as wide
Sternite 7 laterally compressed, resulting in a
strongly developed ventral keel, which is accentu-
ated by a well-developed fringe of cilia (fig. 62) ..
rradeina licia lett R. tropidata
Sternite 7 ventrally flattened, with a low median
keel
Abdominal sternites 5-6 with postero-ventrally
directed tufts of stiff setae (fig 71), paramere as in
GINT GO RL EDER NOLI RIM DI R. wallacei
Abdominal sternites 5-6 with normal pubescence
only, without postero-ventrally directed tufts of
stiff setae (fig 69), parameres as in figs. 3, 75 ...25
Minute black denticles present on pro- and meso-
pleura, mesopleura without coarse punctation ....
TOI Stia RE TEN STE R. minahasa
Minute black denticles absent on pro- and meso-
pleura, mesopleura with coarse punctation .........
MAL PTT AID ANNE 1 R. blogiokommena
About 30 minute teeth in row proximal to large
spine on posterior femur ........................ R. grayi
[Only known by the type series from Sulawesi
Utara, Lake Mala near Mokabang, Polhemus &
Polhemus 1988: 207-208, figs. 208-215]

About 20 minute teeth in row proximal to large
spine on posterior femur
Sternite 7 ventrally halfway with a tuft of caudal-
lidirecedisemel(tip PA en ne R. kalami
Sternite 7 ventrally halfway without a tuft of cau-
dally directed setae
Posterior femur nearly three times as long as wi-
de, proctiger as in fig. 23 ............ R. lorelinduana
[Only known by the type series from Lore Lindu
N. P. in Sulawesi Tengah, Polhemus & Polhe-
mus 1988: 208, figs. 141-148]

Posterior femur about 2.5 times as long as wide,
PIOGL PE HAS MAO E R. horaia
Connexiva caudally folded over abdomen at least
on tergite 7
Connexiva convergent posteriorly but not folded
OE EHE 7 non e 2er 32
Minute black denticles distinctly and abundantly
extending onto thoracic pleura. Connexiva
strongly constricted on segment 5 and conver-
ging in a more or less straight line caudally of
constriction. Caudal apices of connexiva small,
trlangulârd ster me Se RRs: ne R. minahasa
Minute black denticles not distinctly extending
onto thoracic pleura. Caudal apices of connexiva
astenia on. 31
Connexiva only slightly constricted on segments
4/5, more or less straightly convergent caudally.
Middle femur not dorsoventrally flattened .........
PR Re eee R. grayi

263

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

— Connexiva strongly constricted on segment 5,
curved outward again on segment 6. Middle fe-
mur dorsoventrally flattened ....... R. lorelinduana

32. Outer lateral margin of connexiva distinctly con-
stricted on segment 4 (figs. 52, 67) ................ 33

— Outer lateral margin of connexiva more or less
SUIS TE Ts. nen ee ke! 35

33. Meso- and meta-thoracic pleura yellow to pale
orange-like, densely set with minute black denti-
cles. Metanotum caudolaterally with small swel-
lings (fiom EAR ae Mis Se he R. robina

- Meso- and meta-thoracic pleura largely dark with
at most a few minute black denticles. Metanotum
without a pair of caudolateral swellings (fig. 67)

34. Length over 3 mm, connexiva swollen and slight-
ly curved inward on segments 4-5 but not sinuate
(fig. 67), mesopleuron without coarse punctation
hace ee R. tropidata

— Length just under 3 mm, connexiva slightly sinu-
ate and hardly swollen on segments 4-5 (fig. 6),
mesopleuron with coarse punctation ..................
ee ern ae R. blogiokommena

35. Connexiva in dorsal view slightly thickened on
segment 4. Abdominal tergite 7 only slightly lon-
ger than! basall width... R. kalami

— Connexiva not thickened on segment 4. Abdom-
inal tergite 7 over 1.25 times as long as its basal
Width a Me ns deo ea 36

36. Generally a dark grey brown species, yellow
transverse band on pronotum on median line less
than one fifth the median length of pronotum.
Length of body about 3.2, ratio length to maxi-
TALIA CIO ILO ee R. wallacei

— Generally a reddish brown species, yellow trans-
verse band on pronotum on median line more
than one fifth the median length of pronotum.
Length of body about 3.4, ratio length to maxi-
muniwidthel74.. Re R. horaia

Rhagovelia celebensis Polhemus & Polhemus

(figs. 41-42, 81)

Rhagovelia celebensis Polhemus & Polhemus, 1988: 199-
200, figs. 179-186, 229.

Material. — SULAWESI UTARA: Dumoga Bone N.P., Base
Camp, bridge, 16.x.1985 103d 729 (5d 19 macr.); same,
flussabw. Brücke (downstream bridge), 22.x.1985, 2d 39;
same, Base Camp, Stausee (‘barrage lake’), 22.x.1985, 6d
109 (12 macr.); same, Tumpah R.(type locality), Staustufe
(barrage dam) 23.x.1985, 17¢ 189 (36 macr.); same,
Tumpah R., quiet part, 28.x.1985, 34 39; Tumpah R.
beach, 19.x.1985, 36d 369 (1d macr.); Tumpah, Water-
fall oberh. Seitenbach (upstream tributary), 21.xi.1985, 1d
12; Südküste, Strasse, Bach, (South coast, road, stream),
18.xi.1985, 8d 139; all leg. G. Zimmermann; Dumoga
Bone N. P. Tumpah river near confluence Toraut, UTM-

264

XL00631, 210 asl., 18.v.1985, 28 29, same, 23.v.1985,
106 49 (19 macr.); Dumoga Bone N. P., waterfall creek,
tributary of Tumpah, UTM-XL0064, c. 225m asl.,
24.v.1985, 29; same, hygropetric zone of Waterfall Creek,
22.iv.1985, 1d 19; Dumoga Bone N.P., Edward’s sub-
camp, UTM-WL9365, 664m asl., 3.vi.1985, 59 (1 macr.);
all leg. J. van Tol; 20 km E of Kotamobagu, Mt Ambang,
alt. 1210 m, 8.xi.1985, multistr. evergreen forest, soil litter
sifted, 19, leg. J. Krikken. — SULAWESI TENGAH, 50 km SE
of Palu, Lore Lindu N. P., sur. Dongi Dong: shelter, 1°13’S
120°11’E, tributary of Sopu river, edge of multistr. ever-
green forest, 950 m, 4.xii.1985-C, 16 19; Palu, Sopu river
nr. Dongi Dongi, SJ 85, 950m, 5.xii.1985, 59; same,
8.xii.1985, 16, J. van Tol. — SULAWESI SELATAN, c. 10 km
NW Palopo (km 15 along road Palopo-Rantepao), Salo
Tandung, 2°58’S 120°07’E, 300-400 m asl., width 10m,
boulders, torrents and seepage areas, open sec. forest,
27.iv.1991, 3d 109, leg. S. Kofman, 91JvT15; Mamasa,
2°56°S 119°22’E, 1050m, cultivated area, river flowing
through village, rather fast flowing, bottom with boulders
and some coarse sand, 8. IV. 1991, leg. J. van Tol, 26 29.

Distribution. — This species is widespread in
Sulawesi (fig. 81). Its type locality is Dumoga
National Park, Tumpah River.

Rhagovelia minahasa Polhemus & Polhemus

(figs. 70, 73-74, 84)

Rhagovelia minahasa Polhemus & Polhemus 1988: 205-
206, figs. 216-222, 228.

Material. — SULAWESI UTARA: Lakes, Bach unter Sulphur
spr., Z. Dorf, 21.xi.1985, 48d 459 (5d 69 macr.), many
larvae; Dumoga Bone N. P., Tumpah River, 19. X. 1985,
19, leg. G. Zimmermann. Apterous, unless otherwise indi-
cated.

Distribution. — Only known from the eastern part

of North Sulawesi (fig. 84).

Rhagovelia pruinosa Polhemus & Polhemus

(figs. 68, 77, 83)

Rhagovelia pruinosa Polhemus & Polhemus, 1988: 195-196,
figs. 149-157, 229.

Material. - SULAWESI SELATAN: E of Maros, Bantimurung
Area, Sg. Pattunuang Asue (type locality), 5°03’S 119°41’E,
fast flowing stream, ponded areas, through disturbed forest
on limestone, 23.iv.1991, J. van Tol, 68 82 (26 29
macr.). — SULAWESI TENGAH: Luwuk area, Sungai Batui,
Singsing Camp, 19.x.1989, 18 macr.; Sungai Biak, 22.x.
1989, J. van Tol, 26. — Sulawesi Tenggara: Sungai
Mowewe, 28.11.1989, N8922, 38; marsh S of Pomalaa,
2.iii.1989, N8930 36 119; 20 km E Kolaka, mountain
stream, 3.iii.1989, N8934 1 6 , leg. N. Nieser; Konaweha R.
near Sanggona, 7.xi.1989, J. van Tol, 48 109. — BUTON.
Stream N of Baubau, 8.iii.1989, N8935 48 6%, leg. N.
Nieser. Apterous unless otherwise indicated.

Distribution. — Southern and eastern parts of

Sulawesi, Buton (fig. 83).

NIESER & CHEN, Rhagovelia of Sulawesi

Table 1. Measurements of leg segments in Rhagovelia
Tarsal segments 1 and 2 of fore and hind legs have not been measured for most species. When not given, they measure to-
gether 0.05-0.07 and 0.12-0.15 respectively and do not seem to provide specific characters.

femur tibia tars 1 tars2 tars3
R. blogiokommena 3
fore leg 0.80 0.82 = - 0.22
middle leg 1.30 1.05 0.06 0.40 0.62
hind leg 1.20 1.06 = = 0.23
R blogiokommena 2
fore leg 0.81 0.82 - = 0.22
middle leg 1.29 1.01 0.07 0.40 0.61
hind leg 1.11 1.10 = — 0.23
R. chrysomalla 3
fore leg 1.23 1.28 = = 0.30
middle leg 2.00 1.62 0.07 0.72 0.81
hind leg 1.44 1.72 = - 0.38
R. chrysomalla 2
fore leg 1.21 1.25 = = 0.29
middle leg 1.86 1.60 0.08 0.75 0.85
hind leg 1.41 1.73 — - 0.36
R. daktylophora 3
fore leg 0.93 1.01 = = 0.25
middle leg 1.58 1.26 0.07 0.57 0.71
hind leg 1.32 1.41 = = 0.30
R. daktylophora 9
fore leg 0.87 0.98 = = 0.27
middle leg 1.49 1.16 0.07 0.50 0.71
hind leg 122 1.39 = — 0.31
R. horaia 3
fore leg 1.04 1.09 = = 0.30
middle leg 1.60 1.37 0.08 0.49 0.69
hind leg 1.67 1.49 = — 0.40
R. horaia ®
fore leg 1.00 1.03 = = 0.28
middle leg 1.55 1.26 0.007 0.47 0.70
hind leg 1.37 1.43 = = 0.40
R. kalami &
fore leg 0.94 0.97 = = 0.22
middle leg 1.52 1.20 0.08 0.50 0.69
hind leg 1.39 1.20 = = 0.30
R. kalami ?
fore leg 0.93 0.95 = = 0.22
middle leg 1.51 1.18 0.08 0.53 0.69
hind leg 1.31 1.23 = — 0.31
R. kastanoparuphe 3
fore leg 0.93 1.02 = = 0.26
middle leg 1.63 1.28 0.08 0.56 0.70
hind leg 1.47 1.39 — = 0.35
R. kastanoparuphe ?
fore leg 0.95 1.03 = = 0.25
middle leg 1.58 1.23 0.08 0.54 0.70
hind leg 1.32 1.45 — = 0.32

265

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

e)

À 32

30

37

39 2 Dean

45 46

Figs. 30-46. — 30-34. R. kastanoparuphe paratypes; 30-32 male, 30 apex of abdomen, lateral view, 31 paramere, 32 hind fe-
mur; 33-34 female apex of abdomen, 33 dorsal view ‘7’ and ‘8’ abdominal tergites 7 and 8 respectively; 34 lateral view. Scales
30, 32-34 1 mm, 31 0.25 mm. — 35-40. R. pseudocelebensis paratypes; 35-37 male, 35 apex of abdomen, lateral view, 36 pa-
ramere, 37 hind leg; 38-40 female, 38 hind femur, 39 apex of abdomen lateral view, 40 left half of abdomen, dorsal view.
Scales 35, 37-40 1 mm, 36 0.25 mm. — 41-46, parameres of 41-42, R. celebensis (Dumoga Bone N. P.), 43-44, R. pseudoce-
lebensis (Kolaka and Buton respectively), and 45-46, R. sulawesiana (paratype); 45, lateral view, 46, medial view. Scale 0.25

mm.

266

R pseudocelebensis 3

fore leg 0.96 0.99
middle leg 1.48 1.17
hind leg 1.26 1.35
R. pseudocelebensis 9

fore leg 1.00 0.95
middle leg 1.41 1.11
hind leg 1.23 1.35
R. robina 3

fore leg 0.91 0.93
middle leg 1.47 1.11
hind leg 1.37 1.14
R robina 9

fore leg 0.90 0.93
middle leg 1.44 1.08
hind leg 1.29 1.28
R. trichota &

fore leg 0.89 0.91
middle leg 1.41 1.13
hind leg 1.20 1.24
R. trichota 2

fore leg 0.96 0.96
middle leg 1.43 1.1177
hind leg 1.22 1.31
R.tropidata 3

fore leg 0.92 0.96
middle leg 1.52 1.23
hind leg 1.40 1.22
R.tropidata 9

fore leg 0.90 0.93
middle leg 1.42 1.11
hind leg 1.23 1.23

NIESER & CHEN, Rhagovelia of Sulawesi

= - 0.28
0.07 0.53 0.70
= — 0.31

= = 0.23
0.08 0.50 0.69
= — 0.29

En = 0.24
0.08 0.52 0.66
- — 0.30

= = 0.23
0.08 0.51 0.66
- — 0.31

— — 0.20
0.06 0.43 0.65
— — 0.30

= — 0.22
0.08 0.47 0.70
- — 0.32

— - 0.22
0.09 0.48 0.68
— — 0.32

— — 0.22
0.08 0.41 0.64

= — 0.36

Rhagovelia sulawesiana Polhemus & Polhemus

(figs. 61, 69, 81)

Rhagovelia sulawesiana Polhemus & Polhemus, 1988: 198-
199, figs. 158-164, 230.

Material. — SULAWESI SELATAN: E of Maros,
Bantimurung Area, Sg. Pattunuang Asue [type locality],
5°03’S 119°41’E, fast flowing stream, ponded areas,
through disturbed forest on limestone, 23.iv.1991, J. van
Tol, 38 59; SE of Malino, rivulet at N side of
Lompobattang, cultivated area, 20.vi.1985-A, J. van Tol
26 . All apterous.

Distribution. — S Sulawesi (fig. 81).

Rhagovelia wallacei Polhemus & Polhemus
(figs. 71, 76, 82)

Rhagovelia wallacei Polhemus & Polhemus, 1988: 203-205,
figs. 187-193, 230.

Material. — SULAWESI UTARA: Kotamobagu, Caves,
26.x.1985, 116 72; Dumoga Bone N. P., Base Camp,
Toraut R., maze, 7.xi.1985, 78 159; Tumpah R., beach,
19.x.1985, Id 42 (12 macr.); Tumpah R., Waterfall
oberh. Seitenbach, 21.xi.1985, 2d ; Malibagu Z., 8.xi.1985,
12 macr.; Lakes, Kleine See, Abfluss des Flusses (small lake,
drainage of stream), 16.xi.1985, 6d 119 (22 macr.);
Lakes, kleiner See, Fischteich, 16.xi.1985, 26 49;
Südküste, Strasse, Bach, 18.xi.1985, 28 99; all leg. G.
Zimmermann; Dumoga Bone N. P., waterfall creek (type
locality), 24.v.1985, 16; Tumpah river near confluence
Toraut, UTM-XL00631, 210 asl., 21.v.1985, 1d, both J.
van Tol; 20 km E of Kotamobagu, Mt Ambang, alt. 1210
m, 8.xi.1985, multistr. evergreen forest, soil litter sifted, 1d
19, leg. J. Krikken. — SULAWESI TENGAH, 50 km SE of: Lore
Lindu N. P., sur. Dongi Dongi shelter, 1°13’S 120°11’E,
tributary of Sopu river, edge of multistr. evergreen forest,
950 m, 4.xii.1985-C, 19. Apterous, unless otherwise indi-
cated.

Distribution. — Northern and eastern parts of

Sulawesi (fig. 82)

267

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Rhagovelia blogiokommena sp. n.
(figs. 2-6, 82)

Type material. — Holotype d : Sulawesi Tenggara, moun-
tain stream between Tamborasi and Wolo, 1 mar. 1989, leg.
N. Nieser, N8926 (ZMA). — Paratypes, same data as holoty-
pe, 7d 99 (19 allotype ZMA, 36 39 NC, 1d 19 PCHC,
1d 19 Jrrc, 1d 19 MBBJ, 1d 19 RMNH, 1% ZMC). All ap-
terous (fig. 82).

Description. — Apterous form. Dimensions.
Length d 2.80-2.92, 9 2.85-3.00; width d 1.0-1.1,
2 1.1-1.2; width of head d 0.70-0.73, £ 0.72-0.74;
width of pronotum d 0.90-0.95, 2 0.95-1.02.

Colour, generally dark grey to blackish, anterior
quarter of pronotum orange-yellow, connected with
yellowish propleura, and prosternum. Dorsal half
(males) to two thirds (females) of connexivum oran-
ge. Venter dark, juga, prosternum and abdominal
sternite 7 yellowish, genital segments of male yello-
wish. Basal half of first antennal segment, basal part
of rostrum, basal three quarters of fore femur, basal
third of hind femur, acetabula, coxae and trochanters
pale; hind femur in ventral view medium brown.

Minute black denticles virtually restricted to poste-
rior half of jugum of head and propleura on both
sides of rostrum. Mesopleuron in front of acetabulum
with a group of 11-16 coarse punctures. Dorsum
rather bare, abdominal tergites pruinose laterally.
Vertex, interoculus, sides of thorax and abdomen, an-
tennae and legs with the usual pubescence and setae.
Punctures on disk of pronotum indistinct except for a
row at posterior margin of transverse orange-yellow
part.

Length of antennal segments I-IV d 0.70 : 0.37 :
0.47 : 0.41, 2 0.68 : 0.35 : 0.45 : 0.41. Pronotum
long, covering mesonotum, length: width 0.75 : 0.92.
Length of metanotum on midline, 0.10. Abdominal
tergites 1-5 subequal in length (0.15), tergite 6 slight-
ly longer (0.20), tergite 7 0.30 in male, 0.35 in fema-
le.

Male. Posterior trochanter with 1-3 small but dis-
tinct blunt teeth. Posterior femur slightly less than
four times as long as wide (1.20/0.32), on ventral
margin proximally with about 10 minute and closely
set teeth, followed about halfway by a large spine
about as long as half the width of femur, with a row
of about 11 spines of decreasing length distally (fig.
4). Anteroventrally a row of 5 smaller spines located
in distal part. Larger spines black tipped, smaller spi-
nes and teeth entirely black or nearly so. Posterior ti-
bia virtually straight, armed beneath with a row of
about 15 rather coarse pegs, a few proximally paired.
Connexiva more or less horizontal, gradually conver-
ging posteriad. Basal width of abdominal tergite 7
subequal to its median length. Abdominal sternite 7
with a pair of shallow impressions separated by a nar-

268

row and low keel, sternite 8 with an indistinct ventral
keel (fig. 2). Parameres as in fig. 3.

Female. Posterior trochanter without small teeth.
Posterior femur slightly less than four times as long as
wide (1.1/0.3), row of minute teeth absent, row of
larger spines essentially as in male. Posterior tibia with
small and less distinct pegs as in male. Connexiva mo-
re or less vertical, on tergites 5 and 6 somewhat sinu-
ate, otherwise gradually converging caudally, with the
pointed apices of connexiva touching, or nearly so
(fig. 6). One female has the connexiva nearly hori-
zontal. Tergite 8 slanting ventrad at an angle of about
45°, proctiger pointing ventrad. Sternite 7 about half
as long as remaining abdominal sternites (0.5/1.0).
Gonocoxa hidden in segment 7.

Etymology. — Blogiokommenos, greek adjective
meaning pockmarked, referring to the coarse puncta-
tion on mesopleura.

Comparative notes. — Similar to R. tropidata, R.
wallacei and, in males, R. minahasa, all of which have
distinct minute black denticles on pro- and meso-
pleura. À. minahasa lacks mesothoracic punctures, À.
tropidata has fewer (4-9) and indistinct punctures in
front of mesoacetabula, R. wallacei has slightly less
(about 10) and less coarse punctures there. Male ster-
nite 7 and female connexiva are also different (figs. 2,

62, 70-71, 6, 67)

Rhagovelia chrysomalla sp. n.
(figs. 7-11, 83)

Type material. — Holotype d : Sulawesi Tengah, 60 km
SE of Palu: Lore Lindu N. P., UTM-SJ95, Danau Tambing
and brooklets, 1600 m asl., Pandanus, 7 Dec. 1985-A, leg. J.
van Tol (RMNH). — Paratypes 48 89, same data as holotype
(23 39, including allotype RMNH, 26 39 NC, 19 MBB},
12 yrpc). All apterous (fig. 83).

Description. — Apterous form. Dimensions.
Length d 3.80-3.90, 9 4.10-4.33; width d 1.3-1.4,
? 1.4-1.6; width of head d 0.88-0.90, 9 0.90-0.92;
width of pronotum d 1.20-1.28, 9 1.22-1.40.

Colour, dark grey brown to blackish, eyes castane-
ous. Anterior fifth of pronotum orange-yellow.
Propleura, pro- and mesosterna, first antennal seg-
ment except apex, acetabula, coxae and trochanters
anterior femur except apex yellowish. Middle and
hind femur variably yellowish and brown, connexiva
entirely in female, distal two thirds in male orange
yellow. Sternites 7 and 8 in male, tergite 8, proctiger
and gonocoxa in female yellowish. The orange-yellow
anterior transverse band of pronotum tends to be se-
parated from the yellow area of the propleura by a
dark spot behind the eyes which is quite variable and
may be absent.

Thoracic venter with minute black denticles, ex-

Nieser & CHEN, Rhagovelia of Sulawesi

Figs. 47-61. — 47-55. R. robina paratypes; 47 apex of abdomen, lateral view, 48 paramere, 49 hind trochanter and femur; 50-
52 female, 50 hind femur, 51 dorsal part of thorax and abdomen lateral view, 52 left half of thorax and abdomen, dorsal view;
53-55 hemielytra of males (54-55 of same specimen). Scales 47, 49-55 1mm, 48 0.25 mm. — 56-60. R. trichota paratypes; 56-
58 male, 56 apex of abdomen lateral view, 57 paramere, 58 hind trochanter, femur and tibia; 59-60 female, 59 apex of ab-
domen, lateral view, 60 right half of thorax and abdomen, dorsal view. Scales 56, 58-60 1 mm, 57 0.25 mm. - 61, R sula-
wesiana paratype female, right half of thorax and abdomen, dorsal view. Scale 1 mm.

269

TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

tending onto pleura, most distinctly on jugum and
proepisternum. Interoculus, thoracic and abdominal
tergites clothed with dense, appressed golden-yellow
pilosity. (This golden pilosity is also present in larva
V). In addition dorsum of thorax and abdomen with
sparse long erect dark cilia. Vertex sides of thorax an-
tennae and legs with the usual pubescence and setae.

Length of antennal segments I-IV d 0.95 : 0.44 :
0.69 : 0.67, 2 0.99 : 0.47 : 0.68 : 0.59. Pronotum
long, covering mesonotum. Disk of pronotum with
some indistinct punctures, obscured by pilosity.
Length: width of pronotum male 0.93 : 1.22, female
1.18 : 1.32. Length of metanotum on midline, 0.20.
Abdominal tergites 1-5 subequal in length (0.20 in
male, 0.25 in female), tergite 6 subequal in male,
slightly longer (0.30) in female, tergite 7 distinctly
longer (0.43 in male, 0.50 in female).

Male. Posterior trochanter length: width 0.40 :
0.21, without teeth or warts. Posterior femur slender,
slightly less than four and a half times as long as wide
(1.44/0.33), on ventroposterior margin about half-
way with a rather small brown spine, its length so-
mewhat less than one thirds the width of femur, with
a row of about 7 spines of decreasing length distally
(fig. 9). Posterior tibia straight, armed beneath with a
row of about 20 small teeth. Connexiva slanting up-
ward about 45°, convex on segments 3-5 strongly
converging posteriad on segment 7, no caudal points.
Basal width of abdominal tergite 7 1.5 times its me-
dian length (0.6/0.4). Abdominal sternite 7 compara-
tively short, about 1.5 times as long as tergite 6
(0.3/0.2), somewhat flattened medially, without a ca-
rina. Genital segments prominent (fig. 7), fusiform,
tergite 8 slightly longer than tergite 7 (0.45/0.40).
Sternite 8 laterally compressed at base, resulting in a
low blunt carina. Median length of pygophore dis-
tinctly longer than median length of sternite 8
(0.4/0.3). Parameres as in fig. 8.

Female. Posterior femur about five times as long as
wide (1.4/0.23). Posterior femur and tibia without
teeth or spines. Connexiva vertical, sinuate, slightly
curved inward on segments 4-5 and curved outward
on segments 6-7, in some specimens nearly straight
(fig. 11). In lateral view connexiva slightly convex
dorsally on segments 4-6, caudal apex truncate with a
tuft of pilosity at the tip (fig. 10). Basal width of ter-
gite 7 slightly larger than its median length
(0.55/0.50). Tergite 8 nearly horizontal, with caudo-
lateral tufts of bristles. Sternite 7 large, about one
third as long as the remaining abdominal sternites to-
gether (0.6/1.8). Gonocoxa clearly visible, about half
the length of sternite 7, ventro-laterally slightly com-
pressed. Proctiger distinctly visible.

Etymology. — Chrysomallos, greek adjective mean-
ing ‘with a golden fleece’, refers to the golden pubes-

270

cence on dorsum of this species.

Comparative notes. — The size combined with stri-
king golden pubescence dorsally set this species apart
from its Sulawesi congeners.

Rhagovelia daktylophora sp. n.
(figs. 12-16, 79, 81)

Type material. — Holotype d: Sulawesi Tengah, SW
Luwuk, Toptop Camp along Batui River, 1°09'S,
122°31’30”E, 120 m, lowland rainforest, 19-21 Oct 1989,
leg. J. P. Duffels, sample Sul. 18 (ZMA). — Paratypes (inclu-
ding allotype 9) same data as holotype 146 72 (zmA, 2d
29 NC, 1d 19 MBBJ); Batui River at Singsing Camp, SW
of Luwuk, c. 1°09’S 122°31’E, 90m, middle sized stream in
lowland rainforest, 14-17 Oct 1989, sample Sul.14, J.
P.Duffels, 38 (16 RMNH, 26 ZMA); NE Luwuk, Sungai
Bantayan, near road Kayutanyo to Siuna, 0°47’S 123°00’E,
50-100 m, medium sized river in slightly disturbed lowland
rainforest, 7.x.1989, leg. J. P. Duffels, sample Sul. 2, 106
30/2 (ZA 37/2 NI reg SPC aloe) =
Luwuk area, Sg. Batui and tributaries at Singsing Camp,
17.x.1989, J. van Tol 236 119 (48 22 macr., RMNH, 1d
apt., 2ó 12 maer. Ne, 26 22 apt. jrrG Lon lie rapt
SNOW); Luwuk area, Sg. Biak, 21.x.1989, leg. J. van Tol, 8d
92 (12 macr.). Apterous unless otherwise indicated (fig.
81).

Description. — Apterous form. Dimensions.
Length d 3.04-3.25, 2 3.05-3.28; width d 1.2-1.3,
2 1.3-1.4; width of head d 0.80-0.84, © 0.80-0.83;
width of pronotum d 1.02-1.13, 9 1.10-1.20.

Colour, black, pronotum with anterior transverse
orange band usually slightly narrower than interocu-
lar space. Basal third of first antennal segment, aceta-
bula, fore and hind coxae and trochanters and basal
part of anterior femur ventrally pale yellowish. In
some specimens middle coxa medium to light brown,
but in most blackish. Venter largely pruinose ash
grey.

Venter with minute black denticles, which are dif-
ficult to see due to pruinose dark grey background,
extending onto episterna. Dorsum with a few short
golden-yellow setae, longer and denser on genital seg-
ments of male. Vertex sides of thorax antennae and
legs with the usual pubescence and setae.

Length of antennal segments I-IV d 0.80 : 0.40:
0.44 : 0.43, 2 0.80 : 0.39 : 0.43 : 0.42. Pronotum
long, covering mesonotum, usually laterocaudal ang-
les of mesonotum narrowly visible. Disk of pronotum
with comparatively fine punctures, about
30/0.25mm. Length: width of pronotum 0.85 :
1.10. Length of metanotum on midline, 0.15.
Abdominal tergites 1-5 subequal in length (0.12 in
male, 0.15 in female), tergite 6 slightly longer (0.20
in male, 0.30 in female), tergite 7 distinctly longer
(0.40).

Male. Posterior trochanter length: width 0.32:

0.18, pilose but without teeth or warts. Posterior fe-
mur slender, nearly five times as long as wide
(1.4/0.3), on ventroposterior margin about halfway
with a large black spine its length two thirds the
width of femur, with a row of 6 black spines of de-
creasing length distally and three very small spines
proximally of large spine (fig. 14). Posterior tibia
straight, armed beneath with a row of about 16 small
teeth, proximally three or four extra teeth forming
pairs. Connexiva more or less horizontal gradually
converging posteriad, no caudal points. Basal width
of abdominal tergite 7 equal to its median length.
Abdominal venter with long, quite dense, golden yel-
low setae on sternites 3-6 and the base of sternite 7.
Sternite 7 in lateral view distinctly compressed dorso-
ventrally in posterior three fourths. Apical part of
sternite 7 without pubescence, shining black, with a
pair of sublateral impressions, separated by a low, nar-
row median carina (fig. 12). Segment 8 prominent,
elongate, in dorsal view, length 0.55, basal width
0.37, sternite 8 shining black. Parameres relatively
large, as in fig. 13.

Female. Generally with a more compact build as
compared to male, due to compressed genital seg-
ments. Posterior femur nearly five times as long as wi-
de (1.18/0.27). Spines on posterior femur and tibia
essentially as in male. Connexiva vertical, strongly
curved inward on segment 5-6; ending in narrow cau-
dal points with a tuft of pilosity at the tip (fig. 16).
Another tuft of bristles just ventral of apices of con-
nexiva and a smaller group of somewhat more scatte-
red bristles on dorsal margin of connexiva at segment
6, best visible in lateral view (Fig. 15). Tergite 7 with
a well developed mediocaudal projection on hind
margin, with a short tuft of setae apically. Tergite 8
nearly vertical, genital segments retracted into seg-
ment 7. Sternite 7 large, shining black under quite
dense ash white pubescence, slightly over half as long
as the remaining abdominal sternites together
(0.67/1.2), sides of segment 7 pruinose grey.

Macropterous form. — Mostly as apterous except
for development of pronotum and hemielytra.
Length d 3.5-3.7, 2 3.7-3.8; humeral width of pro-
notum of both sexes 1.37-1.45. Submedial carinae at
base of abdomen just not reaching the hind margin of
first abdominal tergite. Female without the finger-li-
ke projection on caudal margin of tergite 7 and the
constriction of connexiva on segments 5-6 less pro-
nounced.

Etymology. — Daktylophora, Greek adjective mean-
ing ‘bearing a finger’ refers to the finger-like projec-
tion on tergite 7 in apterous female.

Comparative notes. — Very similar to À. celebensis
and À. pseudocelebensis, but the connexiva of apterous
females curved inward on segments 5-6 in combina-

NieseR & CHEN, Rhagovelia of Sulawesi

tion with a well developed finger-like projection on
the caudal margin of abdominal tergite 7 (figs. 15-16,
39-40) and the parameres of males, with the broad
apical part longer (figs. 13, 36, 41-44), are distinctive.

Rhagovelia horaia sp. n.
(figs. 17-20, 84)

Type material. — Holotype d: Sulawesi Tenggara,
Centipede Camp, 3.xi.1989, J. van Tol (RMNH). — Paratypes
23 69 (including allotype), same data as holotype (RMNH,
16 22 NC, 12 MBBJ); S. Mokowu, 5.xi.1989, J. van Tol,
19, all apterous (fig. 84).

Description. — Apterous form. Dimensions.
Length d 3.4-3.7, 9 3.5-3.8, width of head d 0.78-
0.81, © 0.78-0.83; width of pronotum ¢ 1.07-1.12,
® 1.08-1.12; thoracic width d 1.12-1.23, 2 1.20-
125:

Colour, reddish brown, eyes dark castaneous to
blackish, anterior quarter of pronotum and entire
propleura pale yellow. Basal two thirds of first anten-
nal segment, basal two thirds of fore femur, coxae and
trochanters, basal half of hind femur especially in fe-
male, distal margin of connexiva more distinctly dor-
sally than ventrally, pale.

Venter with minute black denticles extending onto
thoracic pleura, distinctly on propleura only a few on
meso- and metapleura. Dorsum with normal pubes-
cence and bristles, rather dense yellowish setae on ge-
nital segments of male. Vertex, head, sides of thorax
and apex of abdomen, antennae and legs with the
usual pubescence and setae.

Length of antennal segments I-IV d 0.81 : 0.40:
0.49 : 0.46, 2 0.80: 0.40: 0.47: 0.43. Pronotum
long, covering mesonotum, length: width of prono-
tum 0.85 : 1.1, length of metanotum on midline d
2 0.12. Posterior three quarters of disk with coarse
punctures, about 50/0.25 mm’. Abdominal tergites
1-6 in male and 1-5 in female, subequal in length
(0.20-0.22), tergite 7 in male longer (0.42), tergite 6
in female slightly longer (0.30), tergite 7 distinctly so
(0.55).

Male. Posterior trochanter length : width 0.40 :
0.25, with 7-8 well developed teeth. Posterior femur
strongly inflated, about two and a half times as long as
wide (1.73/0.78 to 1.57/0.60), on posteroventral
margin proximally a row of about 15 minute teeth,
about halfway a large spine about a quarter as long as
the width of femur, with a row of 7 spines of decre-
asing length distally. Anterior to the distal row a row
of 4-5 smaller teeth. Posterior tibia curved (fig. 19),
armed on inner side with a proximal row of about 13
teeth, then a larger tooth, followed by four distal
teeth. In addition a more posterior row of about 12
smaller teeth. Connexiva horizontal slightly convex

271

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 62-80. — 62-67, R. tropidata paratypes; 62-64 male, 62 apex of abdomen, lateral view, 63 paramere, 64 hind leg; 65-67
female, 65 hind leg; 66 abdomen in lateral view, 67 left half of thorax and abdomen dorsal view. Scales 62, 64-67 1 mm, 63
0.25 mm. — 68-73, Apex of abdomen, lateral view. 68, R. pruinosa male, 69 R. sulawesiana male, 70 R. minahasa male, 71 R.
wallacei male, 72 R. grayi female, 73 R. minahasa female. Scale 1 mm. — 74-76. Paramere (75-76 redrawn after Polhemus &
Polhemus 1988); 74 R. minahasa, 75 R. hamjadi, 76 R. wallacei. Scale 0.25 mm. — 77-78, hind femur of female;77 R. prui-
nosa, 78 R. hamjadi (redrawn after Polhemus & Polhemus 1988); 79-80. Right hemielytron; 79 R. daktylophora, 80 R. pseu-

docelebensis. Scale 1 mm.

272

on segment 4-6, otherwise gradually converging to-
ward tergite 7, without apical points. Basal width of
abdominal tergite 7 subequal to its median length
(0.4), centrally more or less bare and somewhat shi-
ning. Abdominal sternite 2 with a well defined me-
dian keel, sternite 7 flattened medially, nearly bare,
shining, without median keel, its median length one
third of median length of remaining sternites together
(fig. 17). Sternites 8 and 9 of subequal length, some-
what shorter (0.25/0.35) than sternite 7. Paramere as
in fig. 18.

Female. Posterior trochanter with three or four
small teeth. Posterior femur four and a half times as
long as wide (1.35/0.31). Posterior femur on ventral
margin proximally with 0-3 small minute teeth, large
spine halfway three fifth the width of femur, followed
by 4-6 smaller spines, hind tibia straight, with a, dis-
tally double, row of about 18 teeth on inner surface.
Connexiva slanting 45° upwards, slightly convex on
segments 3-5, converging caudally with short, nar-
rowly triangular, caudal points. Abdominal tergite 7
distinctly longer than its basal width (0.55/0.30).
Tergite 8 slanting 45° downward, proctiger visible in
dorsal view. Abdominal sternite 7 half as long as pre-
ceding ones together, bare, shining, no minute black
denticles, gonocoxa short but well visible (fig. 20).
Macropterous form, unknown.

Etymology. — Horaios, a greek adverb meaning e. g.
youthfully beautiful.

Comparative notes. — The male is nearly identical
to À. grayi Polhemus & Polhemus, except for the
proctiger which lacks a pair of proximal tubercles
(figs. 21, 22). Females of R. grayi have connexiva up-
turned, expanded and incurvate along tergites 4 and 5
and apically folded over abdomen, whereas in R. ho-
raia the connexiva of female are simple. Another si-
milar species is R. lorelinduana which has in male a
quite different proctiger (figs. 21, 23) and in female
curved and folded connexiva much as in R. grayi.

Rhagovelia kalami sp. n.
(figs. 24-29, 82)

Type material. - Holotype d : Pulau Buton, first stream
along road from Baubau to the North, 8 Mar. 1989,
N8935, leg. N. Nieser (RMNH). — Paratypes, same data as
holotype, 66 172 (29 including allotype RMNH, ZMA 16
22, zme lé 29, rest NC); Buton, small stream near sea,
9.iii.1989, N8941, 146 182 (46 42 PCHC, 16 1 BPUH,
18 19 MBB), 28 32 Jrrc, 1d 12 Zc); Buton, small
stream 16 km E Baubau, 10.iii.1989, N8944, 66 99. All
apterous (fig. 82).

Description. — Apterous form. Dimensions.
Length d 3.08-3.28, 2 3.20-3.50; width d 1.15-
1.19, £ 1.30-1.42; width of head d 0.72-0.80, 9

NIESER & CHEN, Rhagovelia of Sulawesi

0.78-0.80; width of pronotum d 0.98-1.08, 2 1.10-
1.18.

Colour, dark greyish. Eyes dark castaneous to
black, interoculus black, pronotum and dorsum of
abdomen largely black with a greyish pruinose
bloom. Anterior quarter of pronotum, propleura and
prosternum yellowish orange. Connexiva light orange
brown, proximally blackish. First antennal segment,
acetabula, coxae trochanters, anterior femur except
for distal apex, base of posterior surface and most of
anterior surface of hind femur, variable part of anteri-
or surface of middle femur, mesosternum and medio-
caudal part of abdominal venter including genital seg-
ments pale yellow orange. Venter with minute black
denticles which extend onto jugum, proepisternum
and dorsal sides of acetabula. Dorsum of abdomen of
male with golden-yellow setae. Vertex sides of thorax
antennae and legs with the usual pubescence and
setae, except for hind tibia of male which is more hai-
ry than in most other species.

Length of antennal segments I-IV d 0.78 : 0.39 :
0.47 : 0.44, ® 0.77 : 0.38 : 0.47 : 0.42. Pronotum
long, covering mesonotum, in some specimens latero-
caudal angles of mesonotum narrowly visible. Disk of
pronotum with coarse punctures, about 40/0.25mm’.
Length : width of pronotum 0.80 : 1.05. Length of
metanotum on midline, 0.12. Abdominal tergites 1-6
in male and 1-5 in female subequal in length (0.2)

Male. Posterior trochanter length: width 0.3: 0.2,
with 5-6 small teeth of which 2 twice as large as the
others. Posterior femur strongly inflated, less than
three times as long as wide (1.4/0.5), on ventral mar-
gin distally with about 20 minute teeth, about half-
way a large spine its length about one fourth the
width of femur, with a double row of 6-7 spines of
decreasing length distally, the larger spines pale, with
blackish tip. Posterior tibia slightly curved armed
beneath with a row of about 17 short blunt teeth of
which the apical one and the fourth or fifth from apex
are larger (fig. 26), anterior to row a few extra teeth.
Connexiva more or less horizontal, distal margin
straight to slightly convex on segments 4-6, conver-
ging posteriad, no caudal points. Basal width of ab-
dominal tergite 7 three quarters its median length
(0.32/0.40). Sternite 7 in lateral view laterally com-
pressed in posterior three fourths, with a low median
carina marked at its base by a small but dense tuft of
setae resembling a spine (fig. 24). Genital segments
hirsute, tergite 8 as long as wide (0.32/0.33).
Parameres as in fig. 25.

Female. Posterior femur four times as long as wide
(1.3/0.32). Spines on posterior femur essentially as in
male. Posterior tibia straight, armed beneath with a
row of about 17 short blunt teeth all of subequal size.
Connexiva slanting about 45° upwards, distal margin
slightly sinuate, thickened on segment 4, where there

275

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

is a patch of short dense pubescence. Apex of con-
nexiva triangularly produced caudally, accentuated by
a tuft of bristles (fig. 29). Tergite 7 slightly longer
than its basal width (0.40/0.35). Tergite 8 nearly ho-
rizontal, with laterocaudal tufts of pilosity, basal
width equal to its median length (0.32), apical mar-
gin truncate. Sternite 7 large, shining, half as long as
the remaining abdominal sternites together (0.6/1.2).
Gonocoxa not distinctly visible.
Macropterous form, unknown.

Etymology. — Kalami, greek noun, meaning shin,
referring to the modified tibia of male.

Comparative notes. — À. kalami is not very similar
to other species in the region, those with a similar
male tibia (R. grayi, R. horaia) are more slender and
longer. R. tropidata and R. wallacei are of similar sha-
pe, but have the male hind leg less modified. See also
key to species.

Rhagovelia kastanoparuphe sp. n.
(figs. 30-34, 84)

Type material. — Holotype & (ZMA): Sulawesi Tenggara,
NE Kolaka, Mokowu Camp, along Mokowu River, nr.
Gng. Watuwila, c. 3°49°S 121°40’E, 200m, 29-31 Oct.
1989, disturbed lowland rainforest, J. P. Duffels, sample
Sulawesi 24 (ZMA). — Paratypes: same data as holotype, 14¢
269 (78 189 including allotype 9 ZMA, 16 19 BPUH, 19
JTPC, 16 19 MBBJ, 36 32 NC, 16 19 PCHC, 16 19
RMNH); Sungai Mokowu, 30.x.1989, J. van Tol, 29; Rd.
Kendari-Lalimboe, 21.ii.1989, N8903, 22; tributary Sg.
Mowewe, 28.11.1989, N8923, 18 49. — P. Buton, stream
N of Baubau, 8.iii.1989, N8935, 106 199 (68 109 NC,
16 19 BPUH, 16 39 yrec, 16 29 PCHC, 22 RMNH, 29
ZMA, 16 19 ZMC); P. Buton, Sg. Labuhan Tobelo at Jismil
camp, 18.xi.1989, leg. J. van Tol 968 329 (19 macr., 5d
249, incl. 1 macr., RMNH, 48 89 NC). Apterous, unless
otherwise indicated (fig. 84).

Description. — Apterous form. Dimensions.
Length d 3.0-3.3, 2 3.0-3.2; width d 1.0-1.1, ©
1.1-1.2; width of head 6 0.79-0.82, 2 0.82-0.83;
width of pronotum d 0.92-1.01, 9 1.0-1.08.

Colour, black with a greyish tinge. Orange trans-
verse band anteriorly on pronotum only slightly wi-
der than interoculus. Propleurae orange, separated
from pronotal transverse band by a narrow black
mark behind eyes. Distal margin of connexiva orange
brown, more distinctly so in females than in males,
without obvious pilosity. Basal half of first antennal
segment, acetabula, coxae, trochanters, basal half of
fore femur, basal quarter of hind femur pale yellow.
Rostrum castaneous except for darker apex. Abdo-
minal sternites 7 and 8 in male and sternite 7 in fe-
male, castaneous.

Venter with minute black denticles, which do not
extend onto pleura except for a few on metapleuron.

274

Dorsum mostly with a greyish pruinose cover, sparse
pilosity except for some shiny yellowish setae on geni-
tal segments of male and a tuft of long dark setae at
caudal apex of connexiva. Vertex anteriorly with short
dark setae, sparse long dark setae on head, sides of
thorax and apex of abdomen. Antennae and legs with
the usual pubescence and setae.

Length of antennal segments I-IV d 0.81 : 0.40:
0.49 : 0.46, 2 0.80 : 0.40 : 0.47 : 0.43. Pronotum
long, covering mesonotum. Length: width of prono-
tum 0.85: 1.4. Disk of pronotum with about 60
punctures/mm’. Length of metanotum on midline,
0.12. Abdominal tergites 1-5 subequal in length
(0.14), tergite 6 slightly longer (0.25), tergite 7 dis-
tinctly longer (0.45).

Male. Posterior trochanter length: width 0.30:
0.18, without teeth or warts. Posterior femur rather
slender, four and a half times as long as wide
(1.3/0.28), on ventral margin about halfway with a
large spine about 0.6 as long as the width of femur,
with a row of 5-7 spines of decreasing length distally
(fig. 32). Posterior tibia straight, armed beneath with
small blunt teeth proximally shorter and more dense-
ly placed. Connexiva slanting up- and outward, about
45°, gradually tapering downward on tergite 7, with-
out apical points. Basal width of abdominal tergite 7
0.30, its median length 0.45; sternites 6-8 with a well
defined median keel. Segment 8 prominent (fig. 30),
elongate, in dorsal view, length 0.4 width 0.30.
Parameres as in fig. 31.

Female. Generally with a more compact build as
compared to male, due to compressed genital seg-
ments. Posterior femur five times as long as wide
(1.3/0.27). Length of large spine on posterior femur
three quarters the width of femur, followed by 3-5
smaller spines, hind tibia essentially as in male.
Connexiva variable more or less horizontal to vertical,
dorsal margin straight to very slightly convex, apex of
connexiva narrowly triangularly produced dorsocau-
dally, accentuated by a tuft of setae (fig. 34). Posterior
margin of tergite 7 bluntly triangular with a short me-
diocaudal tuft of setae, often suggesting a spine (fig.
33). Tergite 8 nearly vertical, gonocoxa mostly hid-
den within segment 7. Sternite 7 large, shining, half
as long as the remaining abdominal sternites together.

Macropterous form. The single macropterous fe-
male, which has its wings torn off, is identical to the
apterous form except for stronger development of
pronotum, humeral width 1.40, and for stronger de-
velopment of pronotum, humeral width 1.40, and
absence of the mediocaudal tuft of setae on abdomi-
nal tergite 7.

Etymology. — Kastanoparuphe, greek noun mean-
ing brown band or brown rim, referring to the upper
margin of connexiva.

Comparative notes. — At first sight similar to À. ce-
lebensis and related forms, the lack of long pilosity on
abdominal venter in male and the brownish upper
rim of connexiva in female separate this species easily.
The paramere (fig. 31) and the apex of abdomen of
female (fig. 33, 34) are also characteristic.

Rhagovelia pseudocelebensis sp. n.
(figs. 35-40, 80-81)

Type material. - Holotype d, SULAWESI TENGGARA:
Tamborasi-Wolo, mountain stream, 1 Mar. 1989, N8926,
leg. N. Nieser (RMNH). Paratypes: same data as holotype,
138 49 (incl. allotype) (19 macropterous) (allotype
RMNH); Sungai Kolaka, upstream of Kolaka, 27 Feb. 1989,
N8921A, 168 189 apt, 166 209 macr.(39 apt. PCHC);
same, N8921B, 26 (macr.) 39 (29 apt, 19 macr. JTPC);
Tamborasi, mouth of subterranean stream, 1.111.1989,
N8925 816 219 (46 PCHC, 46 JTPC); 20 km E of Kolaka,
mountain stream, 3 Mar. 1989, N8934 29 (all leg. N.
Nieser). — PULAU BUTON, Ist stream road N of Bau-Bau,
8.i11.1989, N8935 4d 49, leg. N. Nieser. All apterous un-
less otherwise indicated (fig. 81). The following museums
received 1d 1@ paratypes each: BPUH, CHC, JTPC, MBBJ,
ZMA; ZMC received a single 9, remaining paratypes in NC.

Description. — Apterous form. Dimensions.
Length d 2.98-3.80, 9 2.78-3.10; width d 1.1-1.2,
2 1.3-1.4; width of head d 0.76-0.82, 2 0.80-0.83;
width of pronotum d 1.00-1.03, 9 1.02-1.13.

Colour, black, pronotum with anterior transverse
orange band only slightly broader than interocular
space, laterally obscured by pale grey pruinose pilosi-
ty. Basal third of first antennal segment, acetabula, fo-
re and hind coxae and trochanters and basal third of
anterior femur pale yellowish. Ventral pilosity of ab-
domen in male yellowish.

Venter with minute black denticles, which are dif-
ficult to see due to pruinose dark grey background,
extending onto episterna and head. Dorsum with a
few short golden-yellow setae, longer and denser on
genital segments of male. Vertex anteriorly with short
dark setae, sparse long dark setae on head, sides of
thorax and apex of abdomen. Antennae and legs with
the usual pubescence and setae.

Length of antennal segments I-IV d 0.83 : 0.42 :
0.46 : 0.48, 2 0.81 : 0.38 : 0.42 : 0.45. Pronotum
long, covering mesonotum, in some specimens is the
posterior margin of mesonotum narrowly visible.
Length: width of pronotum 0.75 : 1.0. Length of me-
tanotum on midline, 0.12. Abdominal tergites 1-5
subequal in length (0.12), tergite 6 slightly longer
(0.17), tergite 7 distinctly longer (0.55).

Male. Posterior trochanter length: width 0.30 :
0.17, without teeth or warts. Posterior femur rather
slender, four times as long as wide (1.2/0.3), on ven-
troposterior margin about halfway with a large spine
about 0.6 as long as the width cf femur, with a row of

NIESER & CHEN, Rhagovelia of Sulawesi

5-6 spines of decreasing length distally (fig. 37).
Posterior tibia straight, armed beneath with five small
blunt teeth proximally and three slightly larger shar-
per teeth apically. Connexiva more or less horizontal
or slightly slanting upward, gradually converging pos-
teriad. Basal width of abdominal tergite 7 0.38, its
median length 0.40. Abdominal venter with long,
quite dense, golden yellow pilosity. Sternite 7 in la-
teral view distinctly compressed dorsoventrally in
posterior three fourths (fig. 35). Apical part of sterni-
te 7 without pubescence, shining black, flattened
with a low, narrow median carina. Segment 8 promi-
nent (fig. 35), elongate, in dorsal view, length 0.50
width 0.30. Parameres as in fig. 36.

Female. Generally with a more compact build as
compared to male, due to compressed genital seg-
ments. Posterior femur five times as long as wide
(1.3/0.25). Spines on posterior femur and tibia essen-
tially as in male. Connexiva more or less vertical, apex
of connexiva triangularly produced caudally, with a
long tuft of medially curved pilosity (figs. 39, 40).
Tergite 7 with a short mediocaudal projection on
hind margin, with a short tuft of setae apically, this
character is, however, quite variable. Especially in
some females from N8921A the projection is hardly
recognizable. Tergite 8 distinctly (about 60°) slanting
ventrad, proctiger pointing ventrad and somewhat re-
tracted into segment 7 in most specimens. Sternite 7
large, shining, half as long as the remaining abdomi-
nal sternites together (1.0/1.8). Gonocoxa not dis-
tinctly visible.

Macropterous form, as apterous except for full de-
velopment of pronotum and wings. Dimensions,
length d 3.5-3.8, 2 3.5-3.7, width of head 0.80-
0.82, humeral width of pronotum d 1.40-1.43, ©
1.37-1.42. Pronotum broadly rounded posteriorly.
Hemielytra dark brown with darker veins, three clo-
sed cells, two basal, reaching about one third the
length of hemielytron, the distal costal cell reaching
well into distal half (fig. 80). Submedial carinae on
base of abdomen reaching to hind margin of tergite 2.

Etymology. — Pseudocelebensis, greek adjective mean-
ing looking like celebensis, refers to the similarity be-
tween R. celebensis and R. pseudocelebensis.

Comparative notes. — Very similar to R. celebensis
but the female has a small caudal projection on tergi-
te 7, the male paramere is different (figs. 36, 41-44).
In R. sulawesiana which is also very similar, the fema-
le has the connexiva folded over the tergites apically,
the paramere of male is different (fig. 45-46). See also
comparative notes under R. daktylophora.

Rhagovelia robina sp. n.

(figs. 47-55, 82)
Type material. — Holotype d: Sulawesi Tenggara: Road

275

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Rhagovelia A Rhagovelia
| 7
R. celebensis ù @ R.minahasa

R. pseudocelebensig @ A. lorelinduana
R. daktylophora
R. sulawesiana
% R horaia

Rhagovelia i Rhagovelia
@ Rpruinosa i R.wallacei
B A hamjadi i R. trichota

% RB chrysomalia È R. tropidata

R. blogiokommena
R. robina

R. kalami

Figs. 81-84. Distribution of Rhagovelia species in Sulawesi. — 81, R. celebensis, R. daktylophora, R. pseudocelebensis, R. sulawe-
siana; 82, R. wallacei, R. tropidata, R. blogiokommena, R. trichota, R. robina, R. kalami; 83, R. pruinosa, R. hamjadi, R. chryso-
malla, 84 R. minahasa, R. unica, R. kastanoparuphe, R. lorelinduana, R. horaia.

276

Kolaka Kendari, 20 km E Kolaka, stream in hilly forest, 3
mar. 1989, N8934, leg. N. Nieser (RMNH). — Paratypes 66
49 apterous, 26 macropterous, same data as holotype (1?
allotype RMNH, 58 29 NC, 1d 19 MBBJ, 1d PCHC, 1d
JIPC, 16 19 ZMA) (fig. 82).

Dimensions. Length d 3.38-3.50, 9 3.58-3.67;
width of head d 0.80-0.81, © 0.82-0.87; width d
1.18-1.20, 2 1.27-1.32, width of pronotum d 1.07-
1.10, £ 1.10-1.12.

Colour, yellow to orange reddish. Eyes dark casta-
neous. Interoculus, posterior three quarters of prono-
tum, abdominal tergites 2-6, proximal and lateral
margins of tergite 7 in male, mesonotum and abdo-
minal tergites except for central spot on tergite 6 in
female, distal half of antennal segment 1 and remai-
ning antennal segments, apical segment of rostrum,
apex of fore femur, most of fore tibia and tarsus, mid-
dle leg except for coxa and trochanter, dorsoapical
streak on hind femur, hind tibia and tarsus dark
brown to blackish.

Body ventrally and dorsally densely covered with
small black denticles. Dorsum without longer pilosity
except on genital segments of male and posterior mar-
gin of abdominal tergite 8 and connexiva in female.
Head, sides of thorax, antennae and legs with the usu-
al setae. Disk of pronotum without coarse punctures.
Length of antennal segments I-IV d 0.70 : 0.40 :
0.49 : 0.41, 2 0.72 : 0.39 : 0.50 : 0.42. Pronotum
long, covering mesonotum, length: width 1.1 : 0.85.
Length of metanotum on midline, 0.12. Abdominal
tergites 1-5 subequal in length (0.15-0.20), tergite 6
slightly (0.22) longer in male more distinctly so(0.30)
in female, tergite 7 longer (0.40-0.45).

Male. Posterior trochanter one and a half times as
long as wide, with one small teeth and a about four
minute teeth or warts. Posterior femur three and a
half times as long as wide (1.37/0.40), on ventral
margin about halfway a large spine with a length just
over half the width of femur, with a row of 8-10 spi-
nes of rapidly decreasing length distally and a row of
16 very small warts proximally (fig. 49). Anteriorly in
distal half a row of 4-8 small warts. Larger spines
black tipped, smaller spines and teeth entirely black
or nearly so. Posterior tibia straight, armed beneath
with a double row of about 20 pairs of warts of which
the posterior ones are largest. Connexiva more or less
horizontal, gradually converging to caudal margin of
tergite 7, connexival points absent. Basal width of ter-
gite 7 0.5, diverging caudally, caudal width 0.6.
Segment 8 prominent in dorsal view, length and
width about 0.6, sides slightly convex. Abdominal
sternite 2 bluntly keeled in midline, length of sterni-
tes 2-5 subequal (about 0.20), length of sternite 6
0.28, and of sternite 7 0.45. Abdominal sternite 7 la-
terally compressed resulting in a broad median keel

(fig. 47). This keel accentuated by a fringe of long pi-

NieseR & CHEN, Rhagovelia of Sulawesi

losity which runs over the midline of all abdominal
sternites. Sternite 8 lateroproximally compressed.
Parameres fig. 48.

Female. Posterior trochanter without small teeth.
Posterior femur four and a half times as long as wide
(1.28/0.28), proximal row of 0-2 minute teeth, row
of larger spines essentially as in male, posteroventral
row of 0-2 smaller spines. Posterior tibia as in male.
Metanotum with a pair of caudolateral broad blunt
tooth-like swellings. Abdominal tergites 1 and 2 swol-
len, in lateral view distinctly rising above the dorsal
margin of connexiva. Tergites 4 and 5 with a broad
blunt median keel. Connexiva (fig. 52) constricted on
segments 4 and 5, caudally ending in blunt triangular
apices fringed with black setae. Tergite 8 only hori-
zontal in basal part, slightly curved ventrad apically,
proctiger well visible. Gonocoxa 1 well visible (fig.
51).

Macropterous form as apterous except for develop-
ment of pronotum, hemielytra and dorsal abdominal
carinae. Pronotum with coarse ill defined pits in pos-
terior part, about 20/0.25 mm. Humeral width of
pronotum 1.35, median length 1.40. Hemielytra
with 4 cells and one or two variable adventitious cells
(figs. 53-55). Dorsal abdominal carinae long, reach-

ing to caudal margin of tergite 4.

Etymology. — Robinos, a greek adjective meaning
rose-coloured, refers to the reddish colour of the spe-
cles.

Comparative notes. — Similar to À. tropidata and À.
wallacei but differing by the orange-like to pale red-
dish metanotum and first abdominal tergite in À. ro-
bina, which are dark in the other species.

Rhagovelia trichota sp. n.
(figs. 56-60, 82)

Type material. — Holotype d, Sulawesi Tenggara,
Tamborasi, creek of subterraneous stream, 1 Mar. 1989,
N8925 (ZMA). — Paratypes: same data as holotype, 50d
329 (16 29 incl. allotype ZMA; 16 12 PCHC, 1d 19
BPUH; 16 19 MBBJ, 1d 19 JTPC; 264 22 RMNH, remaining
specimens NC); Sg. Kolaka, 27.11.1989, 26 ; Sg. Konaweha
at Sanggona, 7.xi.1989, J. van Tol, 38 32 (RMNH, 1d 19
NC). All apterous (fig. 82).

Description. — Apterous form. Dimensions.
Length d 3.10-3.17, 2 3.30-3.40; width 3 1.07-
1.14, 2 1.18-1.36; width of head d 0.78-0.80, 2
0.80-0.83; width of pronotum d 0.99-1.02, 2 1.10-
1.18.

Colour, dull black, orange yellow transverse band
on anterior margin of pronotum laterally reaching
halfway the eyes. Basal half of first antennal segment,
coxae, trochanters and basal half of anterior femur
yellowish. Distal margin of connexiva, brownish

ZII],

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

orange distinct in female, often indistinct in males.
Abdominal venter with variable area along median
line and on posterior sternites medium brown.

Minute black denticles difficult to distinguish, on
the propleura they are, however, usually quite dis-
tinct. Dorsal side of male, except on head, with loose-
ly set long erect setae. Dorsal side of female bare ex-
cept on connexiva, abdominal dorsum especially on
tergites 1-4 with a grey pruinose bloom. Thoracic
pleura with grey pruinose bloom best developed in fe-
males. Vertex, sides of thorax, antennae and legs with
the usual pubescence and setae.

Length of antennal segments I-IV & 0.80 : 0.41 :
0.48 : 0.49, 2 0.80 : 0.43 : 0.49 : 0.49. Pronotum
long, covering mesonotum. Central part of pronotum
smooth, lateral and hind margins with comparatively
fine punctures, along hind margin of orange yellow
transverse band a row of coarser punctures. Length:
width of pronotum 0.8 : 1.0. Length of metanotum
on midline, 0.12. Abdominal tergites 1-5 subequal in
length (0.20-0.22), tergite 6 subequal (0.22) in male,
slightly longer (0.30) in female, tergite 7 distinctly
longer (0.40 in male, 0.50 in female).

Male. Posterior trochanter length : width 0.30 :
0.15, with about 5 small teeth. Posterior femur rather
slender, slightly less than four times as long as wide
(1.20/0.33), on ventroposterior margin about halfway
with a large, yellowish to blackish, usually medium
brown, spine, its length (0.17) half the width of fe-
mur, with a row of 12-13 very small black teeth proxi-
mally and a row of 5-7 black spines of decreasing
length distally (fig. 58). Posterior tibia straight, armed
beneath with an irregular row of about 20 small teeth.
Connexiva more or less horizontal gradually conver-
ging posteriad, no caudal points. Basal width of abdo-
minal tergite 7 about three quarters its median length
(0.28/0.40). Abdominal sternites 2-5 with a low
broad ridge, accentuated by a band of long pilosity
(fig. 56). Sternite 7 slightly less than half as long as the
remaining ones (0.4/1.0), without median keel.
Genital segments relatively small, tergite 8 about three
quarters the length of 7 (0.3/0.4). Sternite 8 flattened
with a low broad carina. Parameres as in fig. 57.

Female. Posterior femur nearly five times as long as
wide (1.20/0.25). One or two small teeth proximally
of large spine, which is about half as long as width of
femur, on hind femur, distal decreasing row of about
six spines. Teeth on posterior tibia essentially as in
male. Connexiva more or less vertical, distal (‘dorsal’)
rim pubescent except on segments 3 and 4 where it is
bare and strongly shiny. Connexiva curved inward on
segments 3 and 4, folded and dorsocaudally touching;
ending in narrowly triangular points (fig. 60). Tergite
7 mostly hidden from view. Tergite 8 and proctiger
nearly vertical, genital segments retracted into seg-

ment 7 (fig. 59). Sternite 7 large, shining, half as long

278

as the remaining abdominal sternites together
(0.6/1.2).

Macropterous form, unknown.

Etymology. — Trichotos, greek adjective meaning
hairy, refers to the hairy aspect of dorsal surface of
male.

Comparative notes. — The long and relatively den-
se dorsal pilosity distinguishes males from its Sulawesi
congeners. Females are similar to those of R. sulawe-
stana but a caudal finger like process is lacking and
the connexiva are different (figs. 60, 61).

Rhagovelia tropidata sp. n.
(figs. 62-67, 82)

Type material. — Holotype d : Sulawesi Tengah, between
Desa Seseba and Singsing Camp, SW of Luwuk, c. 1°09’S
122°31’E, 80m, narrow stream in lowland rainforest,
14.Oct.1989, sample Sul.12, J. P.Duffels (ZMA). —
Paratypes: same data as holotype 7d 69 (4d 49, including
allotype 2 ZMA, 2d 29 NC, 16 PCHC); Batui River at
Singsing Camp, SW of Luwuk, c. 1°09S 122°31’E, 90m,
middle sized stream in lowland rainforest, 14-17 Oct. 1989,
sample Sul. 14, J. P. Duffels, 38 22 (ZMA, 16 NC); SW
Luwuk, Toptop Camp along Batui River, 1°09’S,
122°31’30”E, 120 m, lowland rainforest, 19-21 Oct 1989,
leg. J. P. Duffels, sample Sul. 18, 16; Luwuk area, S. Biak,
21 Oct. 1989, J. van Tol, 38 22 (16 12 NC); Batui River
at Singsing camp, 15 Oct. 1989, J. van Tol, 16 19
(RMNH). All apterous (fig. 82).

Description. — Apterous form. Dimensions.
Length d 3.20-3.32, 9 3.22-3.35; width d 1.2-1.3,
2 1.2-1.3; width of head d 0.76-0.80, 2 0.77-0.80;
width of pronotum d 1.18-1.30, 2 1.26-1.32.

Colour, dorsally dark brown, broad orange yellow
band anteriorly on pronotum extending to propleura,
covering anterior quarter of pronotum. Dorsal two
thirds of connexivum orange yellow, in male caudal
half of abdominal tergite 7 and posterior segments
orange yellow. Venter brownish, laterally (including
acetabula) and caudally orange yellow. First antennal
segment pale yellow, other segments brown. Legs ba-
sally (up to halfway femur) yellowish, apically casta-
neous. ;

Posterior half of jugum of head, anterior half of
propleura, thoracic sternum, dorsal margin of aceta-
bula and abdominal sternites 1-6 with small black
denticles. On darker parts of venter difficult to see be-
cause of lack of contrast. Dorsum rather sparsely
clothed with golden pubescence, longer and thicker
towards apex of abdomen, especially in male. Vertex
anteriorly with short dark setae, sparse long dark setae
on head, sides of thorax and apex of abdomen. Disk
of pronotum with large impressed pits, about 25/0.25
mm. Antennae and legs with the usual pubescence
and setae.

Length of antennal segments I-IV d 0.74 : 0.40:
0.49 : 0.41, 2 0.70 : 0.39 : 0.49 : 0.41. Pronotum
long, covering mesonotum, length : width 0.8 : 1.1.
Length of metanotum on midline, 0.12. Abdominal
tergites 1-6 subequal in length (0.15-0.20), tergite 7
longer (0.35-0.40).

Male. Posterior trochanter with one larger, two
small teeth and a group of about five small warts.
Posterior femur three times as long as wide
(1.35/0.45), on ventral margin proximally with about
20 minute and closely set teeth, followed about half-
way by a large spine about as long as half the width of
femur, with a row of 9-10 spines of decreasing length
distally. Anteroventrally a row of 5 smaller spines lo-
cated in distal part. Larger spines black tipped, smal-
ler spines and teeth entirely black or nearly so (fig.
64). Posterior tibia virtually straight, armed beneath
with a double row of about (2x) 15 pegs. Connexiva
more or less horizontal, gradually converging poste-
riad. Abdominal sternite 7 laterally compressed in
posterior two thirds, resulting in a broad distinct me-
dioventral keel in posterior part. Keel accentuated by
a dense fringe of long pilosity (fig. 62). Genital seg-
ments prominent, 1.5 times as long as sternite 7, ster-
nite 8 with an indistinct ventral keel. Parameres long
(fig. 63), apically curved mediad.

Female. Posterior trochanter without small teeth.
Posterior femur four times as long as wide
(1.30/0.30), row of minute teeth absent, row of larger
spines essentially as in male, anteroventral row of 2-4
smaller spines located in basal part. Posterior tibia as
in male. Connexiva more or less vertical, on tergites 4
and 5 slightly curved inward, thickened and beset
with dense pilosity. Apex of connexiva truncate, dor-
soapically with a small tuft of pilosity (figs. 66-67).
Tergite 8 only slightly slanting ventrad, proctiger
pointing ventrad in most specimens. Gonocoxa |
well visible, laterally somewhat compressed.

Macropterous form, unknown.

Etymology. — Tropidatos, greek adjective meaning
‘provided with a keel’, referring to the well developed
keel on abdominal sternite 7 of male.

Comparative notes. — Similar to À wallacei
Polhemus & Polhemus which differs in the male by a
less well developed keel on sternite 7 and shorter pa-
rameres (fig. 76) and in the female by connexiva not
thickened on segments 4 and 5 and apical segments
(8 and 9) hardly visible, withdrawn into segment 7.
Abdominal segment 8 and the long paramere remind
somewhat of R. meikdelyi Polhemus & Polhemus
from Ambon but in detail there are many differences
of which the more strongly inflated posterior femur
(only slightly over twice as long as wide in males, th-
ree times as long as wide in females) is one of the most
distinct. Finally À. lorelinduana Polhemus & Polhe-

Nieser & CHEN, Rhagovelia of Sulawesi

mus has a strongly developed keel on sternite 7 of
male but this species has an aberrant paramere and
proctiger (fig. 23) in male and a posteriorly prolonged
abdomen with the connexiva bent over its apex in fe-
male.

Discussion

Polhemus & Polhemus (1988) distinguished sever-
al groups within the genus, of which three occur in
Sulawesi. The sarawakensis-group, however, has been
recorded by a single macropterous specimen of R. sa-
marinda only. So we will restrict our discussion to the
orientalis- and papuensis- group.

The orientalis-group is a small group of about eight
species distributed in the Philippines and Sulawesi.
The main synapomorphy for this group is the hemi-
elytral venation, three cells, of which the apical one
reaches into the apical quarter of hemielytron (figs.
79-80). This seems a rather fickle character in view of
the variability of hemielytral venation in some species
and even between hemielytra of a specimen (figs. 53-
55). As the general appearance of at least the Sulawesi
species of this group is homogeneous and, apart from
males of R. sulawesiana, distinct from the species of
the other group, it seems, nonetheless, justified. In
Sulawesi this group is represented by three species: R.
celebensis, R. daktylophora and R. pseudocelebensis.

The remaining Sulawesi species belong to the pa-
puensis-group, a large group with a wide distribution:
India and Sri Lanka through Taiwan, the Philippines,
Northern Borneo, Sulawesi and Maluku (the
Moluccas) to New Guinea, Solomons and Australia.
According to Polhemus & Polhemus (1988) this
group is probably polyphyletic. A more restricted pa-
puensis-group is characterized by a strongly inflated
hind femur, a curved hind tibia with one or more lar-
ger subapical teeth and a comparatively long third an-
tennal segment. R. papuensis Lundblad (Papua New
Guinea), R kawakamii (Matsumura) (Northern
Borneo), R. grayi, R. horaia and R. lorelinduana (Sula-
wesi) and R. kalami (Buton) are all representatives of
this group. But as many species of this group (s. 1.) are
still to be described and the Sulawesi fauna constitu-
tes only a fraction of it an analysis of this group here
would be premature.

The distribution of most species seems to be rather
restricted, although one should not forget, looking at
these small maps, that the distance between the
northern and southern points of the main island of
Sulawesi is over 1000 km and the west-east length of
the northern peninsula c. 600 km. Secondly, more or
less extensive collecting has been done in a rather
small number of restricted areas only. The maps are
based on the data from Polhemus & Polhemus
(1988) and our own.

279

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

The three species of the orientalis-group have, so
far, neatly separated distributions (fig. 81). The most
widely distributed species seem to be R. pruinosa and
R wallacei (figs. 83, 82). R. kastanoparuphe and R.
pseudocelebensis are interesting as they occur with R.
pruinosa both on the main island and on Buton (figs.
81, 83-84), R. trichota has been found on both sides
of Pegunungan Mengkoka. The remaining species are
as yet only known from restricted areas. Nearly all
specimens have been collected by experienced collec-
tors. So we may assume that having been collected
only on one or very few close localities of these local
species indeed reflects restricted distribution rather
than more cryptic habits. Available data suggest that
the widely distributed species tend to be lowland spe-
cies, and most, but not all, of the restricted species
live on higher ground. A definite conclusion in this
respect has to wait until more data become available.
Likewise our data are not sufficient to establish altitu-
dinal separation between species occurring geographi-
cally at the same ‘points.

Polhemus & Polhemus (1990) have postulated
‘centres of species endemism’ in Sulawesi. As average
areas of distribution of species of various genera of
Gerromorpha and Nepomorpha in Malesia are not
yet known, we prefer to avoid the predicates endeme
and endemism for the present. Although their centres
look somewhat like areas in which more intensive col-
lecting has been done compared to the remaining of
Sulawesi, our results with Rhagovelia and Enithares
(Nieser & Chen 1991) coincide with the regions with
different faunal elements of water bugs in Sulawesi
they postulated. Our present data also support their
supposition that Sulawesi Tenggara (apparently in-
cluding Buton) and the eastern peninsula of Sulawesi
Tengah probably are such regions. It should, how-
ever, been pointed out that our results with the gerri-
ne genus Limnometra (Nieser & Chen 1992), which

280

also contains mostly stream inhabiting species, do not
strongly support this pattern. In Zimnometra there are
fewer species of which relatively more occur at least in
two of these regions.

REFERENCES

Bacon, J. A., 1956. A taxonomic study of the genus
Rhagovelia of the Western Hemisphere. — Kansas
University Science Bulletin 38:695-913.

Chen, P. P. & N. Nieser, 1992. Gerridae, mainly from
Sulawesi and Pulau Buton (Indonesia). — Tijdschrift voor
Entomologie 135: 145-162.

Hungerford, H. B. & R. Matsuda, 1961. Some new species
of Rhagovelia from the Philippines. — University of
Kansas Scientific Bulletin 42: 257-279.

Lundblad, ©., 1936. Die altweltlichen Arten der
Veliidengattungen Rhagovelia und Tetraripis. — Arkiv för
Zoologi 28A(21): 1-21, 13 pls.

Lundblad, O., 1937. Einige neue oder wenig bekannte osta-
siatische Rhagovelia-Arten. — Entomologisk Tidskrift 58:
1-9.

Nieser, N., & P. Chen, 1991. Naucoridae, Nepidae and
Notonectidae, mainly from Sulawesi and Pulau Buton
(Indonesia). — Tijdschrift voor Entomologie 134: 47-67.

Nieser, N., & P. Chen, 1992. Revision of Limnometra Mayr
(Gerridae) in the Malay Archipelago. — Tijdschrift voor
Entomologie 135: 11-26.

Polhemus, J. T., & D. A. Polhemus, 1988. Zoogeography,
ecology, and systematics of the genus Rhagovelia Mayr
(Heteroptera: Veliidae) in Borneo, Celebes, and the
Moluccas. — Insecta Mundi 2: 161-230.

Polhemus, J. T., & D.A. Polhemus 1990. Zoogeography of
the aquatic Heteroptera of Celebes: regional relationships
versus insular endemism. — In: Knight, W. J. & J. D.
Holloway (Eds.), Insects and the rain forests of South
East Asia (Wallacea). Royal Entomological Society of
London.

Received: 7 October 1992
Revised manuscript accepted: 1 June 1993

Appendix 1
Rhagovelia species collected in Sabah

Rhagovelia kawakamii (Matsumura)

Kotovelia kawakamii Matsumura, 1913: 99.
Rhagovelia kawakamit, Polhemus & Polhemus 1988: 206-
207, figs. 74-80, 227 (redescription, synonymy).

Material. — EAST MALAYSIA: Sabah, Pulau Ranggi, logging
camp, 7.10N 117.10E, 23. IV. 1987, leg. J. Huisman 2d
1 apterous (1d 19 RMNH, 1d NC).

Distribution. — Taiwan, Philippines, N. Borneo.

Rhagovelia samardaca Polhemus & Polhemus

Rhagovelia samardaca Polhemus & Polhemus, 1988: 200-
201, figs. 194-200, 226.

Material. — EAST MALAYSIA: Sabah, Lahad Datu, 60 km
W of Danum Valley field Centre at junction of Sg. Segama
and Sg. Palum Tambun, 4°58’N 117°48’E, 150m, bridge of
Segama, 26. IV. 1987, 18.30-21.00h, clearing edge of un-
touched evergr. lowl. rainforest, leg. van Tol & Huisman,
46 29 apterous (RMNH, 1d 1 NC).

Distribution. — Sabah.

Rhagovelia silau Polhemus & Polhemus

Rhagovelia silau Polhemus & Polhemus, 1988: 184-186,
figs. 99-105, 227.

NIESER & CHEN, Rhagovelia of Sulawesi

Material. — EAST MALAYSIA: Sabah, 105 km S of Beaufort:
Long Pa Sia area, Sg. Ritan, 4°24’N 115°42’E, 1160m, un-
disturbed evergreen tropical forest, 8.iv.1987, J. van Tol &
J. Huisman, 18 39 (1d 19 apt, 19 macr. RMNH, 19 apt.
NC).

Distribution. — Previously recorded from Sabah in
mountainous terrain, localities 900-2100m asl.

Rhagovelia tawau Polhemus & Polhemus

Rhagovelia tawau Polhemus & Polhemus, 1988: 187-188,
figs. 15-22, 224.

Material. — EAST MALAYSIA: Sabah, 70 km West of Lahad
Datu, Main Trail West 12, 180m, narrow creek, 2.xii.1989,
M. J. & J. P. Duffels, sample Sab. 52, 54 19; same, 4 km
S Main Trail West 5, near Sungai Segama, 150m, middle si-
zed stream and waterfall, 3.xii.1989, M. J. & J. P. Duffels,
sample Sab. 54, 5d 29 (ZMA, 3d 19 NC); 60 km W Lahad
Datu, Danum Valley Field Centre, road DVFC-Kg Silam nr
km 68.3, 4°58 N 117°48’E, at junction of Sg. Sagama and
Sg. Palum Tambun, 150m, disturbed evergreen lowland
rainforest, 26.iii.1987, leg. van Tol & Huisman, 1d. All ap-

terous.

Distribution. — So far only known by the type se-
ries from Tawau (near Kalabakan) which is also in the
eastern coastal region of Sabah, close to the border
with Kalimantan Timur.

281

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

BOOK REVIEW

Kuchlein, J. H. (with collaboration of J. H. Donner), 1993.
De Kleine Vlinders. Handboek voor de Faunistiek van de
Nederlandse Microlepidoptera. - PUDOC, Wageningen.
715 pp., 1372 maps, 8 colour plates, textfigs. [ISBN 90-
220-1038-4] [The small moths. Manual for the Faunistics
of Netherlands Microlepidoptera, in Dutch, with 5 pp.
English summary]. Price Hfl. 150.- (ca US $ 80).

This book deals with the Microlepidoptera of the
Netherlands, the main body being distribution maps of all
1372 species (4 missing maps are provided on a separate er-
ratum sheet).

The first 170 pages contain lengthy general chapters,
dealing respectively with Morphology of adult, larva, feed-
ing traces (mines), Ecology, Taxonomy and zoogeography,
Human influence, Faunistics, and a key to families based on
adults with a bibliography of key-works.

Apart from the maps, the main part of the book contains
a checklist with annotations about faunistic status (incor-
rectly referred to as ecological status), indication of occur-
rence in the periods 1850-1900, 1900-1950 and later than
1950, number of provinces, number of grid squares, indica-
tion of abundance and codenumbers for hostplants. Further
there are annotations for part of the species, usually explain-
ing past identification problems, or indication of first find-
ings.

A final chapter contains some general results and conclu-
sions. Finally there is a long English summary, making the
book also accessible for foreign readers.

The book is attractively made, well printed, and the
colour plates, with mounted moths, mostly from the.collec-
tion of J. C. Koster, are well made, although the black back-
ground sometimes tends to make the colours of the moth
too dark. One colour plate depicts leaf-mines.

The book contains a wealth of interesting information,
and there lies in my opinion the main problem of the book:
it is as if the author could not make a choice whether to
write a general manual about microlepidopteran studies or
to present the results of faunistic research on the species in
the Netherlands. The book would have been much nicer
and easier to use if a choice had been made before writing.
The general introduction chapters are really more a student
textbook, certainly with interesting information, but to be
found in many other textbooks as well, albeit with fewer ex-
amples from the Lepidoptera. The connection between the
lengthy introduction to ecology for instance and the individ-
ual species is difficult to find. In fact there are hardly any
notes about the habitats and ecology of individual species.
The only ecology found is a list of hostplant species (and
then with numbers, which have to be searched in the appen-
dix for the correct name), without any details.

The most interesting and original part of the introducto-
ry chapters is in my opinion the historical chapter on Dutch
Microlepidopterists. The key to families could be useful,
and the colour plates certainly are helpful. It is a pity that the
family names are not given in the captions for the plates.
One photograph has mistakenly been used twice: plate 5: 4
(Mompha epilobiella) shows the photograph of Bucculatrix
ulmella, the same as plate 3: 1.

The bibliography of key works is very long, for new work-
ers disappointingly long. Nowhere a choice has been made,
nor annotations given about the usefulness of a book or arti-

282

cle. Some references seem completely misplaced, like keys
for New Zealand species for groups where recent European
keys are available as well. On the other hand some important
titles are missing.

The checklist is an important summary of present know-
ledge of Dutch faunistics: it is therefore to be regretted that
here and in the species annotations there are many mistakes
or inaccuracies. Despite a well written introduction to zoo-
logical nomenclature, the author nowhere indicates his
source for the nomenclature used. Several recently proposed
changes are not incorporated, but some are (like the generic
name Luquetia Leraut). Notable mistakes are the use of
Episcardia for Haplotinea, shown to be incorrect by
Robinson in 1984, the use of the generic name Glyphipteryx
Curtis, rejected by the ICZN in 1986 and the use of the re-
placement name Nemophora violaria (Razowski), which is
only needed (because of secondary homonymy) when used
in the genus Adela. Such mistakes could easily have been
avoided by closer contact with specialists.

The maps show the distribution in 5 X 5 km grid cells
with a single symbol. Although the databank used contains
information about the date of the records, all records have
the same symbol, so a well dotted map may also show the
distribution of about 100 years ago. It would have been bet-
ter to indicate periods, using different symbols as is general
practice in similar atlasses. Some rapidly declining species
can not be identified from these maps, nor from the text. In
some cases there is considerable doubt about the correctness
of some dots: the large number of dots in the species
Stigmella samiatella and S. ruficapitella could only be ex-
plained by the inclusion of records based on mines.
However, there are no reliable characters separating these
mines, nor the larvae. In some cases I am missing previously
published records, in others there are dots for records which
have been shown to be incorrect in literature.

The presentation chosen is partly impractical because in
order to find all information about a species, one needs to
switch from the map to the list, for which one needs the ap-
pendix to understand the hostcodes, and the introduction to
understand the other codes. The list also indicates whether
there is an annotation for the species, but the caption to the
map does not give this reference.

Finally, I would like to comment on the choice of the spe-
cies treated: it is unfortunate that this book chooses to in-
clude only the Microlepidoptera in the old sense and exclud-
ing such related groups as Psychidae or Sesiidae. On the
other hand one could wonder about the inclusion of single
import events of species found on Bonsai trees imported
from Japan and China. These species have been found by
the Plant Protection Service, and the trees were destroyed
subsequently. Such species do not belong on the list of spe-
cies occurring in The Netherlands, otherwise all species in-
troduced alive in our country could be listed as well with the
same argumentation, including those introduced on pur-
pose by scientists or breeders! The maps of such species do
not give any information (some of the maps even do not
contain any dot, yet they are printed).

In conclusion: a book with much faunistic information,
useful for Microlepidopterists in Europe, but to be used
with care. The long introduction is only of use for readers
with knowledge of Dutch. In my opinion too expensive de-
spite the excellent printing.

[E. J. van Nieukerken]

O. A. SETHER & T. ANDERSEN

Museum of Zoology, University of Bergen

LOBOSMITTIA, A NEW GENUS OF ORTHOCLADS
FROM TANZANIA AND TURKEY (DIPTERA:

CHIRONOMIDAE)

Sæther, O. A. & T. Andersen, 1993. Lobosmittia, a new genus of orthoclads from Tanzania and
Turkey (Diptera: Chironomidae). — Tijdschrift voor Entomologie 136: 283-287, figs. 1-10.
[ISSN 0040-7496]. Published 10 December 1993.

The genus Lobosmittia is erected for a new species from the West Usambara Mts in NE
Tanzania, L. basilobata and for Pseudosmittia invaginata Caspers & Reiss, 1989 from Turkey.
The genus differs from other orthoclads with bare eyes, wings and squama; by completely lack-
ing acrostichals, median microtrichial tuft or less sclerotized median area on scutum; by having
costa not or only slightly extended; fine, triangular, strongly pointed anal point bare of micro-
trichia; and gonostylus with either a basal or preapical lobe. The Parakiefferiella group of gene-
ra or Acamptocladius Brundin and related genera are the most likely closest relatives.
Department of Systematic Zoology, Museum of Zoology, University of Bergen, Musépl. 3,

N-5007 Bergen, Norway.

Key words. — Chironomidae, Orthocladiinae, new genus, Tanzania, Turkey.

INTRODUCTION

During an expedition by the Museum of Zoology,
University of Bergen, to the montane evergreen fo-
rests in the West Usambara Mountains, NE Tanzania
in the autumn of 1990, several interesting new gene-
ra and species of chironomids were collected together
with a few genera new to the African continent
(Andersen & Szther 1993, in press a, b, Sæther &
Wang 1992). One of the new genera had what in
some specimens looked like a double gonostylus.
However, other views showed that it was a basal lobe.
Other characteristics made it clear that the species
could not be placed in any presently recognized ge-
nus. The senior author, together with L. C.
Ferrington jr., Lawrence, Kansas, presently is revising
the genus Pseudosmittia Goetghebuer. Many species
in that genus are tentatively placed since the genus
has not been well defined. When examining types it
was found that Pseudosmittia invaginata described by
Caspers & Reiss (1989) could be placed in the same
genus as the species from Tanzania.

The field work, which included an extensive use of
Malaise traps and sweep nets, was mainly conducted
in the Mazumbai Forest Reserve in the eastern part of
the West Usambara Mts. A thorough description of
the vegetation in these mountains is given by Iversen
(1991). Our Malaise trap localities along the Kaputu
Stream near Mazumbai are described in Andersen &
Johanson (in press).

METHODS AND TERMINOLOGY

The material examined was mounted on slides fol-
lowing the procedure outlined by Sæther (1969). The
general terminology follows Sæther (1980). The me-
asurements are given as ranges followed by mean and
the number measured (n) in parentheses.

The types of Lobosmittia basilobata sp. n. are de-
posited in the Museum of Zoology, University of
Bergen, Norway (ZMBN).

Lobosmittia gen. n.

Type species. — Lobosmittia basilobata sp. n. by
present designation. Other included species:
Lobosmittia invaginata (Caspers & Reiss, 1989: 123,
Figs. 11-13, as Pseudosmittia invaginata), comb. n.

Diagnostic characters. — The bare eyes, wing mem-
brane and squama, combined with complete lack of
acrostichals, microtrichial tuft on median hump or
sclerotized area on scutum, not or only moderately
extended costa, and pointed, triangular anal point,
free of microtrichia will separate the genus from oth-
er orthoclads.

Etymology. — From the Greek lobos, a lobe, and
Smittia, a related genus of Orthocladiinae and a com-
mon ending among orthoclads, referring to the basal
or subapical lobes of the gonostyli of the two includ-
ed species.

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TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Description
Small species with wing length 0.7 - 1.1 mm.
Eyes bare, without dorsomedial elongation.

Coronal suture well developed ending in frontal pro-
jection. Temporal setae consisting of weak inner ver-
ticals and strong outer verticals. Tentorium and stipes
normal. Antennae of male with 13 flagellomeres; an-
tennal groove starting on flagellomere 3; sensilla
chaetica on flagellomeres 2, 3 and 13; apex without
apical setae.

Antepronotum with median lobes narrowed medi-
ally and slightly separated, with 1 lateral seta.
Dorsocentrals several; acrostichals, microtrichial tuft,
hump or unsclerotized median area of scutum absent;
prealars few, in one or two groups; supraalars present
or absent. Scutellum with few setae. Humeral pit not
distinctive.

Wing membrane bare, with very fine punctuation
of microtrichia; veins brown. Anal lobe weak or ab-
sent. Costa not to moderately extended; R,,, running
in the middle between R, and R,,,, ending close to
Rs Ry, ending clearly proximally of end ofM,,;; Cu,
sinuate; FCu far distally of RM; postcubitus extend-
ing beyond FCu; anal vein not reaching FCu.
Brachiolum with 1 seta, other veins bare. Squama
bare. Sensilla campaniformia about 8-10 on base of
brachiolum, 3 below seta, and about 8-10 at apex of
brachiolum; 1 at base of subcosta, 1 on FR and 1 at
base of R..

Tibial spurs and tibial comb normal. Pulvilli ab-
sent, at least in L. invaginata.

Tergite IX with few to several setae at base of trian-
gular, sharply pointed anal point which is bare of se-
tae and microtrichia. Sternapodeme with weak or no
oral projections. Phallapodeme normal. Virga well
developed, consisting of few needle-like spines.
Gonocoxite with long, relatively low inferior volsella.
Gonostylus either with a basal crista dorsalis appear-
ing as a basal appendage in some views or with a long
apically angled crista dorsalis making the gonostylus
appear to have a deep apical invagination.

Female, pupae and larvae unknown.

Systematics

In the key to adult males of Orthocladiinae by
Cranston et al. (1989) Lobosmittia will key to couplet
98. It will not key further since the pulvilli apparent-
ly are absent or vestigial, while the anal point extends
from the posterior margin of tergite IX. Caspers &
Reiss (1989: 12) discussed the placement of their spe-
cies as Pseudosmittia invaginata on the base of the si-

milarity of Pseudosmittia Goetghebuer and Prosmittia
Brundin. Cranston & Oliver (1988) even synonymi-
zed the two genera based on the fact that Prosmittia
nanseni Kieffer (Kieffer 1926: 82) is a good
Pseudosmittia. Oliver (1963: 177) first placed nanseni
in the genus Prosmittia apparently based on the simi-
larities in the hypopygia between nanseni and
Prosmittia jemtlandica (Brundin, 1947). Reexamin-
ation of the type of P. nanseni by Sæther et al. (1984:
270) did not change the placement of the species.
However, the wing venation could not be discerned
on the damaged male. The senior author together
with Dr. L. C. Ferrington jr., Lawrence, Kansas, pre-
sently is revising the genus Pseudosmittia and we have
examined P. jemtlandica as well as a large amount of
P. nanseniand most types of Pseudosmittia. All species
of Pseudosmittia have 2 or occasionally 4-11 (as in P.
nanseni) short, but relatively strong acrostichals, with
distinct sockets, in a pale, less sclerotised median area
on the scutum. There is never a median microtrichial
tuft as in Parakiefferiella Thienemann, and no me-
dian tubercle, but occasionally there is a hump in so-
me teneral specimens. On the wing, vein R,,, may
end slightly distal to end of M,,,, but mostly ends well
proximal of the end of M... In Prosmittia jemtlandi-
ca there is no sign of acrostichals, microtrichal tuft,
hump or pale area on scutum, and R,,, ends far dis-
tally of the end of M,,, almost reaching the wing ap-
ex. The illustration by Brundin (1947: fig. 16) of
Prosmittia jemtlandica (as Pseudosmittia jemtlandica)
is correct and not erroneous as previously presumed.
The genus Prosmittia Brundin (1956: 58, 165) clear-
ly has to be resurrected. The wing venation, however,
appears to place it close to Unniella Sether, but not to
Pseudosmittia nor to Lobosmittia.

Lobosmittia, however, has an anal point placed on
the posterior margin of tergite IX and does not appear
closely related to any of the above mentioned genera.
Boreosmittia Tuiskunen appears to be the most simi-
lar genus. However, without knowledge of the imma-
tures a more definite placement is as yet not possible.

Lobosmittia basilobata sp. n.
(figs. 1-10)

Type material. — Holotype 6, TANZANIA, Tanga region,
West Usambara Mts, Mazumbai, Kaputu Stream, 1640 ma.
s. l., sweep net at waterfall, 28.x.1990, ZMB's Tanzania
Expedition (ZMBN No. 156).— Paratypes: 4d, as holotype;
1d as holotype except Malaise trap loc. 2, 1650 m a.s. |.,
2.-6.xi.1990; 26 as holotype except Malaise trap loc. 10,
1420 m a. s.1., xii.1990.

Figs. 1-10. Male imago of Lobosmittia basilobata gen. n., sp. n. — 1, head; 2, thorax; 3, third palpal segment; 4, tentorium and
stipes; 5, wing; 6, hypopygium, with dorsal aspect to the left, ventral to the right; 7, anal point, lateral view; 8-10, gonosty-

lus, different views.

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SÆTHER & ANDERSEN: Lobosmittia gen. n.

285

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Diagnostic characters. — The basal lobe of the go-
nostylus easily separates L. basilobata from L. invagin-
ata. Other characteristics of L. basilobata include ab-
sence of supraalars, fewer prealars and inner verticals

and more numerous sensilla clavata at apex of the

third palpal segment.

Etymology. — From the Latin basis, base, pedestal,
and New Latin, /obatus, lobed, referring to the shape
of the gonostylus.

Description

Male imago (n = 7-8, except when otherwise sta-
ted). — Total length 1.34-1.46, 1.40 mm (6). Wing
length 0.75-0.83, 0.81 mm. Total length / wing
length 1.57-1.87, 1.73 (6). Wing length / length of
profemur 2.73-3.08, 2.88. Coloration brownish
black with yellowish shoulders and anepisternum.
Legs yellowish brown.

Head (Fig. 1). AR 0.39-0.49, 0.42. Ultimate fla-
gellomere 123-171, 141 um long. One specimen
with division between flagellomere 12 and 13 indis-
tinct, if antenna regarded as 12-segmented AR 0.56.
Temporal setae consisting of 2-3, 2 mostly widely
separated, weak inner verticals; and 3 strong outer
verticals. Clypeus with 4-6, 5 setae. Tentorium (Fig.
4) 80-94, 85 um long, 13-18, 15 um wide. Stipes
(Fig. 4) 57-66, 62 um long, 21-27, 25 um wide. Palp
segments lengths in um: 14-18, 15; 18-23, 21; 27-39,
34; 32-41, 38; 46-66, 59. Third palpal segment (Fig.
3) with 6-7, 6 (6) sensilla clavata. Coronal suture end-
ing in frontal projection.

Thorax (fig. 2). Antepronotum with 1 lateral seta.
Dorsocentrals 8-11, 9; prealars 3; supraalars absent.
Scutellum with 2 setae.

Wing (fig. 5). VR 1.31-1.45, 1.40. Costal exten-
sion 23 -62, 35 mm long.

Legs. Spur of front tibia 29-39, 34 im long; spurs
of middle tibia 14-18, 16 um (6) and 13-18, 15 um
long; of hind tibia 32-39, 35 [um and 11-16, 14 um.
Width at apex of front tibia 16-18, 17 um; of middle
tibia 16-21, 19 um; of hind tibia 25-34, 30 um. Hind
tibial comb of 13-15, 14 setae; shortest setae 18-25,
21 pm long; longest setae 26-34, 31 um long. Lengths
of front, middle and hind femora (in Um) as: 252-
306, 282; 279-315, 306; 288-324, 308; of tibiae:
266-315, 293; 288-360, 325; 315-360, 339.
Tarsomeres 1 and 2 of hind legs 171 um (1) and 90
Um (1) long, other tarsomeres lost.

Hypopygium (fig. 6). Anal point 18-27, 21 pm
long; with 9-14, 10 setae at base (fig. 7); laterosternite
IX with 3 setae. Phallapodeme 39-48, 45 um long,
transverse sternapodeme 34-43, 41 um long. Virga
14-19, 16 um long. Gonocoxite 91-101, 97 um long;
with long inferior volsella apically hooked in some

views. Gonostylus 39-45, 42 pm long; basal lobe end-

286

ing 21-27, 24 um from base (figs. 8-10); megaseta 9-
14, 10 pm long.

Lobosmittia invaginata (Caspers & Reiss) comb. n.

Pseudosmittia invaginata Caspers & Reiss, 1989: 123, figs.
11-13. — Holotype d, TURKEY: Kars province, W.
Sarikamis, Soganli railway station, 2100 m a. s. l., 5.vii.
1985, W. Schacht (Zoologischen Staatssammlung,

Miinchen). [examined]

Diagnostic characters. — The subapical invagina-
tion of the gonostylus easily separates L. invaginata
from L. basilobata. Other characteristics of L. inva-
ginata includes the presence of 1 supraalar, 6 inner
verticals and 1 sensillum clavatum at the apex of the
third palpal segment.

Description

The species is well described by Caspers & Reiss
(1989: 123, figs. 11-13). However, this description
contains a few mistakes. There are 6 weak inner verti-
cals reaching from near the middle of the head to the
position of the outer verticals. There also are 3 stron-
ger outer verticals, apparently 8 setae on the clypeus,
1 lateral antepronotal and 1 supraalar. The first palp
segment is shorter than the second, our measure-
ments are 23 Mm and 30 um. The leg segments
lengths appear to have been measured as total lengths,
not following Schlee (1966), as especially the femora
and ta, are about 15% too long compared with our
measurements, while ta, to ta, are quite close.

ACKNOWLEDGEMENTS

The expedition to Tanzania was funded by The
Norwegian Council for Science and the Humanities
(NAVE). Financial support was also given by
Nansenfondet. Thanks are also 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 sup-
port. Gladys Ramirez made the slide preparations of
Lobosmittia basilobata.

REFERENCES

Andersen, T. & K. A. Johanson, (in press). Caddis flies
(Trichoptera) from a mountain rain forest in NE
Tanzania. — In: Otto, C. Proceedings of the 7th
International Symposium on Trichoptera. — Universal
Book Services, The Netherlands.

Andersen, T. & O. A. Sæther, 1993. Lerheimia, a new genus
of Orthocladiinae from Africa (Diptera, Chironomidae).
— Spixiana 16: 105-112.

Andersen, T. & O. A. Sæther, (in press a). Usambaromyia

nigrala gen. n., sp. n., and Usambaromyiinae, a new sub-
family among the Chironomidae (Diptera). — Aquatic
Insects 15.

Andersen, T. & O. A. Sæther, (in press b). Colosmittia clav-
ata gen. n., sp. n., a new orthoclad from the Usambara
Mts, Tanzania (Diptera: Chironomidae). — Journal of the
Kansas Entomological Society 66.

Brundin, L., 1947. Zur Kenntnis der schwedishen
Chironomiden. — Arkiv for Zoologi 39A (3): 1-95.

Brundin, L., 1956. Zur Systematik der Orthocladiinae
(Dipt. Chironomidae). — Report, Institute of Freshwater
Research, Drottningholm 37: 5-185.

Caspers, N. & F. Reiss, 1989. Die Chironomidae der
Türkei. Teil I: Podonominae, Diamesinae,
Prodiamesinae, Orthocladiinae (Diptera, Nematocera,
Chironomidae).— Entomofauna 10 (8/1): 105-160.

Cranston, P. S. & D. R. Oliver, 1988. Additions and correc-
tions to the Nearctic Orthocladiinae (Diptera:
Chironomidae). — Canadian Entomologist 120: 425-
462.

Cranston, P. S., D. R. Oliver, & O. A. Sæther, 1989. 9. The
adult males of Orthocladiinae (Diptera: Chironomidae)
of the Holarctic region - Keys and diagnoses. — In:
Wiederholm, T. Chironomidae of the Holarctic region.
Keys and diagnoses. Part 3. Adult males. — Entomologica
scandinavica Supplement 34: 165-352.

Iversen, S. T., 1991. The Usambara Mountains, NE
Tanzania: Phytogeography of vascular plant flora. — Acta
Universitatis Upsalensis, Symbolae Botanicae Upsaliensis

29 (3): 1-234.

SÆTHER & ANDERSEN: Lobosmittia gen. n.

Kieffer, J. J., 1926. Chironomiden der 2. Fram - Expedition
(1898-1902). — Norsk entomologisk Tidsskrift 2: 78-89.

Oliver, D. R., 1963. Entomological studies in the Lake
Hazen Area, Ellesmere Island, including lists of species of
Arachnida, Collembola and Insecta. — Arctic 16: 175-
180.

Sæther, O. A., 1969. Some Nearctic Podonominae,
Diamesinae, and Orthocladiinae (Diptera:
Chironomidae). — Bulletin of the Fisheries Research

Board of Canada 107: 1-154.

Sæther, O. A., 1980. Glossary of chironomid morphology
terminology (Diptera: Chironomidae). — Entomologica
scandinavica Supplement 14: 1-51.

Sether, O. A., J. E. Sublette & E. Willassen, 1984.
Chironomidae (Diptera) from the 2nd Fram Expedition
(1898-1902) to Arctic North America described by J. J.
Kieffer. — Entomologica scandinavica 15: 245-275.

Sæther, O. A. & X. Wang, 1992. Euryhapsis fuscipropes sp. n.
from China and Tokyobrillia anderseni sp. n. from
Tanzania, with a review of genera near /risobrillia Oliver
(Diptera: Chironomidae). — Annales de limnologie 28:
209-223.

Schlee, D., 1966. Präparation und Ermittlung von
Messwerten an Chironomiden (Diptera). Gewässer und

Abwässer 41/42: 169-193.

Received: 6 January 1993
Accepted: 20 September 1993

287

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Cr

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SE. OF ENTOMOLOGY

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Florence (Italy), August 25 - 31, 1996

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288

BERNHARD J. van VONDEL

Natuurmuseum Rotterdam

REVISION OE THE LAPATLUS SPECIES @F THE
ORIENTAL REGION EXCLUDING CHINA

(COREORTENA: HALIPEIDAE)

Vondel, B. J. van, 1993. Revision of the Liaphlus species of the Oriental region excluding
China (Coleoptera: Haliplidae). — Tijdschrift voor Entomologie 136: 289-316, figs 1-130.
[ISSN 0040-7496]. Published 10 December 1993.

The Oriental species of the Haliplidae are discussed. The species of the subgenus Liaphlus
Guignot are revised, while the few remaining species of the subgenus Halipluss. str. (in revision
by Holmen) and the genus Peltodytes Régimbart (already revised by Vondel 1992) are only dis-
cussed in general. Twenty-six species of Haliplidae are known from the Oriental region. H.
shanicus Guignot is established as a junior synonym of H. angustifrons Régimbart. Four new
species are described: Haliplus javanicus, H. nedungaduensis, H. samosirensis and H. srilankanus.
Most primary types have been studied. Lectotypes are designated for H. angustifrons Régimbart,
H. oceanicus Régimbart, H. pulchellus Clark and H. pulchellus var. indicus Régimbart. The lat-
ter is raised to specific rank. A check-list of all Oriental Haliplidae is provided, as well as a key
to the Liaphlus species in the Oriental region excluding China, and distribution maps of most

species.

B. J. van Vondel, Roestuin 78, 3343 CV Hendrik-Ido-Ambacht, The Netherlands.
Key words. — Haliplidae, Haliplus, Liaphlus, Oriental region, new species.

Since Zimmermann (1924) published his key to
the Haliplidae of the world, no revisional work has
been done regarding the Oriental species. The status
of the species described since is not always clear, so a
revision is necessary. The type material of most spe-
cies was studied, except for the three species described
by Vazirani, whose types appear to be completely un-
available for examination.

Female genitalia are not described, because they
usually are very uniform. Further research is necessary
to check if there are characters sufficiently reliable to
separate the species.

MATERIAL

About 280 specimens, including type-material, we-
re studied from the following collections: Natural
History Museum, London, UK (BMNH); B. P. Bishop
Museum, Honolulu, Hawaii, USA (BPBM); Canadian
National Collection, Ottawa, Canada (CNCI); Institut
Royal des Sciences Naturelles, Brussels, Belgium
(ISNB); Museum of Comparative Zoology, Cam-
bridge, Massachusetts, USA (MCZC); Muséum Natio-
nal d'Histoire Naturelle, Paris, France (MNHN);
Museum of Zoology, Lund, Sweden (MZLU); Natur-
historisches Museum Wien, Vienna, Austria (NHMV);
Naturhistoriska Riksmuseet, Stockholm, Sweden

(NHRS); Nationaal Natuurhistorisch Museum (Rijks-
museum van Natuurlijke Historie), Leiden, Nether-
lands (RMNH); United States National Museum of
Natural History, Washington D.C., USA (USNM);
Instituut voor Taxonomische Zoölogie (Zoologisch
Museum), Amsterdam, Netherlands (ZMAN); Zoolo-
gisk Museum, Copenhagen, Denmark (ZMUC); Zoo-
logische Staatssammlung, Munich, Germany (ZSMC).
Private collections: L. Hendrich, Berlin, Germany;
P. Mazzoldi, Brescia, Italy; B. J. van Vondel, Hen-
drik-Ido-Ambacht, The Netherlands.

METHODS

Dissecting, preparing and drawing the aedaegi as
well as the use of morphological terms follows the
methods described by Vondel (1991: 76).

Locality names are given, as far as possible, accord-
ing to the Times Atlas of the World (Comprehensive
edition 1983), but when different from original la-
bels, the modern name is given in parentheses. Names
of countries are used with their present-day boundar-
ies. Distribution maps are based on material exam-
ined. Literature records are included using different
symbols.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

SYSTEMATIC SECTION

Checklist of Oriental Haliplidae, including Taiwan
and China south of the river Chiang Jiang

Haliplus Latreille
subgenus Haliplus s.str.
japonicus Sharp, 1873 (*)

sauter: Zimmermann, 1924 (*)

subgenus Liaphlus Guignot
angustifrons Régimbart, 1892
shanicus Guignot, 1956 syn.n.
arrowi Guignot, 1936
chinensis Falkenström, 1932 (**)
diopus Guignot, 1955
diruptus Balfour-Browne, 1946
eximius Clark, 1863
modestus Zimmermann, 1924
emmerichi Falkenström, 1936
holmeni Vondel, 1991 (**)
indicus Regimbart, 1899 stat. n.
Javanicus sp. n.
kapuri Vazirani, 1975
kotoshonis Kano & Kamiya, 1931 (**)
manipurensis Vazirani, 1966
nedungaduensis sp. n.
perroti Guignot, 1950
philippinus Chapin, 1930
pruthii Vazirani, 1966
pulchellus Clark, 1863
oceanicus Régimbart, 1886
samosirensis sp. n.
sharpi Wehncke, 1880 (**)

srilankanus sp. n.

Peltodytes Régimbart
coomani Peschet, 1923 (***)
wui Gschwendtner, 1934
intermedius (Sharp, 1873) (***)
sinensis (Hope, 1845) (***)
variabilis (Clark, 1863)
koreanus Takizawa, 1931
sumatrensis Régimbart, 1885 (***)

The Oriental region as defined in this revision is
according to fig. 122. Species of the subgenus
Liaphlus, which are present in the south of China or
in Taiwan and not further in the Oriental region
(marked ** in the species-list) are already revised by
Vondel (1991) and not included in the following key.
The Oriental species of the genus Peltodytes (marked
*** in the species-list) were revised together with the
Palaearctic species (Vondel 1992). The few species of
the subgenus Haliplus s. str. (marked with * in the
species-list), which are present in the southwest of

290

China (A. japonicus Sharp) and in Taiwan (H. saute-
ri Zimmermann) are subject of a revision of that sub-
genus by Holmen (in prep.) and not further treated
here. Some species are present both in the Orien-
tal and Palaearctic regions. These species are included
in the following key, but only treated here in a gener-
al way, except H. pulchellus Clark, which is fully treat-
ed.

Key to the genera of Oriental Haliplidae

1. Elytra with sutural stripe on about apical half.
Last segment of maxillair and labial palpi longer
FAN PERL ATE SEP ments. E
A CCnUS Pee Oder Pin bart

- Elytra without sutural stripe or at most a weak
trace at the apex. Last segments of maxillair and
labial palpi much shorter than penultimate seg-

TIDE AICS) IE Me genus Haliplus Latreille

Remark. — Species of the genus Peltodytes are al-
ready revised (Vondel 1992) and are only mentioned
in the species-list.

Key to the subgenera of Haliplus

1. Hind tibia with setiferous striole on dorsal face.
Pronotum without basal plicae ....subg. Liaphlus
— Hind tibia without setiferous striole on dorsal
face. Pronotum with basal plicae …

linate da ole subg. Haliplus s.str.

Remarks. — Species of the subgenus Haliplus s.str.
are in the Oriental region present in the southwest of
China: Yunnan (7. japonicus) and in Taiwan (A.
sauteri). The species concerned are only mentioned in
the species-list, because they are subject of a revision
by Holmen.

Guignot (1955b) proposed a division of the subge-
nus Liaphlus into species-groups. In some cases I do
not agree with his interpretation of species. As far as
Oriental species are concerned the following notes
can be made. Haliplus diopus and H. pulchellus are
closely related, but placed by Guignot in different
groups. The latter, however, is placed by him with the
clearly different H. testudo (from Australia) in one
group.

H. arrowi and H. angustifrons are closely related,
but, again, placed by him in different groups.

Key to the Oriental species of the subgenus
Liaphlus:

Species known only from China or Taiwan are not
included in the key, but can be identified with
Vondel (1991).

VAN VONDEL: Oriental Haliplidae

Figs. 1-9. Haliplus angustifrons, lectotype. — 1, dorsal view; 2, antenna; 3, punctures near elytral base and suture; 4, dorsal side
of hind tibia; 5, prosternal process; 6, lateral view of prosternal process; 7, left paramere; 8, penis; 9, right paramere.

291

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Prosternal process with two longitudinal grooves

(OL CR 2
Prosternal process flat or impressed in the middle
(ie DA) n i 9
Éyiraimaculare (he 19) ren. 3
Elytra only narrowly darkened along suture
(ie OS) ere Nae H.nedungaduensis

Margins of elytra serrate in basal part (fig. 46) .4
Margins of elytra not clearly serrate in basal part

Distance between eyes 0.9-1.0X width of one
VE e specimens of H. pulchellus
Distance between eyes 1.1-1.5X width of one
eye
Metasternal process with transverse impression,
in which two pits can be recognised (fig. 50)
RE dou meen EL SN ree H. indicus

_Metasternal process with two separated pits (fig.

DID ORE SIRIA E Di TA eee H. diopus
Mesepimeron with about 4 strong punctures (fig.
92). Length less than 2.6 mm … … H. philippinus
Mesepimeron without or rarely with one or two
punctures. Length more than 2.6 mm
Pronotum usually not continuously margined,
sometimes traces of a margin in posterior part,
posterior corners usually clearly protruding. Basal
black band on elytra reaching fifth primary punc-
ture-row (fig. 98) … … specimens of H. pulchellus
Pronotum usually finely margined except near
anterior corner, posterior corners not or hardly
protruding. Basal dark band on elytra reaching at
least sixth primary puncture row
Elytral basal band brown, reaching margins when
seen from above, black marks very irregular.
Pronotum slightly darker in the middle (fig.
107). Length 3.3-3.5 mm
SE I ia H. samosirensis
Elytral basal band black, reaching sixth primary
puncture-row, black marks well defined.
Pronotum clearly darkened in the middle (fig.
55). Length 2.8-3.0 mm

10

Elytra without any markings
Elytra with markings

. Prosternal process broad and flat, narrowed

beteen procoxae, not narrowed between meso-
(elo) etal bite, Al) en nen. H. eximius
Prosternal process narrowed between pro- and
mesocoxae (see remarks below) ........... H. kapuri

. Prosternal process without anterior margin (fig.

Prosternal process pitted posteriorly (fig. 5)
EE specimens of H. angustifrons
Prosternal process canaliculate (see remarks be-

292

13.

16.

17

low ra aie an ae A N H. manipurensis
Two longitudinal markings on disc of pronotum
(see remarks below) nn H. pruthii
At most small markings on disc of pronotum ..

. Elytral markings not connected with darkened
steuren ion E I I re H. perroti
Elytral markings connected with darkened suture
(fig. 1)

. Prosternal process strongly grooved and with a

deep pit in apical part (fig. 5), elytral base usually

darkened

Prosternal process flat or weakly impressed in ap-

ical part (fig. 32), elytral base usually not dark-

ened (HE IS) er H. diruptus

Distance between eyes 0.8-0.9X width of one

eye. Base of pronotum usually wider than elytral

base, posterior corners rounded. Pronotum short,
width at base 2.1-2.3 X length in the middle,
strong and in basal corners coarse punctures.

Margins thicker than elytral margin near shoul-

ders. Basal dark band usually reaching to fifth

puncture-row (fig. 1). Anterior edge of proster-
num only margined near prosternal process.

About 38 sutural punctures, about 32 punctures

HV HITS CP EMATYÉTO WE H. angustifrons

Distance between eyes 1.0-1.1X width of one

eye. Width of pronotum at base about 2.0 X

length in the middle, margins about as thick as

elytral margins near shoulders . … … 17

Pronotal posterior corners clearly protruding (fig.

113). Anterior edge of prosternum completely

margined. About 33 punctures in first primary

row. Suture darkened to first primary puncture-
row. (felt 3) i ml Ai H. srilankanus

Pronotal posterior corners not protruding.

Anterior edge of prosternum only margined near

prosternal process. About 24 punctures in first

primary row. Suture darkened to first secondary

Pune (i 400) e H. arrowi

Remarks. — The species described by Vazirani (AH.

kapuri, H. manipurensis and H. pruthii) are only

kn

own to me by the original description. As I do not

always know how to interpret some parts of these de-
scriptions, especially concerning the shape of the
prosternal process, some uncertainty remains regard-
ing Vazirani s species.

DESCRIPTION OF THE ORIENTAL LIAPHLUS

Haliplus angustifrons Régimbart
(figs. 1-9)

Haliplus angustifrons Régimbart, 1892: 112. Lectotype d,

‘Konbir [Konbira, Ranchi distr., Bihar, India], P.

VAN VONDEL: Oriental Haliplidae

Figs. 10-18. Haliplus arrowi, holotype. — 10, dorsal view; 11, antenna; 12, punctures near elytral base and suture; 13, dorsal
side of hind tibia; 14, prosternal process; 15, lateral view of prosternal process; 16, left paramere; 17, penis; 18, right para-
mere.

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294

Cardon; Museum Paris, Coll. Maurice Régimbart, 1908;
type’ (MNHN) [examined].

Haliplus shanicus Guignot, 1956: 477. Holotype d, ‘S. Shan
States, Road 40 km E. of Taunggyi, 25.IX-13.X.[19]34,
Malaise’ (NHRS) [examined]. syn. n.

Haliplus angustifrons, Régimbart 1899: 187, Zimmermann
1920: 303, 1924: 140, Guignot 1936: 117, 1954: 563,
1955b: 295, Vazirani 1966: 128.

Haliplus shanicus, Vazirani 1966: 133.

Diagnosis

This species can be distinguished from the other
species with a posteriorly well impressed prosternal
process by the characters mentioned in the key.

Description

Length 2.9-3.8 mm, width 1.6-2.0 mm. Body
oval, widest in the middle (fig. 1).

Head: Yellow-brown to dark brown, vertex some-
times darkened, moderately strongly punctured.
Distance between eyes 0.8-1.0X width of one eye.
Antennae yellow-red (fig. 2), palpi yellow-red.

Pronotum: Yellow-red to rust-coloured. Lateral
borders straight to slightly concave in posterior half
and convex in anterior half, strongly margined.
Posterior corners clearly protruding. Moderately
strongly, in anterior corners and along base strongly
punctured. Impressed along base.

Elytra: Yellow-red to brown-red with extensive
brown markings (fig. 1). Primary puncture-rows
moderately strong, 25-33 punctures in first row.
Secondary punctures relatively strong. All punctures
darkened with yellow centre (fig. 3). Completely
margined.

Ventral side: Body yellow-brown to rust-coloured
or brown, elytral epipleura yellow-brown, legs yellow-
red to brown. Prosternal process strongly impressed
in posterior part, anterior edge usually not clearly
margined, strongly punctured, narrowed near coxae,
anteriorly about as wide as posteriorly (fig. 5), lateral
plicae complete (fig. 6). Metasternal process widely
and strongly impressed in the middle, moderately
strongly to weakly punctured (fig. 5). Metacoxal
plates strongly punctured, along suture with a weakly
punctured area. Setiferous striole on dorsal side of
hind tibia about 15 X length of tibia, longest of the
two tibial spurs about 23 X length of first tarsal seg-
ment (fig. 4).

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on ventral side. Penis and
parameres as in figs. 7-9.

VAN VONDEL: Oriental Haliplidae

Biology and distribution (fig. 123)

In rivers. In mixed primary forest. In watertank.
Up to an altitude of 1500 m.

India: Armachal Pradesh, Bihar, Karnataka,
Madhya Pradesh, Maharashtra, Uttar Pradesh, West
Bengal; Vazirani (1966) also gives Himachal Pradesh
and Orissa. Sri Lanka. Nepal. Burma.

Material examined. — India: Armachal Pradesh: 1 ex.,
Assam, Patkai Mts, Doherty (BMNH); Bihar: 1d, Lectotype;
Id, 12, Konbir [Konbira], P. Cardon, paralectotype
(MNHN); 16, 12, Konbir [Konbira, Ranchi distr.], P.
Cardon, Haliplus pulchellus Régimbart det. 1890 (ISNB); 42
ex., Chota-Nagpore [Chota Nagpur], Nowatoli, R. P.
Cardon, viii-ix.1896, ix-x.1896, iv-v.1897 (MNHN, RMNH);
Karnataka: 18, 19, Shimoga, Maissour, vi-vii.1897
(MNHN); Madhya Pradesh: 1 ex., Barway [Barwah?], P.
Cardon (BMNH); Maharashtra: 1 ex., Lonaula, 80 km E.
Bombay, 13.ix.1991, R. Schuh (NHMV); Uttar Pradesh: 16,
O, Almora Dn., Kumaun, vii.1920, H.G.C (BMNH); West
Bengal: 12, Sarda, F.W.C. (BMNH); no province known:
1d, Indes Or., 22-53 (MNHN). — Nepal: 22, Kathmandu

Valley, 7.u.1981, M. Jäch (NHMV, Vondel); 29,
Kathmandu Distr., Gokarna, 1.viii.1983, M.G. Allen,
4500’, mixed primary forest (BMNH); 12, Ktmd

[Kathmandu], Godavari [?], 15.viii.1967, 5000’; 2d, 29,
nr Birganj, Lothar, 12.ix.1967, 450 Ft. (CNCI); 8 ex.,
Godawari, 15 km S. Kathmandu, 1500 m, 29.iv.1984,
Wewalka (Mazzoldi). — Sri Lanka: 1d, Inginiyagala,
14.x11.1979, V. Mahler Jensen (ZMUC). — Burma: Holotype
3, H. shanicus, 1% allotype, 16 paratype, data as holotype
(NHRS); paratypes of H. shanicus, 16, 29, data as holotype
(MNHN); 1d, 19, Shan States, road 40 km E. of Taung-gyi,
25.ix-13.x.1934; 16, Shan States, Namhkok [?], 20.
viii.1934, Malaise, 900 m.; 1d, Shan States, Taung-gyi,
25.x.1934, Malaise, 1500 m, watertank (MNHN).

Haliplus arrowi Guignot

(figs. 10-18)

Haliplus arrowi Guignot, 1936: 117. — Holotype d, “Type,
3, Calcutta, Atkinson 92-3., Det. Dr. Guignot Haliplus
Arrowi Guign., Type’ (BMNH) [examined].

Haliplus arrow, Guignot 1955b: 294, Vazirani 1966: 131,
1975: 317.

Diagnosis

This species is very closely related to Haliplus an-
gustifrons, but has sparser elytral puncture-rows, the
eyes less close to each other, the posterior corners of
the pronotum less protruding and some other slight
differences, mentioned in the key.

Description
Length 2.8-3.1 mm, width 1.5-1.6 mm. Body
oval, widest in the middle (fig. 10).

Figs. 19-27. Haliplus diopus, holotype. — 19, dorsal view; 22, antenna; 21, punctures near elytral base and suture; 22, dorsal
side of hind tibia; 23, prosternal process; 24, lateral view of prosternal process; 25, left paramere; 26, penis; 27, right para-

mere.

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een
. © © :

bo

296

Head: Brown, moderately punctured. Distance
between eyes 1.1-1.2 X width of one eye. Antenna
(fig. 11) and palpi yellow to yellow-brown.

Pronotum: Yellow-brown. Lateral borders convex,
very finely margined, posterior corners not protrud-
ing. Width at base about 2.0 X length in the middle.
Fairly strongly, along the base very strongly and
densely punctured, punctures not darkened. In the
midddle a transverse somewhat elevated unpunctured
band.

Elytra: Yellow-brown, brown marking along suture
and part of base and on disc, marks in even intervals
(fig. 10). Primary puncture-rows sparse and moder-
ately strong, about 25 punctures in first row.
Secondary punctures moderately strong along suture
and rather weak in intervals. All punctures darkened
with a yellow centre.

Ventral side: Yellow-brown to brown, elytral epi-
pleura yellow, legs yellow-brown, femora brown to
dark-brown near coxae. Prosternal process strongly
impressed in posterior part and less impressed in an-
terior part, slightly narrowed near coxae, moderately
strongly and densely punctured (fig. 14), lateral plicae
complete (fig. 15). Anterior border of prosternum not
completely margined, margin near prosternal process
accompanied by four punctures in a row. Metasternal
process with a wide and very deep pit, weakly punc-
tured (fig. 14). Coxal plates densely punctured, ex-
cept near suture, posterior part of plates dilated.
Setiferous striole on dorsal side of hind tibia about
1/3 X length of tibia and with about 8 punctures,
longest of the two tibial spurs almost as long as first
tarsal segment (fig. 13).

Male: First three tarsal segments of fore and mid-
legs slightly widened, scaly hairs on ventral side. Penis
and parameres as in figs. 16-18.

Biology and distribution (fig. 124)
Biology unknown.
India: Bihar, Madras, Uttar Pradesh, West Bengal.

Burma. Laos.

Material examined. — India: Bihar: 1d, Lectotype
(BMNH); 2d, Calcutta, Atkinson, paratype (BMNH, MNHN);
2 ex., Pusa, Andrewes (BMNH); Uttar Pradesh: 1 ex.
Sirampur [Sitapur?], Elberling (ZMUC); West Bengal: 16,
Calcutta, Haliplus pulchellus det. dr Guignot (BMNH). —
Burma: 1 ex., Maymyo, H.I. Andrewes (BMNH). — Laos: 1
ex., betw. Vientiane and Luang Prabang, end of 1919, R. V.
de Salvara (BMNH). — Unlabelled: 3 ex. (BMNH).

van VONDEL: Oriental Haliplidae

Haliplus chinensis Falkenstròm
Haliplus chinensis Falkenström, 1932: 191.

Fully redescribed by Vondel (1991). In the

Oriental region only known from China.

Haliplus diopus Guignot

(figs. 19-27)

Haliplus diopus Guignot, 1955a: 270. Holotype d,
‘Tonkin, Hoa-Binh [Vietnam], VII-1940 (Coom.)’ [A.
de Cooman] (MNHN) [examined].

Haliplus diopus, Guignot 1955b: 294.

Diagnosis

This species can be distinguished from H. pulchel-
lus by the wider distance between the eyes and from
H. indicus by the clearly separated pits on the metas-
ternal process.

Description

Length 2.8-3.1 mm, width 1.6-1.7 mm. Body
oval, widest in or just before the middle (fig. 19).

Head: Brown, sparsely and weakly punctured,
between eyes partly unpunctured. Distance between
eyes 1.1 X width of one eye. Antennae yellow to yel-
low-red (fig. 20), palpi yellow-red.

Pronotum: Yellow to yellow-red, longitudinal dark
blotch along the middle, this blotch can be vague or
reduced to a blotch on anterior part (fig. 19). Lateral
borders slightly convex, concave just behind anterior
corners, weak margins not allways clearly visible be-
cause of coarse punctures. Strongly, on the disc more
weakly punctured, widened punctures in rounded ap-
ical corners (fig. 19). Usually transverse impression
along base. Width at base 1.8-2.1 length in the
middle and 1.6-1.8 X width in front.

Elytra: Yellow with an extensive dark marking
along suture and along base to fifth puncture-row,
central mark connected to small mark in third inter-
val, a various number of irregular marks spread over
elytra (fig. 19). Rather strong primary puncture-rows,
26-30 punctures in first row. Secondary punctures
fairly strong, accompanied by some very small punc-
tures. All punctures darkened, but distinctly yellow in
centre of punctures (fig. 21). Margins of elytra slight-
ly serrate in basal part (fig. 19).

Ventral side: Yellow-red to rust-coloured, elytral
epipleura yellow-red, legs red-brown, darkened to-
wards coxae. Prosternal process strongly grooved
along margins, slightly narrowed near coxae, strongly

Figs. 28-36. Haliplus diruptus, holotype. — 28, dorsal view; 29, antenna; 30, punctures near elytral base and suture; 31, dor-
sal side of hind tibia; 32, prosternal process; 33, lateral view of prosternal process; 34, left paramere; 35, penis; 36, right par-

amere.

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punctured in both grooves, clearly margined anterior-
ly (fig. 23), lateral plicae complete (fig. 24). Metaster-
nal process flat with strong pit on both sides of the
middle, between pits not or hardly impressed, sparse-
ly, laterally densely punctured (fig. 23). Metacoxal
plates strongly, along suture weakly and sparsely punc-
tured, punctures without a hair, apical sutural corner
slightly margined. Fifth and sixth abdominal segment
with complete row of strong punctures. Setiferous stri-
ole on dorsal side of hind tibia very short, about 1/8 X
length of tibia, longest of two tibial spurs about
4/5 X length of first tarsal segment (fig. 22).

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on ventral side. Penis and
parameres as in figs. 25-27.

Biology and distribution (fig. 128)
Biology unknown.
Vietnam.

Material examined. — Vietnam: Holotype d, 2 allotype,
4 d paratypes, 69 paratypes, Tonkin, Hoa Binh, viii.1940
& viii.1941, A. de Cooman, Haliplus diopus n. sp. dr. F.
Guignot det. (ISNB, MNHN).

Haliplus diruptus Balfour-Browne
(figs. 28-36)

[Haliplus simplex sensu Régimbart 1899, nec Clark 1863.
Misidentification].
Haliplus diruptus Balfour-Browne, 1946: 436.

This species was fully redescribed by Vondel
(991095)

Probably also in India: see under unnamed material.
Guignot records this species from Burma (1954: 563),
but later (1956) changed his view and based the new
H. shanicus on that material. Additional material exam-
ined: 12, SW China, Yunnan, Western Hills nr.
Kunming, 2100 Mt., 7.viii.1940, J. L. Gressitt (BPBM).

Haliplus eximius Clark
(fig. 37-45)

Haliplus eximius Clark, 1863: 418.

Haliplus modestus Zimmermann, 1924: 139.
Haliplus hiogoensis Kano & Kamiya, 1931: 1.
Haliplus emmerichi Falkenstrôm, 1936: 79.

This species was fully redescribed by Vondel
(1991: 97), but erroneously treated under the name
H. eximis.

Additional

material examined: China: 2 ex.

VAN VONDEL: Oriental Haliplidae

Chusan Isl, J. J. Walker (BMNH); 1 ex., Fukien,
Yungan, 5.ix.1940, T. C. Man (BPBM).

Haliplus holmeni Vondel
Haliplus holmeni Vondel, 1991: 109.

This species is only known from China: Yunnan.

Haliplus indicus Régimbart stat. n.
(figs. 46-54)

Haliplus pulchellus var. indicus Régimbart, 1899: 189.
Lectotype d (here designated), ‘Chota-Nagpore [India,
Ranchi Distr., Bihar], Nowatoli, R. P. Cardon, viii-
ix.1896, pulchellus var. indicus Rég., Dr. Régimbart vidit
1898, Museum Paris, ex coll. R. Oberthur’ (MNHN) [ex-
amined].

Haliplus pulchellus var. indicus, Zimmermann 1920: 314.

Haliplus pulchellus ab. indicus, Zimmermann 1924: 140.

Haliplus pulchellus indicus, Vazirani 1966: 133.

Diagnosis

This species, belonging to the group with double
grooved prosternal process, can be distinguished from
H. pulchellus by the wider distance between the eyes.
It is here considered a distinct species.

Description

Length 2.8-3.1 mm, width 1.6-1.7 mm. Body
oval, widest in or just before the middle (fig. 46).

Head: Brown, sparsely and weakly punctured,
between eyes partly unpunctured. Distance between
eyes 1.1-1.2 X width of one eye. Antennae yellow to
yellow-red (fig. 47), palpi yellow-red.

Pronotum: Yellow to yellow-red, dark mark in an-
terior middle, vague central mark before base can be
reduced or absent (fig. 46). Lateral borders slightly
convex, concave just behind anterior corners, weak
margins usually not clearly visible because of coarse
punctures. Strongly punctured, on disc more weakly
so, with widened punctures in rounded posterior cor-
ners. Usually a transverse impression along base op-
posite fourth to sixth elytral puncture-row.

Elytra: Yellow with an extensive dark marking
along suture and along base to fourth or fifth punc-
ture-row, central mark connected with small mark in
fourth interval, a variable number of irregular marks
spread over elytra (fig. 46). Rather strong to strong
primary puncture-rows, 26-35 punctures in first row.
Secondary punctures fairly strong. All punctures
darkened, but distinctly yellow in centre of punctures
(fig. 48). Margins of elytra slightly serrate in basal
part (fig. 46).

Figs. 37-45. Haliplus eximius, China. — 37, dorsal view; 38, antenna; 39, punctures near elytral base and suture; 40, dorsal side
of hind tibia; 41, prosternal process; 42, lateral view of prosternal process; 43, left paramere; 44, penis; 45, right paramere.

299

‘TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Ventral side: Yellow-red to rust-coloured, elytral
epipleura yellow-red, legs red-brown, darkened to-
wards coxae. Prosternal process strongly grooved
along margins, slightly narrowed near coxae, strongly
punctured in both grooves, clearly margined along
anterior edge (fig. 50), lateral plicae complete (fig.
51). Metasternal process with strong transverse impres-
sion, in which two pits can be recognised, sparsely
punctured (fig. 50). Metacoxal plates strongly, along
suture weakly and sparsely punctured. Fifth and sixth
sternite with complete transverse row of strong punc-
tures, last sternite strongly and densely punctured.
Setiferous striole on dorsal side of hind tibia about
1/5 X length of tibia, longest of two tibial spurs about
4/5 X length of first tarsal segment (fig. 49).

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on, ventral side. Penis and
parameres as in figs. 52-54.

Biology and distribution (fig. 127)
Biology unknown.

India: Bihar, West Bengal, Tamil Nadu.

Material examined. — India: 4d, 29, Chota Nagpore
[Ranchi distr. Bihar], Nowatoli, R.P. Cardon, viii-ix.1896
(MNHN, RMNH); 5d, 49, W. Bengal, Santiniketan,
Birbhum District, x.1937, T. C. Maa; 19, Madras State,
Coimbatore, 420 m, iv.1962, P. S. Nathan (BPBM).

Haliplus javanicus sp. n.
(fig. 55-63)

Type material. — Holotype d, ‘Tjilatjap, Java, Drescher,
12.1915; coll. F. C. Drescher’ (ZMAN). — Paratypes, 2d:
1d, data as holotype (coll. Vondel); 16, ‘Java, 1915,
Weltevreden, Neytzell de Wilde; coll. Dr. D. MacGillavry
(ZMAN).

Diagnosis

This species can be distinguished from other spe-
cies with a double grooved prosternal process by the
combination of the central pronotal blotch, the
smooth elytral margin near shoulders and the elytral
basal darkening reaching sixth puncture-row.

Description

Length 2.8-3.0 mm, width 1.5-1.7 mm. Body
oval, widest in the middle (fig. 55).

Head: Brown to dark brown, moderately strongly,
on vertex stronger punctured. Distance between eyes
0.7-1.0 X width of one eye. Antennae yellow-red (fig.
56), palpi yellow-red.

VAN VONDEL: Oriental Haliplidae

Pronotum: Yellow to yellow-brown with large cen-
tral longitudinal mark. Lateral borders straight and
finely margined, near anterior corner concave and not
margined. In posterior corners strongly and densely
punctured, basally strongly punctured, on the disc
more weakly and sparsely punctured. Impressed op-
posite fourth to sixth elytral puncture-rows (fig. 55).

Elytra: Yellow to yellow-red, dark marks along su-
ture and along base to sixth puncture-row, various
marks on intervals (fig. 55). Primary puncture-rows
moderately strong, about 28 punctures in first row.
Secondary punctures strong. All punctures darkened
with yellow centre (fig. 57).

Ventral side: Body yellow-red to brown, elytral epi-
pleura yellow-red, legs yellow-red to brown-red.
Prosternal process with two strong grooves, margined
at anterior edge, narrowed near coxae, moderately
strongly and sparsely punctured (fig. 59), lateral pli-
cae complete (fig. 60). Metasternal process with two
strong pits, weakly punctured (fig. 59). Metacoxal
plates strongly punctured, along suture weakly punc-
tured. Setiferous striole on dorsal side of hind tibia
very short, consisting of about 4 small punctures,
longest of the two tibial spurs about 2/3 X length of
first tarsal segment (fig. 58).

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on ventral side. Penis and
parameres as in figs. 61-63.

Biology and distribution (fig. 130)
Biology unknown.
Indonesia: Java.

Haliplus kapuri Vazirani

(fig. 64-65)

Haliplus kapuri Vazirani, 1975: 317. Holotytpe d, India,
Maharashtra, Fritzgerald, Satara District, 5.x.1972, A. P.

Kapur (in Zoological Survey of India, Calcutta)[not ex-
amined]

Original description

Length 3.5 mm.

Head testaceus; punctation moderate and irregular,
stronger on the vertex; antennal segments 3 and 4
rather thinner than others, 7th segment slightly long-
er than 8th segment.

Pronotum testaceus; punctation larger than on the
head, larger and stronger anteriorly and posteriorly,
less strong on the disc, generally irregular, two longi-
tudinal rows of fine punctures discernible on the disc;

Figs. 46-54. Haliplus indicus, lectotype. — 46, dorsal view; 47, antenna; 48, punctures near elytral base and suture; 49, dorsal
side of hind tibia; 50, prosternal process; 51, lateral view of prosternal process; 52, left paramere; 53, penis; 54, right para-

mere.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 55-63. Haliplus javanicus, holotype. — 55, dorsal view; 56, antenna; 57, punctures near elytral base and suture; 58, dor-
sal side of hind tibia; 59, prosternal process; 60, lateral view of prosternal process; 61, left paramere; 62, penis; 63, right par-
amere.

302

65

VAN VONDEL: Oriental Haliplidae

66 67

Figs. 64-65. Haliplus kapuri, (after Vazirani, 1975). — 64, ventral view of thorax; 65, penis.

Figs. 66. Haliplus manipurensis, penis (after Vazirani 1966).
Figs. 67. Haliplus pruthii, penis (after Vazirani 1966).

lateral margin of the pronotum rebordered.

Elytra pale testaceus without any markings; punc-
tation quite regular except in the apical portion, both
strial and interstrial punctures black; row of sutural
punctures moderate, less strong but more dense than
the strial punctures, interstrial punctures less strong
than the strial punctures but not more numerous
than them.

Ventral side testaceus with legs ferruginous, meso-
sternum darker, metacoxal plates and epipleurae pal-
er. Prosternal process not rebordered laterally but
canaliculate, strongly punctate more so on the lateral
borders, a little narrowed in between the procoxae
and the mesocoxae (fig. 64), anteriorly bordered by a
fine ridge; anterior part of the metasternum (behind
the prosternal process) distinctly swollen/inflated,
with a large median pit behind it and two smaller pits
on the anterolateral corner; punctation on the meta-
sternum irregular except a transverse row of strong
punctures behind the large median pit and a row of
fine punctures along the posterior border of the met-
asternum; metacoxal plates moderately punctate,
punctures irregular and separated by 2-3 X its own
diameter. Penis (fig. 65) rather strongly curved,
rounded at the apex and narrowed in the middle.

Biology and distribution (fig. 129)
Biology unknown.
Only known from the type-locality.

Remarks. — As I have been unable to borrow the
type for examination I only know this species from
the original description. According to Vazirani it
comes close to H. arrowi.

Haliplus kotoshonis Kano & Kamiya
Haliplus kotoshonis Kano & Kamiya, 1931: 2.

This species is fully redescribed by Vondel (1991:
113).

Probably in Laos: see under unnamed material.
Additional notes: Satô (1984) recorded this species
from the Ryukyu Islands.

Additional material examined. — 19, Ryukyu Isl.,
Ishigaki I., 20-30.x1.1952, G.E. Bekart (BPBM).

Haliplus manipurensis Vazirani

(fig. 66)

Haliplus manipurensis Vazirani, 1966: 132. Holotype d,
[India], Manipur, Imphal, 880 m, 24°44’N, 93°58’E,
8.vii.1960, F. Schmid coll. (in Zoological Survey of

India) [not examined].

Original description

Length 3.2-3.5 mm. Form, oval, feebly attenuated
posteriorly.

Head testaceus, finely and moderately punctate,
punctures on the vertex separated by their own diam-

303

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

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Figs. 68-76. Haliplus nedungaduensis, holotype. — 68, dorsal view; 69, antenna; 70, punctures near elytral base and suture; 71,

dorsal side of hind tibia; 72, prosternal process; 73, lateral view of prosternal process; 74, left paramere; 75, penis; 76, right
paramere.

Figs. 77-86. Haliplus perroti, holotype. — 77, dorsal view; 78, antenna; 79, punctures near elytral base and suture; 80, dorsal

side of hind tibia; 81, prosternal process; 82, lateral view of prosternal process; 83, metacoxal plate; 84, left paramere; 85, pe-
nis; 86, right paramere.

304

VAN VONDEL: Oriental Haliplidae

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 87-97. Haliplus philippinus (87, 89-91, 93, 95-97, holotype; 88, 92, 94, Laos). — 87, dorsal view; 88, antenna; 89, punc-
tures near elytral base and suture; 90, dorsal side of hind tibia; 91, prosternal process; 92, mesepimeron; 93, lateral view of
prosternal process; 94, head and pronotum; 95, left paramere; 96, penis; 97, right paramere.

306

eter. Antennal segments 7-11 subequal and broader
than the sixth.

Pronotum testaceous, paler towards sides; puncta-
tion moderate and irregular, stronger along anterior
and posterior margins, sparse on the disc; a few punc-
tures on postero-lateral margins strong and confluent.
Anterior angles not projecting beyond the middle of
anterior margin. Sides moderately rebordered, slight-
ly convex, posterior angles slightly prominent, not
projecting beyond the thoraco-elytral angles.

Elytra testaceous with irregular ferruginous mark-
ings, more so along the suture and extending laterally
at base. Sutural line of punctation irregular, not so
closely punctate; punctures separated by twice their
own diameter. Strial punctures moderate, shallow,
black; on the disc quite close and not separated by
their own diameter. Each interstrial space with a row
of much smaller and deeper, but not more numerous
black punctures.

Ventral side testaceous, epipleurae paler. Prosternal

process and legs darker. Prosternal process margined

laterally but not anteriorly, canaliculate, slightly nar-
rowed adjacent to the front coxae; anterior end a little
wider than apex; lateral margins punctate but not
roughened, punctures fine, separated by more than
their own diameter. Apical part of metasternum al-
most flat, pitted, moderately but irregularly punctate.
Metacoxal plates with moderate, shallow and dense
punctures, punctures separated by half of their own
diameter.

Male with anterior tarsi moderately broadened and
furnished with ‘ciliae’ underneath; claws broken. Penis

very slightly curved, rounded at the apex (fig. 66).

Biology ne inbucon (fig. 129)
Biology unknown.

Only known from the type-locality.

Remarks. — As I have been unable to borrow the
type for examination I only know this species from
the original description.

Haliplus nedungaduensis sp. n.
(figs. 68-76)

Type material. — Holotype à , ‘South India, Nedungadu,
7.1934, Hathach [?]’ (MNHN).

Diagnosis

This species can be distinguished from the other
species with double grooved prosternal process by the
absence of elytral markings.

Description
Length 2.7 mm, width 1.5 mm. Body oval, widest
in the middle (fig. 68).

VAN VONDEL: Oriental Haliplidae

Head: Yellow-brown, vertex slightly darkened,
rather weakly punctured, unpunctured band between
the eyes not always present. Distance between the
eyes 1.2-1.4 the width of one eye. Antennae yel-
low-red (fig. 69), palpi yellow-red.

Pronotum: Yellow to yellow-red. Lateral borders
about straight, fine margins a little undulate. Modera-
tely strongly and sparsely punctured. Base opposite
fourth to sixth elytral puncture-row impressed and
strongly punctured (fig. 68).

Elytra: Yellow to yellow-red, suture slightly dark-
ened to secondary punctures (fig. 68). Completely
margined, margin serrate in anterior part. Primary
puncture-rows fairly weak, about 29 punctures in
first row. Secondary punctures strong. All punctures
darkened with a yellow centre (fig. 70).

Ventral side: Body yellow to yellow-red, elytral epi-
pleura yellow, legs yellow-red to brown-red, slightly
darkened towards coxae. Prosternal process with two
weak grooves, margined at anterior edge, narrowed
near coxae, anteriorly wider than posteriorly, moder-
ately strongly punctured (fig. 72), lateral plicae com-
plete (fig. 73). Metasternal process with deep trans-
verse impression in which two pits are recognisable,
weakly punctured (fig. 72). Metacoxal plates moder-
ately strongly punctured, along suture with a weakly
punctured area. Setiferous striole on dorsal side of
hind tibia about 1/3 X length of tibia, longest of the
two tibial spurs about 2/3 X length of first tarsal seg-
ment (fig. 71). |

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on ventral side. Penis and
parameres as in figs. 74-76.

Biology and distribution (fig. 127)
Biology unknown.
Only known from the type-locality.

Haliplus perroti Guignot
(figs. 77-86)

Haliplus perroti Guignot, 1950: DI Holotype 6, “Tam Dao,
Tonkin, H. Perrot, 6, Type’ (MNHN) [examined].

Diagnosis

This species can be recognised by the longitudinal
elytral marks, which are not connected to the darke-
ned suture.

Description

Length 3.2-3.4 mm, width 1.7-1.9 mm. Body
oblong oval, widest in the middle (fig. 77).

Head: Yellow-brown, vertex slightly darkened,
rather weakly punctured, unpunctured band between
eyes not always present. Distance between eyes 1.2-
1.4X width of one eye. Antennae yellow-red (fig.

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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

78), palpi yellow-red.

Pronotum: Yellow with two small marks on disc
before base. Lateral borders convex, finely margined.
In anterior corners strongly and densely punctured,
basally with a continous row of darkened punctures
getting strong towards sides, on disc more weakly and
sparsely punctured. Slightly impressed in front of
central base (fig. 77).

Elytra: Yellow to yellow-red, suture completely
darkened to secondary punctures, oblong marks on
even intervals (fig. 77). Primary puncture-rows fairly
weak, 31-35 punctures in first row. Secondary punc-
tures weak. All punctures darkened with yellow
centre (fig. 79).

Ventral side: Body yellow-red to yellow-brown,
elytral epipleura yellow, legs brown-red, darkened to-
wards coxae. Prosternal process impressed in the mid-
dle, less impressed at apex, margined at anterior edge,
narrowed near coxae, moderately strongly punctured
(fig. 81), lateral plicae complete (fig. 82). Metasternal
process widely and strongly impressed in the middle,
weakly punctured (fig. 81). Metacoxal plates densely
and rather strongly punctured, along suture with a
wide weakly punctured area, slightly impressed to-
wards apical point (fig. 83). Setiferous striole on dor-
sal side of hind tibia about 1/3 X length of tibia, con-
sisting of about 12 small groovelike connected
punctures, longest of the two tibial spurs about 2/3 X
length of first tarsal segment (fig. 80).

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on ventral side. Penis and
parameres as in figs. 84-86.

Biology and distribution (fig. 125)
Biology unknown.
Vietnam. India: West Bengal, Rajasthan.

Material examined. — Vietnam: Holotype d; 19, allo-
type, 2 d, paratype, Tam Dao, Tonkin, H. Perrot (MNHN).
— India: 1 ex., Pusa, Bengal, R. D. D., xi.1908 (BMNH); 19,
Rajasthan, Sariaka, 16.viii.1989, A. Riedel (Hendrich).

Haliplus philippinus Chapin
(figs. 87-97)
Haliplus philippinus Chapin, 1930: 9. Holotype d, ‘Bacoor

Nov PI, P. L. Stangl Collector, Type No. 41757
U.S.N.M., Haliplus philippinus Type Chap.’ (USNM)
[examined].

Diagnosis

This species can be distinguished from the related H.
pulchellus by the four punctures on the mesepimeron.

Description

Length 2.4-2.7 mm, width 1.3-1.5 mm. Body
broad, more or less parallel in basal half of elytra (fig.
87).

Head: Brown-red, margins dark, weakly and
sparsely punctured. Usually with dark arrowlike mark
(fig. 94). Distance between eyes about 1.2 X width of
one eye. Antennae yellow-red (fig. 88), palpi yellow-
red.

Pronotum: Yellow-red to yellow in lateral parts.
Usually wich dark central mark (fig. 94). Lateral bor-
ders slightly convex, not margined. Posterior corners
broadly rounded, anterior corners bent outwards.
Anterior margin clearly protruding in the middle.
Rather weakly punctured, two rows of strong punc-
tures along the slightly impressed base, coarse punc-
tures together forming a strong impression in each
hind corner (fig. 87). A

Elytra: Yellow-red, brown markings along base and
suture and a clear mark in posterior half between
fourth and sixth puncture-row, often additional
marks on intervals (fig. 87). Primary punctures mod-
erately strong, standing in irregular sometimes diffi- _
cult to recognise rows. About 28 punctures in first
row. Secondary punctures relatively strong, some-
times nearly as strong as primary punctures. All punc-
tures slightly darkened with a yellow centre (fig. 89).

Ventral side: Body brown-red with several strongly
darkened parts, elytral epipleura yellow-red, legs yel-
low-red to brown-red. Mesepimeron with about 4
strong punctures (fig. 92). Prosternal process parallel,
slightly narrowed near coxae, sparsely punctured,
margins ridgelike, about flat in the middle (fig. 91).
Metasternal process flat, small clear pit on each side,
sparsely punctured. Metacoxal plates strongly, near
suture very sparsely and weakly punctured, apical cor-
ner of suture sharp. Last sternite largely covered with
strong punctures, fifth and sixth sternite with a con-
tinuous row of strong punctures. Setiferous striole on
dorsal face of hind tibia about % X length of tibia,
longest of tibial spurs about 4 X length of first tarsal
segment (fig. 90).

Male: First three segments of front and midlegs
slightly widened, scaly hairs on ventral side. Claws of
forelegs equal in shape and length. Penis and para-
meres as in figs. 95-97.

Biology and distribution (fig. 128)
Attracted to light. Up to an altitude of 920 m.
Philippines. Indonesia: Java. Thailand. Laos.

Vietnam. Burma.

Figs. 98-106. Haliplus pulchellus, lectotype. — 98, dorsal view; 99, antenna; 100, punctures near elytral base and suture; 101,
dorsal side of hind tibia; 102, prosternal process; 103, lateral view of prosternal process; 104, left paramere; 105, penis; 106,

right paramere.

308

van VONDEL: Oriental Haliplidae

309

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Material examined. — Philippines: Holotype d (USNM). —
Vietnam: 19, Cochin China, Travinh, Travinh Prov,
vii.1932, K.M. Tsui (BPBM). — Laos: 2d, Vientiane, 31.v-
3.vi.1960, S. Quale & L. Quale, light trap; 18, Muong
Sing, NW of Luang Prabang, 6-10.vi.1960, L.W. Quale,
light trap, 650 m.; 2 ex, 39 km S. Djiring, 810 m,
29.iv.1969, J.W. Quale; 2 ex., Di Linh, Djiring, 920 m,
light trap, 22-28.iv.1960, S. & J. Quale; 1 ex., Cochin
China, French Indo China, Travinh, Travinh Province,
vii.1932, K. H. Tsui (BPBM). — Thailand: 2d, 19, Bangkok
(BMNH, Vondel); 2 ex., Bangkok, Aynttaya, 28.viii.1988, S.
Schödl (NHMV, Vondel); 2 ex., Bangkok (MNHN); 2 ex., 25
km NW Lan Sak, 210 km NW Bangkok, viii.1991, W.
Thielen (RMNH). — Burma: 1 2, Victoria Point, Tenasserim,
viii. 1887, E.T. Atkinson (BMNH); 8 ex., Rangun, 16.v.1976,
Wewalka (Mazzoldi). — Indonesia: 16, Batavia [Jakarta],
ii.1949, C. v. Nidek (ZMAN).

Haliplus pruthii Vazirani
(fig. 67)

Haliplus pruthii Vazirani, 1966: 130. Holotype d, Madhya
Pradesh (Narmada Survey); Rewa State (=Rewa District),
Khetagao, stn. 45, c 880 m, 24°31N, 81°19E;
Mar.1927, H.S. Prithi coll. (in: Zoological Survey of
India) [not examined].

Diagnosis

The character of the two longitudinal marks on
pronotum seems to offer a possibility to distinguish
this species from related species.

Original description
Length 3.0-3.2 mm.

Head testaceus; punctation on the vertex fine,

moderately close; antennal segments 7-11 subequal,
segment 7 not significantly broader than preceding
segment.

Pronotum testaceus, with two ferruginous longitu-
dinal parallel lines on the disc coinciding with punc-
tures; punctation irregular, moderate, rather effaced
on the disc (excepting those coinciding with the fer-
ruginous markings); sides narrowly rebordered, a lit-
tle oblique, very slightly convex; posterior angles ob-
tuse. Anterior angles almost in line with middle of the
anterior margin.

Elytra testaceus, with irregular and indeterminate
ferruginous markings, more prominent along the su-
ture and at base. Sutural line of punctures regular,
punctures almost separated by their own diameter,
more numerous than the strial punctures, Strial punc-
tures moderate, rather close and shallow, narrowly
black. Each interstrial space with a row of small deep
punctures not more than the strial punctures.

Ventral side testaceus; sternum and legs ferrugi-
nous. Prosternal process moderately margined lateral-
ly, narrowly margined anteriorly, moderately nar-
rowed adjacent to the front coxae, anterior end and
apical end subequal, lateral margins punctate, median

310

area hardly inflated, punctation indistinct. Apical
part of metasternum slightly inflated and pitted.
Metasternum weakly and sparsely punctate.
Abdominal sternites with a single transverse row of
punctures. Apical abdominal sternite with a few pos-
tero-lateral punctures (less than in angustifrons or ar-
rowi). Metacoxal plates more or less regular and fine-
ly punctate, separated by two or three times their own
diameter.

Male with protarsi moderately dilated and fur-
nished with cilia underneath. Protarsal claws thin,
slightly curved and more than half the apical seg-
ments length. Penis (fig. 67) of paratype moderately
curved.

Biology and distribution (fig. 129)

Up to an altitude of 880 m. Probably attracted to
light.

India: Madhya Pradesh, probably also in Uttar

Pradesh and Bihar (see under unnamed material).

Remarks. — As I have been unable to borrow the
type for examination I only know this species from
the original description.

Haliplus pulchellus Clark
(fig. 98-106)

Haliplus pulchellus Clark, 1863: 418. Lectotype d (here des-
ignated), ‘Browning, Siam [Thailand], Co-type’ (BMNH)
[examined].

Haliplus oceanicus Régimbart, 1886: 139. Lectotype (here
designated), ‘Ed. Everts, W. Dates, Medan, Sumatra;
Haliplus oceanicus Régb., type; Type; Museum Leiden,
Haliplus pulchellus det. Clark (RMNH) [examined].

Haliplus pulchellus, Régimbart 1889: 147, 1892: 112, 1899:
188, Zimmerman 1920: 314, 1924: 140, 1927: 2,
Guignot 1955b: 295, Vazirani 1966: 133.

Haliplus oceanicus, Zimmermann 1927: 3.

Haliplus pulchellus var. oceanicus, Régimbart 1899: 189,
Zimmermann 1920: 314, 1924: 140.

Diagnosis

This species can be distinguished from the other
species with a double grooved prosternal process by
the characters mentioned in the key.

Description

Length 2.5-3.4 mm, width 1.5-2.0 mm. Body
oval, widest just before the middle (fig. 98).

Head: Brown to dark brown, darkened along the
eyes and near antennae, sometimes with-a weak ar-
row-like darkening in the middle. Sparsely and weak-
ly, behind eyes stronger punctured. Distance between
eyes 0.9-1.1 X width of one eye. Antennae yellow to
brown-yellow (fig. 99), palpi yellow-red to brown.

Pronotum: Yellow to yellow-red, large medial dark
blotch that can be reduced to a round blotch on ante-

VAN VONDEL: Oriental Haliplidae

Figs. 107-112. Haliplus samosirensis, holotype. — 107, dorsal view; 108, antenna; 109, punctures near elytral base and suture;
110, dorsal side of hind tibia; 111, prosternal process; 112, lateral view of prosternal process.

rior half with a partly reduced or vague blotch on ap-
ical half (fig. 98). Lateral borders straight to slightly
convex, anterior corners concave, clearly, vaguely or
not visibly margined, margin never reaching anterior
corner. Apical corners rounded, middle of base usual-
ly finely impressed. Strongly, anteriorly weaker punc-
tured, near basal corners with one or two stronger
punctures in a slight depression. Basal punctures
darkened.

Elytra: Yellow to yellow-red, clear and extensive
markings along suture and along base untill fifth pri-
mary puncture-row, a central mark in third interval
connected to suture and various small marks in later-
al part (fig. 98). Primary punctures of average
strength, stronger towards base, about 30 punctures

in first row. Secondary punctures relatively strong ac-
companied by some very small punctures. All punc-
tures darkened except the central hole. Lateral bor-
ders sometimes slightly serrate in basal part.

Ventral side: Body yellow-brown to dark brown,
elytral epipleura yellow to yellow-brown, legs yellow-
brown with brown to dark-brown femora and coxae.
Prosternal process with two strong grooves, anterior
edge margined, punctured, sometimes strongly, espe-
cially in the grooves, weakly narrowed near coxae,
sligthly diverging towards apex (fig. 102, 103).
Metasternal process flat with two pits, these pits
sometimes very strong and than almost forming one
confluent transverse pit, sparsely punctured (fig.
102). Coxal plates rather sparsely, near suture weakly

a

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Figs. 113-121. Haliplus srilankanus, holotype. — 113, dorsal view; 114, antenna; 115, punctures near elytral base and suture;

116, dorsal side of hind tibia; 117, prosternal process; 118, lateral view of prosternal process; 119, left paramere; 120, penis;
121, right paramere.

52

punctured, sutural corners more or less rectangular.
Sternites with complete puncture-rows. Last sternite
almost completely covered with strong punctures, ap-
ical point with a very short ridge. Setiferous striole on
dorsal side of hind tibia very short, very close to the
lateral row of setae and often hard to distinguish from
it, longest of the tibial spurs about 4 X length of first
tarsal segment (fig. 101).

Male: First three tarsal segments of fore- and mid-
legs slightly widened, scaly hairs on ventral side. Penis
and parameres as in figs. 104-106.

Biology and distribution (fig. 126)

Collected up to an altitude of 920 m. Attracted to
light.

China along the coast north to Xiamen, Thailand,
Vietnam, Indonesia: Sumatra, Java, Sulawesi, India:

West Bengal, Sri Lanka.

Material examined. — China: 1 d, 12, Amoy [Xiamen],
Galatea (ZMUC). — Thailand: 16, Lectotype; 19,
Paralectotype, Browning, Siam; 1d, Siam; 2 ex., Bangkok,
Castlenau (BMNH); 16, Cotype, Wallace, Mal.’ [Malacca],
67-56, Paralectotype (BMNH); 1 ex., Pattaya, 4.x11.1979
(MZLU); 16, Banna, Chawang nr. Nabon, 70 m., 5.ix.1958,
J.L. Gressitt, light trap (BPBM); 6 ex., 25 km NW Lan Sak,
210 km NW Bangkok, viii.1991, W. Thielen (RMNH). —
Vietnam: 1 2, Annam, Chuan-an [nr. Hue], Museum Paris,
Coll. Maurice Régimbart, 1908, TYPE, oceanicus Rég. [not
belonging to syntype-series]; 12, Annam, Chuan-an [nr.
Hue]; 16, Cochin Chine, Harmand 1872; 1 m, Annam,
Rég. de An-Ninh [?], R. P. M. Maunier 1903 (MNHN). — Sri
Lanka: 16, Polonnaruwa, 28.xi.1980, M. Jäch (NHMV);
16, Maha-illerpulma [Madakalapuwa ?], 30.xi.1957, C. H.
Fernando, light (BMNH). — India: 19, Calcutta, Atkinson
coll. (BMNH). — Indonesia: Sumatra: 16, Lectotype of H.
oceanicus (RMNH); 6 ex., Fort de Kock [Bukittinggi], 920 m,
E. Jacobson, 1.1922, 1924, 1925, 1926 (BMNH, RMNH,
ZMAN); 1 ex., Solok, exped. Kleiweg de Zwaan 1907; 1 ex.,
Deli, de Bussy (ZMAN); 8 ex., Medan, vii. 1904, 1906 (-
RMNH); Java: 22 ex., Weltevreden [nr. Jakarta], 1915,
Neytzell de Wilde (RMNH, ZMAN); 4 ex., G. Oengaran [G.
Ungaran], v.1915, xii.1907, F. C. Drescher; 39 ex, Batavia
[Jakarta], C. v. Nidek, 25.ix.1947, xii.1948, 11.1949,
v.1949, vi.1949, xii.1949, 1.1950, v.1950; 2 ex., Tengger
Mountains, Drescher; 1 ex., Tjilatjap, xii.1914, F. C.
Drescher (ZMAN); 1 ex., Batavia [Jakarta], v.1913, J. B.
Corporaal; 1 ex., Weltevreden [nr. Jakarta], 1.1919, P. B.
Buytendijk (RMNH); Sulawesi: 1d, Makassar, I-74 [1.1874
?] O. Beccari (Vondel); 3 ex., Makassar, L. de Vos (RMNH);
19, Sharp (MNHN).

Haliplus samosirensis sp. n.

(fig. 107-112)

Type material. — Holotype 9, ‘Sumatra [Indonesia],
Samosir Isl., 2°28’N 98°49’E, Tuk-Tuk, L. Toba, 2800 Ft.
4.11.1978, M. Holmen; 13.’ (zMuc). — Paratypes, 22: 19,
same data as holotype (coll. Vondel); 12, ‘J. Corporaal,
Sumatra’s O.K., Medan, 1.10.1921, 20 M, Haliplus pul-
chellus v. oceanicus det. A. Zimmermann (ZMAN).

VAN VONDEL: Oriental Haliplidae

Diagnosis
This species can be distinguished from related spe-
cies by the elytra, which are provided with a basal dark

band reaching margins and very irregular dark marks.

Description

Length 3.3-3.5 mm, width 1.9-2.0 mm. Body
oval, widest in the middle (fig. 107).

Head: Red-brown, frons vaguely marked, moder-
ately strongly, on vertex stronger punctured. Distance
between eyes about 1.1 X width of one eye. Antennae
yellow-red (fig. 108), palpi yellow-red.

Pronotum: Yellow-brown, gradually darkening to-
wards the middle and base. Lateral borders convex,
finely margined, anterior corner not margined and
bending outwards. Strongly but not very densely
punctured, all punctures darkened. Impressed oppo-
site fourth to sixth elytral puncture-row (fig. 107).

Elytra: Yellow to yellow-red, suture weakly dark-
ened to first secondary punctures, base darkened to
seventh row, darker towards shoulders, various dark
marks (fig. 107). Completely margined, margin
sometimes invisible from above. Primary puncture-
rows strong and somewhat irregular, 22-26 punctures
in first row. Secondary punctures strong, sometimes
almost as strong as adjacent primary punctures. All
punctures darkened with a yellow centre (fig. 109).

Ventral side: Body yellow-red to brown, elytral epi-
pleura yellow to yellow-brown, legs brown-red, dark-
ened towards coxae. Prosternal process with two
grooves, margined at anterior edge, narrowed near
coxae, weakly punctured (fig. 111), lateral plicae
complete (fig. 112). Metasternal process with trans-
verse impression, in which two pits are recognisable,
weakly punctured (fig. 111). Metacoxal plates dense-
ly and rather strongly punctured, along suture with a
weakly punctured area. Setiferous striole on dorsal
side of hind tibia very short, longest of the two tibial
spurs about % X length of first tarsal segment (fig.
110). Sternites strongly punctured.

Male: Unknown.

Biology and distribution (Be. 130)
Up to an altitude of 850 m.
Indonesia: Sumatra.

Haliplus sharpi Wehncke

Haliplus sharpi Wehncke, 1880
Haliplus tsukushiensis Yoshimura, 1932

This species is fully redescribed by Vondel (1991:

129).
Additional notes: also recorded from Taiwan.
Additional material examined: 16, Taiwan,

Taihoku, 4.vi.1928, F. C. Hadden (BPBM).

515

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

Fig. 122. Oriental region as treated in this revision. Fig. 124. Distribution of Haliplus arrowi (dots: examined,
Fig. 123. Distribution of Haliplus angustifrons (dots: exam circles: literature record). Fig. 125. Distribution of Haliplus
ined, circles: literature record). perroti (dots) and Haliplus srilankanus (triangles).

Fig. 126. Distribution of Haliplus pulchellus (dots: exam-
ined, circles: literature records).

Fig. 127. Distribution of Haliplus indicus (dot) and Haliplus
nedungaduensis (triangle).

Fig. 128. Distribution of Haliplus philippinus (dots) and
Haliplus diopus (square).

Fig. 129. Distribution of Haliplus pruthii (dots: examined,
circle: literature record), Haliplus kapuri (triangle) and
Haliplus manipurensis (square).

Fig. 130. Distribution of Haliplus samosirensis (dot) and
Haliplus javanicus (triangles).

314

Haliplus srilankanus sp. n.
(figs. 113-121)

Type material. — Holotype à, ‘Sri Lanka, Tissawewa,
10.xii.1979, V. Mahler Jensen leg. (ZMuc). Paratype 6,
same data as holotype (Vondel); Paratype 2, ‘Sri Lanka,
Tissawewa, 10.xii.1979, V. Mahler Jensen leg., Haliplus
(Liaphlus) pulchellus Clark det. T. G. Vazirani 1981, SLO’
(ZMUC). Paratype, 2d, ‘India, Rajasthan, Bharatpur, 11.
viii.1989, A. Riedel’ (Hendrich, Vondel).

Diagnosis

This species is related to A. angustifrons and H. ar-
rowi, but can be distinguished from them by the char-
acters given in the key.

Description

Length 3.2 mm, width 1.8 mm. Body oval, widest
in the middle (fig. 113).

Head: Red-brown, moderately strongly punctured.
Distance between eyes about 1.1 X width of one eye.
Antennae yellow-red (fig. 113), palpi yellow-red.

Pronotum: Yellow-red, slightly and vaguely dark-
ened in the middle near anterior side and before base.
Lateral borders straight, finely margined, posterior
corner protruding (fig. 113). In posterior corners
strongly and densely punctured, impressed along
base. Strongly and except on disc densely punctured,
punctures not darkened except in the darkened cen-
tral areas near base and anterior edge.

Elytra: Yellow, suture completely darkened to first
primary punctures, base darkened to fifth row,
hooked marks on disc and in apical part connected to
suture, marks in sixth interval (fig. 113). Completely
margined. Primary puncture-rows moderately strong
and not very dense, stronger in basal part, about 30
punctures in first row. Secondary punctures moder-
ately strong, sparse or partly absent in even intervals.
All punctures darkened with a yellow centre (fig.
11159).

Ventral side: Body yellow-red to red-brown, elytral
epipleura yellow, legs yellow-brown, darkened to-
wards coxae. Prosternal process impressed in the mid-
dle, margined at anterior edge, slightly narrowed near
coxae, strongly and coarsely punctured (fig. 117), lat-
eral plicae complete (fig. 118). Metasternal process
widely and strongly impressed in the middle, weakly
punctured (fig. 117). Metacoxal plates moderately
strongly punctured, along suture more weakly punc-
tured. Setiferous striole on dorsal side of hind tibia
about 4 X length of tibia, consisting of about 12
small punctures, longest of two tibial spurs about 7

X length of first tarsal segment (fig. 116). Sternites
weakly punctured.

Male: First three tarsal segments of fore- and mid-
legs widened, scaly hairs on ventral side. Penis and
parameres as in figs. 119-122.

VAN VONDEL: Oriental Haliplidae

Biology and distribution (fig. 125)
Biology unknown.
Sri Lanka. India: Rajasthan.

Unnamed material

The specimens I failed to determine are listed be-
low. Some of them might belong to still undescribed
species.

Species near H. diruptus. — 16, [India], Calcutta,
Type, Atkinson Coll., BM 92-3, Haliplus falsificus n.
sp. [MS-name] Type! J. Balfour-Browne det. (BMNH)

Species near H. kotoshonis. — 19, Laos, Vientiane,
9.v.1965, P.D. Ashlock, at light (BPBM)

Species close to H. indicus, but with unmaculated
pronotum. — 1®, Sarda, Bengal, F. W. C. (BMNH)

Specimens probably representing H. pruthii.—1°,
[India], Uttar Pradesh, Haldwani Divn., Kumaun,
vii.1923, H. G. C., at light, 3956 (BMNH); 19,
(India, Bihar], Chota-Nagpore, Nowatoli, R. P.
Cardon, viii-ix.1896 (MNHN); 19, N. India, U.P.,
Rishikesh, 450 m, viii.1988, Werner (NHMV).

ACKNOWLEDGEMENTS

I wish to express my sincere thanks to the following
persons for placing material or information at my dis-
posal: R. Angus (Egham), L. Baert (Brussels, ISNB),
M. J. D. Brendell (London, BMNH), B. Brugge
(Amsterdam, ZMAN), S. P. Cover (Cambridge,
MCZC), R. Danielsson (Lund, MZLU), K. Desender
(Brussels, ISNB), J. P. Duffels (Amsterdam, ZMAN),
M. Hansen (Copenhagen, ZMUC), L. Hendrich
(Berlin), S. J.Hine (London, BMNH), M. Holmen
(Copenhagen, ZMUC), J. Huijbregts (Leiden, RMNH),
M. Jäch (Vienna, NHMV), J. Krikken (Leiden, RMNH),
P. Lindskog (Stockholm, NHRS), E. Matsui (Hondo
City), P. Mazzoldi (Brescia), H. Perrin (Paris,
MNHN), G. A. Samuelson (Honolulu, BPBM), G.
Scherer (Munich, ZSMC), A. Smetana (Ottawa, CNCI),
P. J. Spangler (Washington DC, USNM). The Uytten-
boogaart-Eliasen Foundation is acknowledged for fi-
nancial support.

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of Haliplus subgenus Liaphlus Guignot (Coleoptera:
Haliplidae). -Tijdschrift voor Entomologie 134: 75-144,

figs 1-312.

Vondel, B. J. van, 1992. Revision of the Palaearctic and
Oriental species of Peltodytes Régimbart (Coleoptera:
Haliplidae). — Tijdschrift voor Entomologie 135: 275-
267, figs 1-101.

Wehncke, E., 1880. Neue Haliplus. — Stettiner Entomolo-
gische Zeitung 41: 72-75.

Yoshimura, S., 1932. On a new species of Haliplus from
Japan. — Mushi 5: 102-103.

Zimmermann, A., 1920. Dytiscidae, Haliplidae, Hygrobii-
dae, Amphizoidae. — Coleopterorum Catalogus 71: 1-
325.

Zimmermann, A., 1924. Die Halipliden der Welt. —
Entomologische Blatter (fiir Biologie und Systematik der
Kafer) 20 (1): 1-16; (2): 65-80; (3): 129-144; (4): 193-
213.

Zimmermann, A., 1927. Fauna sumatrensis. Beitrag Nr. 45.
Revision der Haliplidae et Dytiscidae von Sumatra. —
Supplementa Entomologica 16: 1-44.

Received: 3 March 1993
Accepted: 4 May 1993

Volume 136 1775

Tijdschrift
voor |
Entomologie

A journal of systematic and evolutionary
entomology since 1858

Published by the Nederlandse Entomologische Vereniging

Tijdschrift voor Entomologie

A journal of systematic and evolutionary entomology since 1858

Scope

The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long
tradition in the publication of original papers on insect taxonomy and systematics.
The editors particularly invite papers on the insect fauna of the Palaearctic and
Indo-Australian regions, especially those including evolutionary aspects e.g.
phylogeny and biogeography, or ethology and ecology as far as meaningful for
insect taxonomy. Authors wishing to submit papers on disciplines related to
taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied
entomology, are requested to contact the editorial board before submitting.
Usually, such papers will only be published when space allows.

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

Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)

Advisory board
M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff).

The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
Amsterdam.

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c/o National Museum of Natural History,
Postbus 9517, 2300 RA Leiden, The Netherlands.

Correspondence regarding membership of the society, subscriptions and
possibilities for exchange of this journal should be addressed to:

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Special rate for members of the society. Please enquire.

Instructions to authors
Published with index of volume 136 (1993).

Graphic design
Ontwerpers B.V., Aad Derwort, 's-Gravenhage

Tijdschrift voor Entomologie

Contents of volume 136

Articles

113

125

133

137

147

235

23

55

257

Andersen, T.: see Saether

Beccaloni, G. W.
A new species of Extatosoma Gray (Phasmatodea: Phasmatidae) from Papua
New Guinea, with remarks on the species in the genus.

Chen, P. P.: see Nieser

Cuppen, J. G. M.
Distribution and ecology of Hydraena in The Netherlands (Coleoptera:
Hydraenidae).

Donnelly, T. W.
Two new genera of isostictid damselflies from New Britain, Bougainville, and
the Solomon Islands (Odonata: Zygoptera).

Haitlinger, R.
A new species of the genus Diplothrombium Berlese (Acari, Prostigmata,
Johnstonianidae) from Poland, based on the larva.

Hämäläinen, M.
Description of Neurobasis daviesi sp.n. from Palawan, with taxonomic notes
on other species of the N. chinensis group (Odonata, Calopterygidae).

Häuser, C. L.
An annotated checklist of the species of the Parnassiinae (Lepidoptera:
Papilionidae).

Hurkmans, W.
A monograph of Merodon (Diptera: Syrphidae). Part |.

Jong, H. de
The phylogeny of the Nephrotoma flavescens species group (Diptera:
Tipulidae).

Lansbury, I.
Strongylovelia (Veliidae) and Metrobatopsis (Gerridae) and associated pleus-
ton Hemiptera of West New Britain.

Lansbury, I.
Rhagovelia of Papua New Guinea, Solomon Islands and Australia (Hemiptera -
Veliidae).

Mathis, W. N. & T. Zatwarnicki
Revision of the shore-fly genus Chlorichaeta Becker (Diptera: Ephydridae).

Neboiss, A.

New species of the genus Molanna Curtis from Sulawesi (Trichoptera:
Molannidae).

259 Nieser, N. & P. P. Chen
The Rhagovelia (Heteroptera: Veliidae) of Sulawesi (Indonesia). Notes on
Malesian aquatic and semiaquatic bugs (Heteroptera), IV.

77 Pape, T. & H. Shima
A new genus of Tachinidae from the Phillipines (Diptera).

83 Roth, L. M.
Revision of the cockroach genus Ctenoneura Hanitsch (Blattaria,
Polyphagidae).

283 Sæther, O. A. & T. Andersen
Lobosmittia, a new genus of orthoclads from Tanzania and Turkey (Diptera:
Chironomidae). -

Shima, H.: see Pape

289 Vondel, B. J. van
Revision of the Liaphlus species of the Oriental region excluding China
(Coleoptera: Haliplidae).

Zatwarnicki, T.: see Matthis

Announcements and book reviews

14 Ossiannilson, F., 1992. The Psylloidea (Homoptera) of Fennoscandia and
Denmark. [J. van Tol].

82 Scoble, M. J., 1992. The Lepidoptera: Form, function and diversity. [E. J. van
Nieukerken]. | Fibiger, M., 1993. Noctuidae Europaeae. Volume 2, Noctuinae
Il. [E. J. van Nieukerken].

Ill Publications Netherlands Entomological Society.

112 Announcement: 5th European Congress of Entomology, University of York,
U. K.

124 Announcement: XX International Congress of Entomology.

282 Kuchlein, J. H. (with collaboration of J. H. Donner), 1993. De Kleine Vlinders.
Handboek voor de Faunistiek van de Nederlandse Microlepidoptera. [E. J. van
Nieukerken].

288 Announcement: XX International Congress of Entomology, intent form.

Referees for volume 136

R. A. Angus (London), P. Ashe (Dublin), D. A. L. Davies, P. L. T. Beuk (Leiden), P. E.
Bragg (Nottingham), P. Goeldlin (Lausanne), R. W. Holzenthal (St Paul), D. K.
McAlpine (Sydney), N. Méller Andersen (Copenhagen), R. V. Southcott (Adelaide),
M. C. D. Speight (Dublin), I. Tangelder (Leiden), J. A. L. Watson (Canberra)

Dates of Publication
Volume 136 (1), pages 1-112, | July 1993
Volume 136 (2), pages 113-316, 10 December 1993

© Nederlandse Entomologische Vereniging, Amsterdam _IssN 0040-7496

NEW TAXA DESCRIBED IN TIJDSCHRIFT VOOR

ENTOMOLOGIE, VOLUME 136

ACARI

Diplothrombium ludwinae Haitlinger .................... 12
BLATTARIA

Ctenoneura crassistyla Roth ......... 107
Ctenoneura gigantea ROLE 0014 RE AU 87
Ctenoneura kinabaluana Roth … … nn. 104
Ctenoneura luma Roth... 101
Ctenoneura murudensis Roth................................ 106
Ctenoneura parascutica Roth … … … … 103
@ienoneurayparınga Roth. 2.7. 100
Ctenoneura propannulicornis Rother wert settee ae 88
Ctenoneura scutica Roth... 102
Cicnoncuratspinasiyla Roth: ne inn 98
Ctenoneura triprocessa Roth … … … … … n 106
Ctenoneura uncata Roth ........................................ 99
COLEOPTERA

Haliplus javanicus van Vondel … … 301
Haliplus nedungaduensis van Vondel..................... 307
Haliplus samosirensis van Vondel … … 313
Haliplus srilankanus van Vondel … … 315
DIPTERA

vaponcormyia Pape 82 Shninmas seated senaat: 77
Aporeomyia antennalis Pape & Shima .................... VAI
Azpeytia shirakii Hurkmans............................... 208

nom. nov. for Merodon scutellaris Shiraki, 1968)
Chlorichaeta africana Mathis & Zatwarnicki ......... 62

Chlorichaeta mais Mathis & Zatwarnicki ............... 66
Chlorichaeta orba Mathis & Zatwarnicki ............... 70
Lobosmittia Sæther & Andersen... 283
Lobosmittia basilobata Sæther & Andersen........... 284
Merodon aberrans isperensis Hurkmans ................ 177
Merodon altinosus Hurkmans............................... 164
Merodon ankylogaster Hurkmans ........................ 169
Merodon aureotibia Hurkmans … … 203
Merodon auronitens Hurkmans............................ 170
Merodon bequaerti Hurkmans … ne 194
Merodon cupreus Hurkmans …............................. 179
Merodon elegans Hurkmans ...................... 195
Merodon hypochrysos Hurkmans........................... 165
Merodon isperensis Hurkmans (ssp. of aberrans) … 177
Merodon kaloceros Hurkmans .......... 187
Wierodon lucas: AUTRMANS 2... 198
Merodon lusitanicus Hurkmans … … 181
Merodon marginicornis Hurkmans............. 166
Merodon mariae Hurkmans … … un 160
Merodon nitidifrons Hurkmans.........................- 199
Merodon oidipous Hurkmans … … … iii 171

Merodon ottomanus Hurkmans … … 161
Merodon persicus Hurkmans ............ n 171
Merodon satdagensis Hurkmans............................ 200
Merodon schachti Hurkmans … … … 201
Merodon sophron Hurkmans … annae 168
Merodon splendens Hurkmans....................... 182
Merodon tangerensis Hurkmans............................ 172
Merodon taniniensis Hurkmans ........................... 201
Merodon testaceoides Hurkmans … … … … … …… … 162
Merodon toscanus Hurkmans … … … 202
Merodon vandergooti Hurkmans … … 188
Merodon warnckei Hurkmans … … i 184
Merodon xanthipous Hurkmans … nennen 175
HEMIPTERA

Rhagovelia amnicus Lansbury … … nennen eneen 31
Rhagovelia aureospicata Lansbury........................... 41
Rhagovelia blagiokommena Nieser & Chen … … 268
Rhagovelia browni Lansbuty … nennen 33
Rhagovelia caesius Lansbury ........................2..........I0
Rhagovelia chrysomalla Nieser & Chen … 268
Rhagoveliatenmtallansbury an. 46
Rhagovelia daktylophora Nieser & Chen .............. 270
Rhagovelia fulvus Lansbury............22e22220222.20220202200e 36
Rhagovelia herzogensis Lansbury........................ 43
Rhagovelia hirsuta Lansbury......... i 39
Rhagovelia horaia Nieser & Chen ....................... 271
Rhagovelia kalami Nieser & Chen ....................... 273
Rhagovelia kastanoparuphe Nieser & Chen ......... 274
Rhasovelapriordbansbun sees eee eee 29)
Rhagovelia pseudocelebensis Nieser & Chen … … 275
Rhagovelia robina Nieser & Chen........................ 275
Rhagovelia thysanotos Lansbuty … nennen 48
Rhagovelia trichota Nieser & Chen....................... 277
Rhagovelia tropidata Nieser & Chen.................... 278
Strongylovelia priori Lansbuty ............. 15
ODONATA

@OnenvsticaDonnell ge CREO 128
Cnemisticta angustilobata Donnelly...................... 128
Cnemisticta latilobata Donnelly … … 129
Neurobasis daviesi Hämäläinen … 135
Ian: Donne 126
Titanosticta macrogaster Donnelly … … 126
PHASMATODEA

Extatosoma carlbergi Beccaloni … … … 115
TRICHOPTERA

Molanna jolandae Neboiss.................. 257

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993

INSTRUCTIONS TO AUTHORS

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Tijdschrift voor Entomologie
Volume 136, no. 2

Articles

113 G. W. Beccaloni
A new species of Extatosoma Gray (Phasmatodea: Phasmatidae) from Papua
New Guinea, with remarks on the species in the genus.

125 T. W. Donnelly
Two new genera of isostictid damselflies from New Britain, Bougainville, and
the Solomon Islands (Odonata: Zygoptera).

133 M. Hämäläinen
Description of Neurobasis daviesi sp.n. from Palawan, with taxonomic notes on
other species of the N. chinensis group (Odonata, Calopterygidae).

137 C. L. Häuser
An annotated checklist of the species of the Parnassiinae (Lepidoptera:
Papilionidae).

147 W. Hurkmans
A monograph of Merodon (Diptera: Syrphidae). Part |.

235 H. de Jong
The phylogeny of the Nephrotoma flavescens species group (Diptera: Tipulidae).

257 A. Neboiss
New species of the genus Molanna Curtis from Sulawesi (Trichoptera:
Molannidae).

259 N. Nieser & P. P. Chen
The Rhagovelia (Heteroptera: Veliidae) of Sulawesi (Indonesia). Notes on
Malesian aquatic and semiaquatic bugs (Heteroptera), IV.

283 O. A. Sæther & T. Andersen
Lobosmittia, a new genus of orthoclads from Tanzania and Turkey (Diptera:
Chironomidae).

289 B. J. van Vondel
Revision of the Liaphlus species of the Oriental region excluding China
(Coleoptera: Haliplidae).

Announcements and book reviews

124 Announcement: XX International Congress of Entomology.

282 Kuchlein, J. H. (with collaboration of J. H. Donner), 1993. De Kleine Vlinders.
Handboek voor de Faunistiek van de Nederlandse Microlepidoptera. —
PUDOC, Wageningen. [E. J. van Nieukerken].

288 Announcement: XX International Congress of Entomology, intent form.

© Nederlandse Entomologische Vereniging, Amsterdam

Published 10 December 1993 ISSN 0040-7496

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