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Full text of "Revue suisse de zoologie"

ANNALES 



de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

et du 

MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



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fascicule 4 
2005 






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REVUE SUISSE DE ZOOLOGIE 



TOME 112— FASCICULE 4 

Publication subventionnée par: 

Académie suisse des Sciences naturelles (SCNAT) 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 

Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, etnologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 

SUISSE Fr. 225.— UNION POSTALE Fr. 250.- 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P 6434, CH-1211 Genève 6, Suisse 



ANNALES 

de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 

et du 

MUSÉUM D'HISTOIRE NATURELLE 

de la Ville de Genève 



tome 1 1 2 
fascicule 4 
2005 




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JëL GENEVE DECEMBRE 2005 ISSN 0035 -418 X </5 ■■■■ 



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REVUE SUISSE DE ZOOLOGIE 



TOME 112-FASCICULE 4 

Publication subventionnée par: 

Académie suisse des Sciences naturelles (SCNAT) 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 

Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités 
suisses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 

SUISSE Fr. 225.- UNION POSTALE Fr. 250. 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse 



Revue suisse de Zoologie 112 (4): 771-805; décembre 2005 



Additional notes on the fauna of Agromyzidae (Diptera) 
in Switzerland 

Milos CERNY 

CZ-763 63 Halenkovice 1, Czech Republic. 

E-mail: cerny.milos@centrum.cz 

Additional notes on the fauna of Agromyzidae (Diptera) in Switzerland. 

- New faunistic data on the Swiss Agromyzidae are given. 144 species are 
recorded including 92 species found in Switzerland for the first time. Each 
species included is accompanied by information on its host plants and 
geographical distribution. The current list of the Swiss Agromyzidae thus 
embraces 232 species. 
Keywords: Diptera - Agromyzidae - faunistics - biology - Switzerland. 

INTRODUCTION 

The Agromyzidae, with its 1165 species described from the Palaearctic Region 
(Scheirs et al., 1999) and 843 species from Europe (Martinez, 1998) is one of the larger 
families of Diptera. The Swiss fauna of Agromyzidae is little studied with 140 recorded 
species in 19 genera (Martinez, 1998). On the other hand, the study of Agromyzidae in 
Europe and especially in the other countries of Central Europe is relatively advanced 
(cf. e.g. Griffiths, 1980; Nowakowski, 1973; Spencer, 1964, 1966, 1972b, 1976; 
Zlobin, 1992, 1993a, 1993b, 1993c, 1993d, 1994, 1995, 2003a, 2003b). 

The following species were described directly from the territory of Switzerland 
(type locality in parentheses, abbreviation of cantons in following chapter): Agromyza 
alnibetulae Hendel, 1931 (type locality: Switzerland); Chromatomyia gentianella 
(Hendel, 1932) (Switzerland); Ch. hoppiella Spencer, 1990 (GR: Lenzerheide); 
Liriomyza polygalae Hering, 1927 (TI: Lugano); Napomyza dehor ii Spencer, 1966 
(VS: Orsières); Phytobia mallochi Hendel, 1924 (GR: Peist); Phytomyza abdita 
Hering, 1927 (ZH: Zürichberg); P. alpestris Hendel, 1920 (GR: St. Moritz); P. hoppi 
Hering, 1925 (SG: Pfäfers); P. narcissiflorae Hering, 1928 (GR: Parpan); P. hoppiana 
Hering, 1931 (GR: Zernez) as a synonym of P. pulsatiallae Hering, 1924 (Spencer, 
1976); P. thalictri Escher-Kiindig in Rougemont, 1912 (Switzerland); P. virgaureae 
Hering, 1926 (SZ: Rigi); P. thymi Hering, 1928 (GR: Lenzerheide) and 
Pseudonapomyza europaea Spencer, 1973 (VS: Stalden). P. atragenis Hendel, 1931 
(GR: Zernez) and P. philactaeae Hering, 1932 (GR: Vais) were synonymized with 
P. kaltenbachi Hendel, 1922 (Hendel, 1931-6; Papp, 1984). P. prenanthidis Hering. 
1932 (SG: Pfäfers) was synonymized with P. sonchi Robineau-Desvoidy, 1851 by 
Hendel (1931-6). Later Spencer (1965) synonymized P. sonchi with P. marginella 
Fallen, 1823. 



Manuscript accepted 07.02.2005 



772 M. CERNY 

The new Material examined by the author provided records on 144 species 
including 92 species which represent faunistic novelties for Switzerland. In total 
232 species are now known from Switzerland representing 28% of the European fauna. 

Agromyzids are mostly small to very small flies with a dark body sometimes 
showing a light metallic shine (Melanagromyza spp.), or a distinct yellow pattern (e.g. 
Liriomyza spp., Phytoliriomyza spp., Cerodontha spp.). The length of wings varies 
betwen 0.9 and 4.5 mm. The adult thorax is massive and the relatively broad abdomen 
consists of 6 visible segments. The eyes are broadly separated in both sexes, 1-2 rows 
of short orbital setulae are developed in addition to the regular orbital setae (except for 
Selachops). 

Trophic preferences are phytophagous, larvae typically feed upon tissues of 
living plants and form characteristic mines. The larvae are cylindrical, slightly tapered 
on both ends, their body consists of the cephalic segment, 3 thoracic and 8 abdominal 
segments. The length of body varies between 1.5-3.0 mm but the body of Phytobia is 
unusually long, sometimes reaching 25 mm. Larvae of most species live in the 
parenchym of leaves, some mine the surface of stems. Larvae of Phytobia feed upon 
bast of some wood plants. Hexomyza larvae form galls on twigs of trees and bushes. 
Larvae of Melanagromyza, Napomyza and Ophiomyia (partially) live within stems and 
on roots of host plants. Some Phytomyza larvae are specialized for life in flowers. 

Most Agromyzidae are oligophagous and their larvae prefer closely related 
species, genera or families of host plants. Some species are even monophagous and 
mine only one host plant. Only a relatively small group of agromyzids are widely 
polyphages and their spectrum of host plants includes numerous plants from many 
different families. Chromatomyia horticola (Goureau, 1851) may be mentioned as the 
best known and widely distributed polyphagous species mining a large series of plants. 

A review of Central European agromyzids with a certain economic importance 
was published by Spencer (1973). Some species used to be included among serious 
pests of crops, e.g. Liriomyza huidobrensis (Blanchard, 1926) recorded also from 
Switzerland (Martinez, 1998). However, many species damage only the general or 
aesthetical appearance of plants, and economical losses are not too serious. More 
important economic losses are known only during the mass occurrence of some species 
living on cereals, (Agromyza megalopsis Hering, 1933 or A. nigrella [Rondani, 1875]), 
on Fabaceae and Brassicaceae (Phytomyza rufipes Meigen, 1830) and some ornamental 
plants (Ch. horticola, L. huidobrensis). Larvae of Phytobia carbonaria (Zetterstedt, 
1848) bore the cambium of apple twigs. 

The density of agromyzid populations is very often seriously influenced by 
hymenopteran parasites from families Braconidae, Chalcididae, Eulophidae and 
Ichneumonidae. The strictly monophagous species with their close relations to host 
plants may be used as suitable models for ecological research. 

MATERIAL AND METHODS 

Species originating from Switzerland were found in the following collections: 

CBM private collection Bohuslav Mocek, Hradec Krâlové, Czech Republic, 8 spe- 
cimens. 



AGROMYZIDAE IN SWITZERLAND 773 

CMB private collection Miroslav B arrak, Praha, Czech Republic, 75 specimens. 

These specimens were collected with a Malaise Trap of the type M. Bartâk 

(Rohâcek et al., 1998) above the tree line on 2000 m a.s.l. in the Alps of the 

Canton Graubünden. 
MHNG Muséum d'histoire naturelle, Genève, Switzerland (Bernhard Merz), 284 

specimens . 
ZMUC Zoological Museum, University Copenhagen, Denmark (Rudolf Meier), 33 

specimens. 

Genera and species are arranged alphabetically in the two subfamilies 
Agromyzinae and Phytomyzinae. All relevant data from locality labels are included. 
Only selected important synonymy and new References (with regard to Switzerland) 
are quoted here, for full data see Martinez (1998), Papp (1984), Spencer & Martinez 
(1987) and Spencer (1990, 1992). 

Nomenclature of Agromyzidae and plant names follow Spencer (1990). 

The localities are grouped according to cantons (districts) of Switzerland and 
the following abbreviations are used: AG = Aargau; BE = Bern; FR = Fribourg; GE = 
Genève; GL = Glarus; GR = Graubünden (= Grisons); JU = Jura; NE = Neuchâtel; SG 
= St. Gallen; SH = Schaffhausen; SZ = Schwyz; TI = Ticino; VD = Vaud; VS = Valais; 
ZH = Zürich. 

MT on locality lables means Malaise Trap. 

The species which are new to Switzerland are marked with an asterisk (*) 
before the name. 

LIST OF SPECIES 
Subfamily Agromyzinae 
Agromyza abiens Zetterstedt, 1848 

Reference: Martinez, 1998: 269. 

Material examined: GE: Cartigny, Moulin de Vert, 350 m a.s.l., 1 9 , 2.vi.2002. 
GR: Lenzerheide near Sanaspans, 1500 m a.s.l., 1 9 , 7.viii.l992. VS: Leuk Bahnhof, 
625 m a.s.l., 1 ó\ 15.V.2000. ZH: Zürich, 500 m a.s.l., 1 9 , 10.viii.1991. AU B. Merz 
leg. (MHNG). 

Notes: This relatively large species with the orange frons and a broad epistoma 
was described from Sweden. A. abiens is generally distributed and locally common in 
Europe. Its occurrence in Japan was also confirmed. Larvae form broad leaf mines on 
Boraginaceae. Records include the genera Anchusa, Asperugo, Borago, Cynoglossum, 
Echium, Lycopsis, Pentaglottis, Pulmonaria and Symphytum. 

Agromyza albipennis Meigen, 1830 

Reference: Martinez, 1998: 269. 

Material examined: GE: Chancy Bord du Rhône, 350 m a.s.l., 1 â , 28.vii.2002. 
B. Merz leg. (MHNG); Russin, Les Baillets, 405 m a.s.l., 1 <? , l.vii.2002, B. Merz & 
Herrmann leg. (MHNG). 

Notes: A. albipennis is a Holarctic species which is common in Europe, espe- 
cially in its temperate and northern parts. It is less known from the Mediterranean area 



774 M. CERN Y 

(only Italy and Spain). Its occurrence in Japan, Kamchatka and North America (United 
States and Canada) is also documented. Larvae feed singly, forming linear-blotch mine 
on Gramineae, particularly on Phalaris arundinacea, less commonly on Poa and 
Hordeum, rarely on other grasses. 

*Agromyza bicaudata (Hendel, 1920) 

Material examined: VS: Leuk Pfynwald, 630 m a.s.l., 1 S , 21 .iv.1998, B. Merz 
& Botta leg. (MHNG). 

Notes: The type series includes localities in Germany and Austria. It is distri- 
buted in temperate Europe (Austria, British Isles, the Czech Republic, Germany, 
Hungary, Lithuania, Poland and Slovakia). Hendel (1931-6) mentioned this species 
from Finland a South Russia. This species is unusual in having a strongly developed 
presuturai de and specific male terminalia. Very probably a grass-feeder though its 
biology is not precisely known. New for Switzerland. 

Agromyza bromi Spencer, 1966 

Reference: Martinez, 1998: 269. 

Material examined: VS: Leuk Platten, 630 m a.s.l., 1 6 , 22 .iv.1998, B. Merz & 
B. Botta leg. (MHNG). ZH: Zürich Irchel, 500 m a.s.l., 1 6 , 16.viii.1998, B. Merz leg. 
(MHNG). 

Notes: The species is known from temperate Europe. The only recorded host is 
Ceratochloa unioloides (= Bromus catharticus) but other grasses are doubtless also 
attacked. 

* Agromyza cinerascens Macquart, 1835 

Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 cT , 21.iii.1999; Chancy, 
La Laire, 350 m a.s.l., 1 S , 1 .iv.2002; Russin, Les Baillets (Allondon), 390 m a.s.l., 1 
6, 24.iii.2003. ZH: Zürich Albisgütli, 490 m a.s.l., 1 S, 9.iv.l997; Zürich Ahmend, 
440 m a.s.l., 1 9 , 17.iv.1996. All B. Merz leg. (MHNG). 

Notes: This Palaearctic species was described from France; it is known from 
many countries of Europe (Austria. Belarus, Belgium, Croatia, the Czech Republic, 
Denmark, England, Estonia, Finland, Germany. Hungary, Italy incl. Sicily, Lithuania, 
the Netherlands, Norway, Poland, Romania, Slovakia, Spain, Russia, and Sweden), 
North Africa (Egypt and Tunisia) and Japan. Host plants are Gramineae, most 
frequently Dactyl is glomerata, sometimes Secale cereale. New for Switzerland. 

* Agromyza frontella (Rondani, 1875) 

Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 S , 8.vii.2001 , B. Merz 
leg. (MHNG). VS: Leuk Brentjong, 920 m a.s.l., 1 6, 15.V.2000, B. Merz leg. 
(MHNG). 

Notes: This Holarctic species was described from Italy (Parma). It is common 
in Austria, Belarus, the Czech Republic, Denmark, England. France, Germany, 
Hungary, Italy, Lithuania, Poland, Russia, Slovakia, Spain, Sweden, Canada, United 
States, Afghanistan. Israel, and Turkey. A. frontella belongs to the Agromyza species 
feeding on Leguminosae. The larva forms a characteristic mine on Medicago sativa 
and other Medicago spp., but also on Melilotus. New for Switzerland. 



AGROMYZIDAE IN SWITZERLAND 775 

*Agromyza hendeli Griffiths, 1963 

Material examined: ZH: Zürich Irchel, 500 m a.s.l., 1 S , 16.viii.1998, B. Merz 
leg. (MHNG). 

Notes: A. hendeli is distributed in temperate Europe and known from Austria, 
Belgium, the Czech Republic, Denmark, England, Estonia, Germany, North Italy, 
Latvia, Lithuania, the Netherlands, Poland, Scotland and Slovakia. The larvae form a 
broad blotch mine on Phragmites communis. New for Switzerland. 

*Agromyza idaeiana Hardy, 1853 

Material examined: GR: San Vittore, Rebberg, 290 m a.s.l., 1 S , 8.iv.l997, 
B. Merz leg. (MHNG). VS: Leuk Platten, 625 m a.s.l., 1 6, 30.V.2002, B. Merz leg. 
(MHNG). 

Notes: A Holarctic species known in the literature under the junior synonym 
Agromyza potentillae (Kaltenbach, 1864) (Bland, 2000). The species seems to be 
common in Europe (Austria, Belgium, the Czech Republic, England, Estonia, Finland, 
Germany, Hungary, Italy (Sardinia), Lithuania, the Netherlands, Norway, Poland, 
Russia, Slovakia, Spain, Sweden) as well as in Central and East Palaearctic 
(Uzbekistan, Kuril Islands and Japan). Largely distributed also in North America 
(Canada and United States). The larva forms a linear mine which later widens to a 
conspicuous blotch on Rosaceae (Agrimonia, Alchemilla, Comarum, Filipendula, 
Fragaria, Geum, Potentilla, Rosa, Rubus and Sanguisorba) . New for Switzerland. 

* Agromyza lucida Hendel, 1920 

Material examined: GIL: Cartigny, Moulin de Vert, 350 m a.s.l., 1 a , 9.VÜ.2002, 
B. Merz leg. (MHNG). 

Notes: A. lucida is known from the Holarctic Region and distributed especially 
in Europe (Austria, the Czech Republic, Denmark, England, Finland, France, 
Germany, Hungary, Ireland, Italy, Latvia, Lithuania, the Netherlands, Poland, Russia, 
Scotland, Slovakia, Spain, Sweden) and North America (Canada). Its occurrence is 
also confirmed in India. The larvae form a broad blotch mine on Gramineae, known 
foodplants are Glycera maxima and Deschampsia caespitosa. New for Switzerland. 

* Agromyza marionnae Griffiths, 1963 

Material examined: ZH: Zürich Albisgiitli, 500 m a.s.l., 1 <3\2.iv.l997,B.Merz 
leg. (MHNG). 

Notes: The species was described from England and later recorded from 
Germany, Ireland, Lithuania and Spain. According to Pakalniskis (1996) the larva 
mines stems of Vicia cracca and V. tetrasperma at borders of mixed forest and in urban 
cenoses. New for Switzerland. 

Agromyza mobilis Meigen, 1830 

Reference: Martinez, 1998: 269. 

Material examined: GE: Bernex Chante Merle, 420 m a.s.l., 1 â , 7.viii.2000, 
B. Merz & G. Bächli leg. (MHNG). GR: Zernez, Gondas, 1480 m a.s.l., 1 o\ 
8.viii.l996, B. Merz & G. Bächli leg. (MHNG). SH: Bargen-Mülital, 650 m a.s.l., 1 6 , 



776 M. CERNY 

28.viii.1992, B. Merz leg. (MHNG). TI: Gordola, 210 m a.s.l., 1 S, 14.ix.1989, B. 
Merz leg. (MHNG). ZH: Zürich Irchel, 500 m a.s.l., 1 6, 24.V.1996, B. Merz leg. 
(MHNG). 

Notes: A. mobilis is a species with a Palaearctic distribution, known to occur 
especially in Europe but also in China and Japan. It occurs namely in countries of 
temperate and northern Europe. In the Mediterranean area it was recorded only from 
Italy, Spain and the former Yugoslavia. Triticum represents the only confirmed known 
host plant but the larvae mine very probably also other species of Graminae. 

Agromyza nana Meigen, 1830 

Reference: Martinez, 1998: 269. 

Material examined: GR: Lenzerheide. 2000 m a.s.l., 1 6, 14. -21. vii. 2000, 
2 S S, 23.-3 l.viii. 2000, pasture, MT, B. Merz leg. (CMB). SH: Merishausen, Ladel, 
700 m a.s.l., 1 S, 13.V.1991, B. Merz leg. (MHNG). VS: Eggerberg, 850 m a.s.l., 1 S, 
24. vi .2000, B. Merz leg. (MHNG); Visperterminen Rothorn, 2250 m a.s.l., 1 S, 
26.viii.2001, B. Merz & Landry leg. (MHNG); Saas-Fee, 2400 m a.s.l., 1 S, 
22 .vii. 1965, 2 SS, 23 .vii. 1965, O. Lomholdt leg. (ZMUC). 

Notes: The Palaearctic A. nana belong to the species of the orobi-group (Zlobin, 
2000), which live in Leguminosae (Medicago sativa, Trifolium, Melilotus). With its 
morphology and the structure of male genitalia it is very similar to other species of this 
group, especially to A. frontella (Rondani, 1875), but stridulation border is species- 
specific (Tschirnhaus, 1971). Generally common in the Palaearctic Region including 
Egypt, Marocco, Tunisia, Turkey and eastern Siberia. Also recorded from India. 

Agromyza nigripes Meigen, 1830 

References: Griffiths. 1963: 132; Martinez, 1998: 269. 

Material examined: GE: Jussy, Prés-de- Villette, 475 m a.s.l., 1 â , 2.viii.2002, 
B. Merz leg. (MHNG). ZH: Zürich Albisgütli, 470 m a.s.l., 1 S, 19.viii.1997, B. Merz 
& Botta leg. (MHNG); Zürich Hönggerberg, 600 m a.s.l., 1 o* , 18. vi. 1992, B. Merz leg. 
(MHNG); Zürich Zürichberg, 650 m a.s.l., 1 S , 26.vii.1995, S. Ungricht leg. (MHNG). 

Notes: The species is known mainly from the Holarctic Region, (Europe and 
Canada), but it is also recorded from the Oriental Region (India). The larva forms a 
long, widening mine on the upper leaf surface of Glyceria maxima and the species also 
commonly occurs on Holcus. 

Agromyza poly goni Hering, 1941 

Reference: Martinez, 1998: 269. 

Material examined: GR: Zuoz, Nüd, 1700 m a.s.l., 1 S , 7 .viri. 1996, B. Merz & 
G. Bächli leg. (MHNG). 

Notes: The species was described from Germany and recently it was also 
recorded from the Czech Republic, France, Lithuania, Poland and Switzerland. The 
larva forms on Polygonum bistorta an irregular linear-blotch mine, which is not 
associated to the midrib. 

* Agromyza prespana Spencer, 1957 

Material examined: TI: Gordola. Bolle di Magadino, Zeltfalle, 220 m a.s.l., 
1 S , 19.vi.1995. B. Merz & G. Bächli leg. (MHNG). 



AGROMYZIDAE IN SWITZERLAND 777 

Notes: Spencer (1957) described this species from Macedonia. It is also known 
from Austria, British Isles, Croatia (Dalmatia), the Czech Republic, France, Hungary, 
Poland, Slovakia and Sweden. The larva forms a broad mine on Triticum aestivum and 
other grasses. New for Switzerland. 

*Agromyza pseudoreptans Nowakowski, 1967 

Material examined: GE: Chancy, La Laire, 350 m a.s.L, 1 S, 1 .vii .2001; 
Dardagny, Roulave, 420 m a.s.L, 1 6 , 30.vi.2001 . GR: Valbella, Casoja, 1550 m a.s.L, 
1 S , 13.vii.1996. SH: Merishausen-Ladel, 600 m a.s.L, 1 9 , 28.vii.1992. ZH: Flaach, 
Thurauen, 350 m a.s.L, 1 2, l.ix.1993; Zürich Waldgarten, 450 m a.s.L, 2 SS, 
24.V.1996; 460 m a.s.L, 1 S, l.v.1997. All B. Merz leg. (MHNG). 

Notes: This Holarctic species is common in Austria, Belgium, Canary Islands, 
the Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Hungary, 
Ireland, Italy, Lithuania, the Netherlands, Norway, Poland, Russia, Scotland, Slovakia, 
Sweden and North America (Canada and United States, incl. Alaska). The larva forms 
elongate linear-blotch mine, normaly adjoining margin of leaves of Urtica dioica. New 
for Switzerland. 

*Agromyza pseudorufipes Nowakowski, 1964 

Material examined: GR: Valbella, Casoja, 1550 m a.s.L, 1 S, 13.vii.1996, 
B. Merz leg. (MHNG). 

Notes: The species was described from Poland and relaible records are known 
from the Czech Republic, England, Germany, Lithuania, Poland, Romania, Slovakia 
and Japan. New for Switzerland. 

*Agromyza quadriseta Zlobin, 2001 

Material examined: GE: Bernex, Signal, 510 m a.s.L, 1 S , 24.V.2002. ZH: 
Embrach, Haumiili, 430 m a.s.L, 1 S, 29 .v. 1998, B. Merz leg. (MHNG). 

Notes: This species was recently described from Estonia, Lithuania, Finland and 
Russia (holotype). A. quadriseta belongs to the large group of Palaearctic species 
having long presuturai dorsocentrals and the costal vein strongly extending to Mj + 2- 
Biology of this species is unknown. New for Switzerland. 

*Agromyza rondensis Strobl, 1900 

Material examined: GE: Cartigny, Moulin de Vert, 350 m a.s.L, 1 S , 4.V.2003. 
B. Merz & M. Eggenberger leg. (MHNG). 

Notes: The species was described from Spain and later on recorded in other 
countries of West Palaearctic (Austria, Belarus, Belgium, Canary Islands, Crete, 
Croatia, the Czech Republic, England, Estonia, France, Germany, Hungary, Italy, 
Lithuania, Norway, Poland, Portugal, Romania, Scotland, Slovakia, Sweden and 
Turkey). The larva forms a leaf mine on Gramineae (Arrhenaterum, Bromus, 
Calamagrostis, Dactylis, Hordeum, Poa, Secale and Triticum). New for Switzerland. 

Agromyza sulfuriceps Strobl, 1898 

Reference: Martinez, 1998: 269. 

Material examined: VS: Saas-Fee, 2400 m a.s.L, 1 â , 22.vii.1965, 0. Lomholdt 
leg. (ZMUC). 



778 M. CERNY 

Notes: This small species with yellow frons and bright yellow antennae is 
known especially from the Holarctic Region but also recorded from tropical Africa 
(Nigeria). Host plants of this species are Potentilla and Sanguisorba. The larva forms 
a narrow upper surface gallery to start, then broadening and zigzagging to create a false 
blotch. 

*Melanagromyza astragali Spencer, 1976 

Material examined: SZ: Brunni, 900 m a.s.L, 1 6 , 26.vi.1990, B. Petersen leg. 
(ZMUC). VS: Leuk Brentjong, 930 m a.s.L, 1 6 , 3 .v. 1990, 900 m a.s.L, 1 S , 1 .vi. 1991 , 
B. Merz leg. (MHNG). 

Notes: This species was described by Spencer (1976) from Sweden from stems 
of Astragalus glyciphyllus and later on it was recorded from other countries of Europe 
(Czech Republic, France, Germany, Lithuania and Slovakia). New for Switzerland. 

Melanagromyza cunctans (Meigen, 1830) 

Reference: Martinez, 1998: 269. 

Material examined: VS: Visperterminen Dorf, 1400 m a.s.L, 1 S , 26.viii.2001, 
B. Merz & Landry leg. (MHNG). ZH: Zürich Irchel, 500 m a.s.L, 1 S , 16.viii.1998, B. 
Merz leg. (MHNG). 

Notes: The species was described from Spain and its known distrubution area 
includes other countries of temperate Europe and Scandinavia (Sweden) but it seems 
to be more common in the Mediterranean area (Canary Islands, Corsica, Crete, Croatia, 
Egypt, Italy, Maltese Islands and Turkey) and the Afrotropical Region (including 
Madagascar). Its occurrence was also confirmed in Central Palaearctic (Uzbekistan) 
and the Oriental Region (India). The larva forms a slender stem-gall on Lotus corni- 
culatus (Leguminosae). 

*Ophiomyia aeneonitens (Strobl, 1893) 

Material examined: VS: Leuk Pfynwald, 600-650 m a.s.L, 1 6, 15.V.1996, 
B. Merz & G. Bächli leg. (MHNG). 

Notes: The sole member of the genus with typically white squamae and fringes, 
and lacking the posterior crossvein on the wing. This is the second record of this 
species in Europe. It was described from a warm area of Austria. Biology of this 
species is not known. New for Switzerland. 

*Ophiomyia campanularum Stary, 1930 

Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.L, 1 S , 19.V.2001 , B. 
Merz leg. (MHNG). 

Notes: This species was described from the the Czech Republic and now it is 
known only from Germany, Slovakia and Sweden. The larva forms an external stem- 
mine with frass in large, widely-spaced grains on Campanula rotundifolia. New for 
Switzerland. 

*Ophiomyia cunctata (Hendel, 1920) 

Material examined: ZH: Zürich Oerlikon, 420 m a.s.L, 1 9 , vi.-x.1987, B. Merz 
leg. (MHNG). 



AGROMYZIDAE IN SWITZERLAND 779 

Notes: Widespread throughout much of Europe (Albania, Austria, Canary 
Islands, the Czech Republic, Denmark, England, Finland, France, Germany, Hungary, 
Italy, Lithuania, Norway, Poland, Russia, Scotland, Spain, Sweden, Turkey and the 
former Yugoslavia). The larva forms a whitish blotch mine along the midrib, with 
irregular offshoots into the leaf-blade on Crépis, Hypochoeris, Lapsana, Mycelis, 
Picris, Sonchus and Taraxacum. New for Switzerland. 

Ophiomyia curvipalpis (Zetterstedt, 1848) 

Reference: Martinez, 1998: 269. 

Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 â , 24.V.2002, B. Merz 
leg. (MHNG); Corsier-Port, 19, 16.-3 l.viii.2003, C. Besuchet leg. (MHNG). TI: 
Biasca Loderio, 300 m a.s.l., 1 o\ 5. vii. 1991, B. Merz leg. (MHNG). VS: Leuk 
Pfynwald, 680 m a.s.l., 1 ? , 7. vi. 200 1 , B. Merz & Landry leg. (MHNG); Leuk Platten, 
625 m a.s.l., 1 S , 2.V.1999, B. Merz leg. (MHNG); St. German, Brücke, 624 m a.s.l., 
1 6 , 3 .viri. 1998, B. Merz & G. Bächli leg. (MHNG). 

Notes: This species is distributed chiefly in the West Palaearctic but it is also 
known from Japan. The larva forms a narrow, inconspicuous stem- mine on Achillea 
millefolium, A. ptarmica, Anthémis tinctoria, Artemisia vulgaris, Matricaria inodora, 
and possibly also on Medicago sativa. 

* Ophiomyia galii Hering, 1937 

Material examined: VS: Visperterminen, Kreuz, 1400 m a.s.l., 1 3 , 3.vi.2003, 
B. Merz leg. (MHNG). 

Notes: The species was recorded from Corsica, the Czech Republic, England, 
France, Germany, Lithuania, Poland, Slovakia, and Spain. O. galii was described from 
Germany, and the type series was reared from Galium mollugo. The larva forms an 
external stem mine, with frass in large, widely spaced grains. According to Pakalniskis 
(1998b) it may also attack G. verum. New for Switzerland. 

* 'Ophiomyia inaequabilis (Hendel, 1931) 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 3 , 2.vii.2001 , B. Merz 
leg. (MHNG). 

Notes: This Mediterranean species was described from a single male originating 
from Turkey; later it was also recorded from the Czech Republic. Our record represents 
the westernmost boundary of the distribution area in Europe. Its biology is unknown. 
New for Switzerland. 

Ophiomyia nasuta (Melander, 1913) 

Reference: Martinez, 1998: 269. 

Material examined: SH, Merishausen, Ladel, 600 m a.s.l., 1 3, 28.iii.1992, 
B. Merz leg. (MHNG). 

Notes: This species occurs in the Holarctic Region. It is known especially from 
temperate and northern Europe, North America (Canada, United States) and Japan. Its 
only known host plant genus is Taraxacum sp. 

Ophiomyia orbiculata (Hendel, 1931) 

References: Spencer, 1964: 800; Martinez, 1998: 269. 



780 M. CERNY 

Material examined: GE: Bemex, Signal, 510 m a.s.l., 1 3 , 24.V.2002, B. Merz 
leg. (MHNG). NE: St. Blaise Les Riedes, 470 m a.s.l., 2 6 6, 19.V.2001, B. Merz leg. 
(MHNG). VS: Leuk Pfynwald, 600-650 m a.s.l., 1 6 , 15.V.1996, B. Merz & G. Bächli 
leg. (MHNG); Leuk Platten, 630 m a.s.l., 1 6, 22.iv.1998, B. Merz & Botta leg. 
(MHNG). 

Notes: The species was described from Austria based on two males and its 
distribution area includes mainly temperate and northern Europe. In the Mediterranean 
area it is known only from the former Yugoslavia and Turkey. The larva feeds as a 
stem-miner on Pisum sativum, and probably also on Vicia. 

Ophiomyia pinguis (Fallen, 1820) 

References: Spencer, 1973: 140; Martinez, 1998: 269. 

Material examined: GR: Valbella Casoja, 1500 m a.s.l., 1 6, 19.vii.1997, 
B. Merz leg. (MHNG). VS: Saas-Fee, 2300 m a.s.l., 1 6,1 9, 19.vii.1965, 1800 m 
a.s.l., 1 ? , 21. vii. 1965, O. Lomholdt leg. (ZMUC). 

Notes: O. pinguis is known to occur in the Palaearctic Region. The species is 
recorded from European countries including the Mediterranean area (Italy, Spain and 
the former Yugoslavia). It is also confirmed from Egypt and Turkey and penetrates 
through Tajikistan and Uzbekistan to China. The natural hosts of this species are 
Cichorium intybus, C. endivia, Lactuca sativa and Leontodon. 

*Ophiomyia submaura Hering, 1926 

Material examined: GR: Ardez, Bahnhof, 1350 m a.s.l., 1 6, 6.viii.l996, 
B. Merz & G. Bächli leg. (MHNG). 

Notes: O. submaura occurs in the West Palaearctic. It is confirmed from the 
Czech Republic, Germany, Hungary, Lithuania, Poland. Spain and Turkey. Pakalniskis 
(1996) reared it from Medicago falcata at borders of mixed forests and in brighter pine 
woods. New for Switzerland. 

*Ophiomyia vimmeri Cerny, 1994 

Material examined: GR: Valbella, Casoja, 1500 m a.s.l., 1 6 , 14.vii.1998, 
B. Merz leg. (MHNG). 

Notes: O. vimmeri was described from the Czech Republic, it is also known to 
occur in Slovakia. Its biology is unknown. New for Switzerland. 

Subfamily Phytomyzinae 

*Amauromyza (Cephalomyza) flavifrons (Meigen, 1830) 

Material examined: SH: Rtidlingen, 370 m a.s.l., 1 6 , 7.ix.l996, B. Merz leg. 
(MHNG). TI: Gordola. Bolle. 220 m a.s.l., 1 6, l.viii.1993, B. Merz & M. • 
Eggenberger leg. (MHNG). ZH: Zürich Oerlikon, 430 m a.s.l., 1 6, 23.V.1992, 
B. Merz leg. (MHNG). 

Notes: This Holarctic species is distributed in Alabania, Austria, Belgium, 
Corsica, the Czech Republic, Denmark, England, Finland, France, Germany, Hungary, 
Kyrgyzstan, Lithuania, the Netherlands, Norway, Poland, Romania, Sardinia, Scotland, 
Spain, Sweden. Turkey and North America (Canada and United States). The larva 



AGROMYZIDAE IN SWITZERLAND 781 

forms a white linear-blotch mine on many genera of Caryophyllaceae, particularly on 
Dianthus, Lachnis, Melandrium, Saponaria, Silène, Stellaria but also on Beta vulgaris 
and Spinacia oleracea (Chenopodiaceae). New for Switzerland. 

*Amauromyza (Cephalomyza) mihalyii Spencer, 1971 

Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.l., 2 SS, 19.V.2001 , 
B. Merz leg. (MHNG). 

Notes: This species was described from Hungary and later recorded from the 
Czech Republic. This record is a further evidence of A. (C.) mihalyii in Central Europe 
and represents the westernmost boundary of its occurrence. Its biology is unknown. 
New for Switzerland. 

Amauromyza (Cephalomyza) monfalconensis (Strobl, 1909) 

References: Spencer, 1992: 142; Martinez, 1998: 269. 

Material examined: GR: Lenzerheide Sundroina, 1600 m a.s.l., 1 S, 
16. vii. 2000, B. Merz leg. (MHNG). 

Notes: This species is distributed especially in temperate Europe. It is also 
recorded from Scandinavia (Sweden) and some countries of the Mediterranean area 
(Italy and Spain). Host plants and immature stages unknown; larva almost certainly 
feeds as internal stem-borer. 

*Aulagromyza luteoscutellata (de Meijere, 1924) 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 S, 25.viii.2001, 
B. Merz & Landry leg. (MHNG). 

Notes: The species is distributed in temperate and northern Europe (Belgium, 
the Czech Republic, Denmark, Finland, France, Germany, Lithuania, the Netherlands, 
Norway, Poland, Sweden) and Canada. It has not been recorded from the 
Mediterranean area, but it is known from India. The larva forms a short, funnel-shaped 
leaf mine, filled centrally with dark-green frass on Lonicera and Symphoricarpos . New 
for Switzerland. 

*Aulagromyza orphana (Hendel, 1920) 

Material examined: ZU: Zürich Katzensee, 440 m a.s.l., 1 S, 25.V.1999. 
B. Merz leg. (MHNG). 

Notes: Locally distributed particularly in Central Europe. Its occurrence is 
confirmed for Austria, Belgium, the Czech Republic, Denmark, England, France, 
Germany, Hungary, the Netherlands, Poland, Scotland, Slovakia, Spain and Turkey. 
The larva forms an external stem mine on Galium aparine, probably also on 
G. palustre. New for Switzerland. 

*Aulagromyza similis (Brischke, 1880) 

Material examined: SH: Merishausen, 550 m a.s.l., 1 $, 26.iv.1990, B. Merz 
leg. (MHNG). 

Notes: A European species occurring in Austria, the Czech Republic, England, 
Finland, France, Germany, Hungary, Ireland, Italy, Lithuania, Norway, Poland, 
Romania, Sweden and Wales. The larva forms a large whitish linear blotch leaf mine. 



782 M. CERNY 

frequently associated with the midrib on Knautia arvensis and Succisa pratensis. New 
for Switzerland. 

Aulagromyza trivittata (Loew, 1873) 

Reference: Martinez, 1998: 270. 

Material examined: ZH: Zürich Albisgütli, 490 m a.s.L, 1 S , 9.iv.l997, B. Merz 
leg. (MHNG). 

Notes: This species is distributed mainly in temperate Europe. In Scandinavia it 
is recorded from Norway and Sweden. It is not known from the Medirranean area. The 
larva feeds as internal stem borer on Galium mollugo. 

Calycomyza humeralis (von Roser, 1 840) 

Reference: Martinez, 1998: 270. 

Material examined: GR: Lenzerheide, 2000 m a.s.L, 3 S S , viii.2000, pasture, 
MT, B. Merz leg. (CMB). VS: Leuk Platten, 630 m a.s.L, 1 S, 22 .iv. 1998, B. Merz & 
Botta leg. (MHNG); Visperterminen Giw-Gebidemsee, 1900-2200 m a.s.L, 1 S, 
28.viii.2001, B. Merz & Landry leg. (MHNG). 

Notes: A semicosmopolitan species which is locally distributed in Europe but 
also known from East Palaearctic, North and South America, the Afrotropical and the 
Oriental Regions, Australia and the Hawaiian Islands. The larva forms a leaf mine on 
Aster, Bellis and Erigeron. 

*Cerodontha (Butomomyza) angulata (Loew, 1869) 

Material examined: ZH: Zürich Zürichberg, 500-600 m a.s.L, 3 ââ, 
16.viii.1992, B. Merz leg. (MHNG). 

Notes: The species is distributed in the Holarctic Region including Japan and 
Kuril Islands. The larva forms a long greenish leaf mine on Carex, rarely on Scirpus 
silvaticus. New for Switzerland. 

*Cerodontha (Butomomyza) rohdendorfi Nowakowski, 1967 

Material examined: ZH: Embrach, Haumüli, 400 m a.s.L, 1 S, 28.V.1997, 
B. Merz leg. (MHNG). 

Notes: Described from Poland and later recorded from the Czech Republic, 
Finland and Italy. The larva forms an upper or lower surface mine, filling the entire 
width of the leaf of Poa chaixii, Festuca pratensis and probably other related grasses. 
New for Switzerland. 

Cerodontha (Cerodontha) affinis (Fallen. 1823) 

References: Spencer, 1976: 177; Martinez, 1998: 270. 

Material examined: ZH: Horgen, 550 m a.s.L, 1 S , 24.V.1989, B. Merz leg. 
(MHNG). 

Notes: The species is common in Europe, particularly in the temperate zone. Its 
biology is unknown. 

Cerodontha (Cerodontha) denticornis (Panzer, 1 806) 

Reference: Martinez, 1998: 270. 

Material examined: GR: Lenzerheide, 1600 m a.s.L, 1 $ , 8.x. 1991, B. Merz & 
M. Eggenberger leg. (MHNG); Lenzerheide, 2000 m a.s.L, 1 â, 23.viii.-9.ix.2000, 



AGROMYZIDAE IN SWITZERLAND 783 

pasture, MT, B. Merz leg. (CMB). SH: Merishausen, Gräte, 520 m a.s.l., 1 S, 
1. vi. 1996, B. Merz & M. Eggenberger leg. (MHNG). VS: Saas-Fee, 1800 m a.s.l., 1 S , 
3 9 9, 21.vii.1965, O. Lomholdt leg. (ZMUC). TT: Lavertezzo, 560 m a.s.l., 1 9, 
12.ix.1989, B. Merz leg. (MHNG). ZH: Zürich AUmend, 460 m a.s.l., 2 SS, 
17.V.1995; Zürich Irchel, 500 m a.s.l., 1 S , 24.V.1996; Zürich Katzensee, 440 m a.s.l., 
1 S , 25 .v. 1996, B. Merz leg. (MHNG). 

Notes: A common Palaearctic species described from Germany. Known to occur 
also in the Afrotropical and Oriental Regions including Taiwan. The larva feeds mainly 
in the leaf sheath of Gramineae. 

Cerodontha (Cerodontha) fulvipes (Meigen, 1830) 

References: Spencer, 1976: 177, 180; Martinez, 1998: 270. 
Material examined: GL: Pragelpass, 1500 m a.s.l., 1 S , 5.viii.l991, G. Bächli, 
Beuk & B. Merz leg. (MHNG). GR: Lenzerheide, 2000 m a.s.l., 1 ó\ viii.2000, 
pasture, MT, B. Merz leg. (CMB). TI: Gordola, Bolle d. M., 205 m a.s.l., 2 SS, 
6.VÜ.2001, B. Merz leg. (MHNG). VS: Baltschieder, Rotten-Ufer, 650 m a.s.l., 1 S, 
19.V.1996, B. Merz leg. (MHNG); Saas-Fee, 2000 m a.s.l., 1 S , 1 9 , 18.vii.1965, 2300 
m a.s.L, 2 9 9, 19.vii.1965, 1800 m a.s.l., 1 9 , 21 .vii. 1965, 0. Lomholdt leg. (ZMUC). 
ZH: Embrach Haumuli, 400 m a.s.L, 3 SS, 24.vi.1995, 1 S , 19.vi.1998, Zürcher leg. 
(MHNG); Zürich Irchel, 500 m a.s.l., 1 S , 16.viii.1998, B. Merz leg. (MHNG). 

Notes: A Palaearctic species which is generally distributed in Europe and 
recorded also from China. Poa trivialis is known as a host plant but the larva feeds 
probably on other grasses as well. 

^Cerodontha (Cerodontha) unguicornis Hendel, 1932 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 S, 23 .-31. viii.2000, 
pasture, MT, B. Merz leg. (CMB); Valbella, Casoja, 1600 m a.s.l., 1 9 , 1 8. vii. 2000, B. 
Merz leg. (MHNG). VS: Saas-Fee, 1800 m a.s.L, 1 S, 21. vii. 1965, O. Lomholdt leg. 
(ZMUC). 

Notes: The species was described from Central Europe. It is known from 
Austria, the Czech Republic and Germany. Its biology is still unknown. New for 
Switzerland. 

^Cerodontha (Dizygomyza) bimaculata (Meigen, 1830) 

Material examined: BE: Tramelan, La Tourbière, 995 m a.s.L, 1 S , 4.vi.2003, 
Merz, Haenni & Rapp leg. (MHNG). GR: S. Vittore, Rebberg, 300 m a.s.L, 1 S, 
4.VÜL1997, B. Merz leg. (MHNG). TI: Biasca, 350 m a.s.L, 1 S, 18.V.1991, B. Merz 
leg. (MHNG); Mte. Caslano, 400 m a.s.l., 1 S , 20 .v. 1991, B. Merz leg. (MHNG). VS: 
Morgins, Portes du Soleil (Monthey), 1700-1950 m a.s.l., 1 S , 22 .vi. 2003, B. Merz leg. 
(MHNG). 

Notes: This species is common in Europe. It is also known from the Kuril 
Islands and Japan. The larva forms a narrow mine on Luzula, particularly on L. pilosa. 
New for Switzerland. 

^Cerodontha (Dizygomyza) crassiseta (Strobl, 1900) 

Material examined: GR: Lenzerheide, 2000 m a.s.L, 1 S, viii.2000, pasture, 
MT,B. Merz leg. (CMB). 



784 M. CERNY 

Notes: A European species which is distributed especially in temperate Europe. 
In Fennoscandia known only from Sweden, and in the Mediterranean area recorded 
only from Italy and Spain. Dactylis glomerata and Poa compressa are known as host 
plants of this species. New for Switzerland. 

*Cerodontha (Dizygomy za) fasciata (Strobl, 1880) 

Material examined: VS: Leuk Pfynwald, 630 m a.s.l., 1 S , 21 .iv.1998, B. Merz 
& Botta leg. (MHNG); Visperterminen Dorf, 1400 m a.s.l., 1 S , 26.viii.2001, B. Merz 
& Landry leg. (MHNG). 

Notes: A species with a Holarctic type of distribution, known from temperate 
and northern Europe. It has not been recorded from the Mediterranean area. In North 
America occurring in Canada and the United States. Its biology is unknown. New for 
Switzerland. 

*Cerodontha (Dizygomy za) griffithsi Nowakowski, 1967 

Material examined: VS: Saas-Fee, 2000 m a.s.l., 1 S, 22.vii.1965, 1 6, 
27 .vii. 1965, O. Lomholdt leg. (ZMUC). 

Notes: This species was described from one male originating from the Austrian 
Alps. Later it was found in France. Its discovery in the Swiss Alps is thus not 
surprising. Recently it was captured in the High Tatra Mts in Slovakia (Cerny & Vaia, 
2005). Its biology is unknown. New for Switzerland. 

*Cerodontha (Dizygomyza) luctuosa (Meigen, 1830) 

Material examined: GR: Zernez, Gondas, 1480 m a.s.l., 1 S, 4.viii.l996, 
B. Merz & G. Bächli leg. (MHNG). 

Notes: The species occurs in the Holarctic Region. It is known particularly from 
Europe, but also from Tunisia, China, Japan and North America (Canada, United 
States, incl. Alaska). The larva feeds on Juncus effusus. New for Switzerland. 

*Cerodontha (Dizygomyza) luzulae (Groschke, 1957) 

Material examined: ZH: Zürich Albisgiitli, 450 m a.s.l., 1 6,1 .v. 1995, B. Merz 
leg. (MHNG). 

Notes: This European species was recorded from some Central European 
countries (the Czech Republic, Germany. Hungary and Poland) and the British Isles. 
Luzula sylvatica is the only known host plant of C. (D.) luzulae. New for Switzerland. 

Cerodontha (Dizygomyza) morosa (Meigen, 1830) 

References: Hendel, 1931-6: 92; Martinez, 1998: 270. 

Material examined: GR: S. Vittore Rebberg, 300 m a.s.l., 1 â , 4.viii.l997, 
B. Merz leg. (MHNG). 

Notes: This Holarctic species is common in Europe and known to occur also in 
the East Palaearctic (China and Japan) and the Oriental Regions (Philippines). The 
larva forms a relatively short and broad mine, which not extends to the leaf base of 
Carex. 

*Cerodontha (Dizygomyza) spinata (Groschke, 1954) 

Material examined: VS: Visperterminen Rothorn, 2250 m a.s.l., 2 ôâ, 
26.viii.2001, B. Merz & Landry leg. (MHNG). 



AGROMYZIDAE IN SWITZERLAND 785 

Notes: The species is known from Austria, the British Isles, the Czech Republic, 
Germany and Poland. The larva feeds on Car ex sylvatica and C. pilo sa. New for 
Switzerland. 

*Cerodontha (Poemyza) alpina Nowakowski, 1967 

Material examined: GR: Lenzerheide, St. Cassian, 1450 m a.s.l., 1 S, 
12.vii.1998, B. Merz leg. (MHNG). 

Notes: This mountain species was described from the upper forest zone of the 
East Carpathians (Poland) and later it was found in Austria. Our record of C. (P.) alpina 
in the Swiss Alps could be expected. The larva forms a leaf mine on Poa alpina and 
Trisetum alpestre. New for Switzerland. 

Cerodontha (Poemyza) atra (Meigen, 1830) 

References: Hendel, 1931-6: 38; Nowakowski, 1973: 91; Martinez, 1998: 270. 

Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 â , 24.V.2002, B. Merz 
leg. (MHNG). 

Notes: This species is common in Europe and known to occur in Japan. Some 
species of Gramineae are host plants of this species, in particular Calamagrostis 
epigeios and Agrostis alba. 

^Cerodontha (Poemyza) beigerae Nowakowski, 1973 

Material examined: GE: Russin, Les Baillets, 405 m a.s.l., 1 â , 30.vi.2001, 
B. Merz leg. (MHNG). 

Notes: Originally known only from Central Europe (the Czech Republic, 
Germany, Hungary and Poland). Recently Zlobin (1986, 1992) recorded C. (P.) 
beigerae also from East Siberia and the Far East. The larva feeds on Calamagrostis 
canescens, C. arundinacea, C. villosa and Agrostis canina. New for Switzerland. 

^Cerodontha (Poemyza) calamagrostidis Nowakowski, 1967 

Material examined: GE: Bernex, Chante-Merle, 415 m a.s.l., 1 S , 20.V.2002, B. 
Merz leg. (MHNG). 

Notes: This Holarctic species is distributed in Europe but it has not been found 
in the Mediterranean area. In North America known from Canada. Calamagrostis sp. 
and Alopecurus pratensis are the known host plants. New for Switzerland. 

* Cerodontha (Poemyza) incisa (Meigen, 1830) 

Material examined: ZH: Zürich Zürichberg, 450-650 m a.s.l., 1 â , 20.viii.1992, 
B. Merz leg. (MHNG). 

Notes: This Holarctic species is common in Europe but also known from China. 
Pakistan, Japan and North America. The larva forms a leaf mine on Gramineae, most 
commonly on Agropyron, Calamagrostis, Festuca and Phalaris but Nowakowski 
(1973) recorded 17 additional European genera. New for Switzerland. 

* Cerodontha (Poemyza) lateralis (Macquart, 1835) 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 6, 23.-31.viii.2000, 
pasture, MT, B. Merz leg. (CMB). VS: Branson, Follatères, 450-950 m a.s.l., 1 6, 
29.iii.2002;LeukPfynwald,630ma.s.l., 1 S, 25.V.1997, 600 m a.s.l., 1 o\ 2. vii. 2001. 
AHB. Merz leg. (MHNG). 



786 M. CERNY 

Notes: This Palaearctic species is common in the whole of West Palaearctic and 
penetrates to North Africa (Tunisia), Turkey, China and Japan. The larva forms a linear 
mine running towards the apex of the leaf and this can widen and become almost 
blotch-like on Gramineae. New for Switzerland. 

*Cerodontha (Poemyza) lyneborgi Spencer, 1972 

Material examined: GE: Chancy, La Laire, 350 m a.s.L, 1 a, 24.iv.2003, 
B. Merz & F. Amiet leg. (MHNG); Russin, Les Baillets, 405 m a.s.l., 1 S , 30.vi.2001, 
B. Merz leg. (MHNG). 

Notes: This species was described by Spencer (1972a) from Spain and later on 
it was found in other countries of Europe (the Czech Republic, Hungary, Italy, 
Lithuania and Russia). Its biology is unknown. New for Switzerland. 

Cerodontha (Poemyza) muscina (Meigen, 1830) 

Reference: Martinez, 1998: 270. 

Material examined: GR: Lenzerheide, 2000 m a.s.L, 2 â â , viii.2000, pasture, 
MT, B. Merz leg. (CMB); Lenzerheide, gegen Sanaspans, 1500 m a.s.L, 1 S, 
7.VÜL1992, B. Merz leg. (MHNG). VS: Visperterminen Dorf, 1400 m a.s.l. , 1 6, 
26.viii.2001, B. Merz & Landry leg. (MHNG). ZH: Zürich Katzensee, 450 m a.s.L, 
1 â, l.viii.1995; Zürich Ziegelhütte, 460 m a.s.L, 1 S, 29.vi.1997, B. Merz leg. 
(MHNG). 

Notes: This Holarctic species is common in Europe including the Mediterranean 
area (Spain, Italy and Slovenia), but also in North America. The larva forms a leaf mine 
on Gramineae. Mines were recorded on Dactylis, Festuca, Holcus, Milium and Poa. 

*Cerodontha (Poemyza) pygmaea (Meigen, 1830) 

Material examined: GR: Lenzerheide, 2000 m a.s.L, 1 S, 23 .-31. viii.2000, 
pasture, MT, B. Merz leg. (CMB). TI: Biasca, Loderio, 350 m a.s.L, 1 3 , 26.iii.1998, 
B. Merz leg. (MHNG). 

Notes: This species from the mma-group shows a Holarctic type of distribution 
and it is well known from a greater part of Europe. The larva forms a leaf mine on 
Gramineae. New for Switzerland. 

*Cerodontha (Poemyza) spencerae Zlobin, 1993 

Material examined: BE: Berner Alpen, Grindelwald env., 2500 m a.s.L, 4 S 6 , 
16.vi.1997, B. Mocek leg. (CBM). 

Notes: This Holarctic species is based on the type series originating from 
Russia, Estonia and Kazakhstan. In the Palaearctic Region C. (P.) spencerae was 
recorded from the British Isles, the Czech Republic, Finland, Italy, Latvia, Lithuania, 
Slovakia, East Siberia and the Russian Far East incl. Sakhalin. In Europe it was once 
bred from Agropyron repens. New for Switzerland. 

*Cerodontha (Xenophytomyza) atronitens (Hendel, 1920) 

Material examined: VS: Leuk Pfynwald, 600-650 m a.s.L, 1 ó\ 19.V.1996, 
B. Merz & G. Bächli leg. (MHNG). 



AGROMYZIDAE IN SWITZERLAND 787 

Notes: This species is distributed in temperate and northern Europe, but not yet 
found in the Mediterranean area. Host plants are unknown but certainly it is an internal 
feeder in the leaf-sheath or stem of Gramineae. New for Switzerland. 

Cerodontha (Xenophytomyza) biseta (Hendel, 1920) 

Reference: Martinez, 1998: 270. 

Material examined: GE: Cartigny, Moulin de Vert, 350 m a.s.l., 1 S , 9.vii.2001 , 
B. Merz leg. (MHNG). VS: Leuk Pfynwald, 600 m a.s.l., 1 o\ 6.vi.2001, 1 S, 
7.vi.2001, B. Merz & Landry leg. (MHNG). ZH: Zürich Hönggerberg, 530 m a.s.l., 1 
S , 9.vi.l998; Zürich Waldgarten, 460 m a.s.l., 1 S , 7.vi.l997; Zürich Ziegelhütte, 460 
m a.s.1., 2 S S , 29.vi.1997, B. Merz leg. (MHNG). 

Notes: C. (X.) biseta shows a Holarctic distribution and it is known to occur 
particularly in Europe including the Mediterranean area but also in Japan and North 
America (Canada and United States). The single specimen of C. (X.) biseta collected 
from the Jamaican Blue Mountains represents the first Neotropical record (Boucher, 
2003). Host plants unknown, certainly an internal feeder in Gramineae. 

*Chromatomyia fuscula (Zetterstedt, 1838) 

Material examined: GR: Lenzerheide, 2000 m a.s.l., pasture, 1 S, 23.- 
31.viii.2000, MT, B. Merz leg. (CMB). SZ: Unteriberg, O. Weid, 1600 m a.s.l., 1 S , 
24.vii.2001. TI: Biasca, Loderio, 350 m a.s.l., 1 S, 15.iii.1994, 1 S, 23.iii.1996, 1 S, 
26.iii.1998. VS: Branson, Follatères, 450-950 m a.s.1., 2 S S , 29.iii.2002, B. Merz leg. 
(MHNG); Leuk Pfynwald, 600 m a.s.l., 1 o\ 6. vi .2001, B. Merz & Landry leg. 
(MHNG); Visperterminen Dorf, 1550 m a.s.l., 1 ó\ 27.viii.2001, B. Merz & Landry 
leg. (MHNG). 

Notes: Ch. fuscula has a Holarctic distribution. In Europe this species is known 
to occur chiefly in temperate and northern parts. Also recorded in the East Palaearctic: 
Russia (Asian part), Kamchatka and Japan. The larva forms leaf-mines on Gramineae. 
New for Switzerland. 

Chromatomyia gentianella (Hendel, 1932) 

References: Hendel, 1931-6: 311; Spencer, 1990: 396; Martinez, 1998: 270. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 2 SS, 21-31.viii.2000, 
pasture, MT, B. Merz leg. (CMB). 

Notes: This mountain species is only known from Central Europe (Austria, 
North Italy, Poland and Switzerland). Larva forms a linear mine on Gentiana. 

Chromatomyia hoppiella Spencer, 1990 

References: Spencer, 1990: 396; Martinez, 1998: 270. 

Material examined: GR: Ausserferrera, 1300 m a.s.l., 1 S, 15.viii.1991, 
B. Merz & A. Freidberg leg. (MHNG); Lenzerheide, 2000 m a.s.l., 1 S, viii.2000. 
pasture, MT, B. Merz leg. (CMB); Valbella Casoja, 1500 m a.s.l., 1 S , 1 9. vii. 2000, B. 
Merz leg. (MHNG). 

Notes: Ch. hoppiella was described recently from Switzerland (Lenzerheide), 
based on the male holotype which was bred from Gentiana excisa. Our records 
represents the first additional faunistic data after the description. 



788 M. CERNY 

Chromatomyia hordeola (Goureau, 1851) 

Reference: Martinez, 1998: 270. 

Material examined: VS: Branson Follatères, 450-950 m a.s.l., 1 3 , 29.iii.2002, 
B. Merz leg. (MHNG). ZH: Zürich Oerlikon, 430 m a.s.l., 1 $ , 23.V.1992, leaf mine 
ex Centaurea sp., B. Merz leg. (MHNG). 

Notes: A polyphagous species, common in the Palaearctic, Afrotropical and 
Oriental Regions. Ch. horticola is a well known and common species mining a large 
spectrum of host plants, Griffiths (1967) and Spencer (1973) recorded host plants 
belonging to 34 families. The larva forms a long whitish upper surface corridor, which 
may go to the lower surface. 

* Chromatomyia isicae (Hering, 1962) 

Material examined: TI: Biasca, Loderio, 300 m a.s.l., 1 3 , 9.iv.l995, B. Merz 
leg. (MHNG). 

Notes: This species was described from Austria. It is also recorded from the 
Czech Republic, Finland, Germany, Ireland, Lithuania, Norway and Sweden. Its 
biology is unknown. New for Switzerland. 

Chromatomyia milii (Kaltenbach, 1864) 

Reference: Martinez, 1998: 270. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 3 , viii.2000, pasture, 
MT, B. Merz leg. (CMB); Valbella Casoja, 1550 m a.s.l., 1 3, 13.vii.1996, 1 3, 
15.vii.1996, B. Merz leg. (MHNG). TI: A. d'Arena, 1700 m a.s.l., 1 3 , 20.vii.1983, C. 
Besuchet leg. (MHNG); Biasca Loderio, 350 m a.s.l., 1 3 , 23.iii.1996, 300 m a.s.l., 1 
3, 9.ÌV.1995, B. Merz leg. (MHNG); Gordola, Bolle d. M., 205 m a.s.l., 1 3, 
6.VÜ.2001, B. Merz leg. (MHNG). ZH: Embrach Haumüli, 430 m a.s.l., 1 3, 
3 .vii . 1 996 , B . Merz leg . (MHNG) ; Zürich Albisgütli , 450 m a .s .1 . , 1 3 , 27 .iii . 1 994 , 500 
m a.s.l., 1 3 , 2.iv.l997, B. Merz leg. (MHNG); Zürich Ahmend, 450 m a.s.l., 2 3 3 , 
30.iii.1994, B. Merz leg. (MHNG); Zürich Katzensee, 440 m a.s.l., 1 3 , 19.iv.1997, B. 
Merz leg. (MHNG); Zürich, 1 3 , 26. vii. 1987, B. Merz leg. (MHNG). 

Notes: A Holarctic species which is common in the West Palaearctic and in 
North America (Canada) but it is also recorded from India. This species feeds in the 
larval stage on a variety of soft-leaved wild grasses in mesic and woodland habitats. It 
is not a pest of cultivated cereals, and large marsh grasses, such as Phragmites; 
Phalaris and Glyceria are also apparently avoided. 

Chromatomyia nigra (Meigen, 1830) 

Reference: Martinez, 1998: 270. 

Material examined: GR: Lenzerheide Sundroina, 1550 m a.s.l., 1 3, 
18.vii.1997, B. Merz leg. (MHNG). SG: Chäserrugg, 1850 m a.s.l., 1 3 , 15.viii.1997, 
B. Merz leg. (MHNG). 

Notes: A Holarctic species. Its distribution area in the Palaearctic and Nearctic 
Regions seems to be larger than in the related Ch. milii. It was also recorded in the 
Oriental Region. The larva forms a narrow, whitish linear mine on many genera of 
Gramineae. 



AGROMYZIDAE IN SWITZERLAND 789 

*Chromatomyia opacella (Hendel, 1935) 

Material examined: GR: Lenzerheide, Sanaspans, 2150 m a.s.L, 1 S, 
14.vii.1996, B. Merz leg. (MHNG). 

Notes: This mountain species is distributed up to the subalpine and alpine zones 
of the Polish Tatra Mts. It penetrates far to the North (Iceland and Faroes Islands). 
Recently recorded from Central Europe (Cerny & Vaia, 2005), known also from the 
British Isles and North Italy. One record from Greenland is mentioned by Griffiths 
(1980). Immature stages and the mine are unknown. Sesleria tatrae and Trisetum 
fuscum are recorded as host plants. Our record extends its range of alpine localities 
from Austria and Italy to Switzerland. New for Switzerland. 

*Chromatomyia pseudomilii Griffiths, 1980 

Material examined: VS: Visperterminen Dorf, 1550 m a.s.L, 1 6 , 27.viii.2001, 
B. Merz & Landry leg. (MHNG). 

Notes: This mountain species was described from Germany; it is also known 
from mountains of the Czech Republic (Cerny & Vaia, 1996), from the High Tatra Mts 
in Slovakia (Cerny & Vaia, 2005) as well as from North America (United States and 
Alaska). Its occurrence in further European mountains is very likely. The larva forms 
leaf mines on Gramineae and Carex. New for Switzerland. 

Chromatomyia ramosa (Hendel, 1923) 

References: Spencer, 1992: 142; Martinez, 1998: 270. 

Material examined: JU: Alle, 450 m a.s.L, 1 S, 6.ix.l989, B. Merz leg. 
(MHNG). ZH: Volketswil, Homberg, 500 m a.s.L, 1 S, 7.vi.l995, B. Merz leg. 
(MHNG). 

Notes: A common species in Europe which was also recorded from China. The 
larva feeds primarily along midrib, forming short offshoots into the leaf-blade in 
Dipsacus, Knautia and Succisa. 

*Galiomyza galiivora (Spencer, 1969) 

Material examined: GE: Jussy, Prés-de- Villette, 475 m a.s.L, 1 â , 2.viii.2002, 
B. Merz leg. (MHNG). 

Notes: G. galiivora was described by Spencer (1969) from specimens bred from 
Galium in the United States. It is also recorded in Canada and Europe (Belgium, 
Corsica, the Czech Republic, Germany, Lithuania, Poland and Slovakia). New for 
Switzerland. 

Galiomyza morio (Brischke, 1880) 

Reference: Martinez, 1998: 270. 

Material examined: NE: St. Blaise Les Riedes, 470 m a.s.L, 1 S , 19.V.2001, 
B. Merz leg. (MHNG). 

Notes: Also this species is mining Galium, but it was also bred from Asp end a 
odorata (= Galium odoratum). Its distribution area includes Europe and Japan. The 
larva forms a narrow linear mine which may largely fill small leaves and produce a 
secondary blotch. 



790 M. CERNY 

*Liriomyza approximata (Hendel, 1920) 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 â, 21-31.viii.2000, 
pasture, MT, B. Merz leg. (CMB). 

Notes: This species of mixed forests is known from Central and Northern 
Europe where it was found in Austria, Finland, Germany, Hungary, Northern Italy and 
Poland. The larva forms an upper surface blotch mine on Daphne mezereum. New for 
Switzerland. 

Liriomyza congesta (Becker, 1903) 

Reference: Martinez, 1998: 270. 

Material examined: SH: Merischausen, Ladel, 700 m a.s.l., 1 S , 13.V.1991, 
B. Merz leg. (MHNG). ZH: Zürich Katzensee, 440 m a.s.l., 1 S , 25.V.1996, B. Merz 
leg. (MHNG). 

Notes: Generally distributed in the West Palaearctic but apparently ranging to 
Japan. Common in the Mediterranean area (Albania, Corsica, Egypt, Italy incl. 
Sardinia, Maltese Islands, Spain and Turkey). The larva forms an upper surface linear 
mine exclusively on Leguminosae, with many genera being attacked, in particular 
Medicago, Lathyrus, Pisum and Vicia. 

'"Liriomyza erucifolii de Meijere, 1944 

Material examined: TI: Biasca, Loderio, 300 m a.s.l., 1 S , 9.iv.l995, B. Merz 
leg. (MHNG). 

Notes: This species is based on the male holotype from France and later 
recorded from the Czech Republic, Denmark, England, the Netherlands and Poland. 
Recently found on the Maltese Islands (Cerny, 2005b). The larva forms a relatively 
long linear mine, normally beginning near the apex of the leaf on Senecio erucufolius 
and S.jacobaea. New for Switzerland. 

Liriomyza flaveola (Fallen, 1823) 

Reference: Martinez, 1998: 271. 

Material examined: GE: Chancy, La Lake, 350 m a.s.l., 2 SS, 24.iv.2003, 
B. Merz & F. Amiet leg. (MHNG). ZH: Zürich, 600 m a.s.l., 1 S , 10.vi.1991, B. Merz 
leg. (MHNG). 

Notes: A common European species but known to occur also in Turkey, eastern 
Russia (East Siberia, Far East, Kamchatka. Kuril Islands), Uzbekistan, Japan and India. 
The larva forms a narrow, whitish leaf mine on Gramineae, occurring on many genera, 
particularly Bromus, Dactylis, Holcus and Poa, also on cultivared Avena sativa and 
Hordeum vulgare. 

'"Liriomyza lutea (Meigen. 1830) 

Material examined: VS: Visperterminen, 1460 m a.s.l., 1 S, 23 .vii. 1992, 
B. Merz leg. (MHNG). 

Notes: Widespread in Europe but local, frequently occurring in large numbers 
in association with the food-plants. The larva feeds in individual seeds on Angelica 
sylvestris, Heracleum sphondylium and Pastinaca sativa, probably also on 
Laserpitium. New for Switzerland. 



AGROMYZIDAE IN SWITZERLAND 791 

*Liriomyza obliqua Hendel, 1931 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 S, 21-31.viii.2000, 
pasture, MT, B. Merz leg. (CMB). 

Notes: L. obliqua is distributed in temperate Europe and in the Mediterranean 
area (Austria, Bulgaria, Germany, Lithuania, Romania, Slovakia, former Yugoslavia 
and Ukraine). Its biology is unknown. New for Switzerland. 

Liriomyza polygalae Hering, 1927 

Reference: Martinez, 1998: 271. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 2 S â , viii.2000, pasture, 
MT,B. Merz leg. (CMB). 

Notes: The species was described by Hering (1927) from Lugano (TI) in 
Switzerland. It is known to occur mainly in Central Europe but also in England. This 
is the second record from Switzerland. The larva forms an irregular linear-blotch mine 
on Poly gala vulgaris. 

*Liriomyza taraxaci Hering, 1927 

Material examined: VS: Leuk Pfynwald, 630 m a.s.l., 2 SS, 23.iv.1998, 
B. Merz & Botta leg. (MHNG). 

Notes: A Holarctic species which is common in Europe, but known only from 
Slovenia and Spain in the Mediterranean area. In North America recorded from Canada 
and United States. The larva forms a somewhat irregular, elongate blotch leaf mine on 
Taraxacum. New for Switzerland. 

*Metopomyza flavonotata (Haliday, 1833) 

Material examined: VS: Leuk Platten, 625 m a.s.l., 1 S, 30.V.2002; 
Visperterminen Kreuz, 1400 m a.s.l., 1 S , 3.vi.2003, B. Merz leg. (MHNG). 

Notes: This is a type species of the genus Metopomyza. It is common in Europe 
as well as in Russia (incl. Siberia and Far East) and Japan. Its biology is unknown, 
Pakalniskis (1998a) recorded Deschampsia caespitosa as a host plant for the first time. 
New for Switzerland. 

* Metopomyza nigriorbita (Hendel, 1931) 

Material examined: GE: Bernex, Chante-Merle, 415 m a.s.l., 1 6 , 16.viii.2002, 
B. Merz leg. (MHNG). 

Notes: A Palaearctic species which is common in Europe but not known in 
southern parts. Recorded also from Japan. The puparium is dark brown, with posterior 
spiracles each having a bunch of about 6 irregular bulbs (Spencer, 1976) but the host 
plant is unknown. New for Switzerland. 

*Metopomyza scutellata (Fallen, 1823) 

Material examined: GE: Jussy, Prés-de- Villette, 475 m a.s.l., 1 S , 2.viii.2002, 
B. Merz leg. (MHNG); Russin, Les Baillets, 405 m a.s.l., 1 S, l.vii.2002, B. Merz & 
Herrmann leg. (MHNG). VS: Leuk Pfynwald, 630 m a.s.l., 1 S, 6.vi.l997, B. Merz 
leg. (MHNG), 625 m a.s.l., 1 S, 16.V.2000, B. Merz & Ulrich leg. (MHNG). ZH: 



792 . M. CERNY 

Zürich, 500 m a.s.L, 2 Sa, 10.viii.1991, B. Merz leg. (MHNG); Zürich Waldgarten, 
460 m a.s.L, 1 S , 30.vii.1997, B. Merz leg. (MHNG). 

Notes: A Palaearctic species which is, in addition to Europe, also recorded from 
Kazakhstan, Kyrgyzstan, eastern Russia (East Siberia, Far East) and Japan. Species of 
Carex are host plants. New for Switzerland. 

*Metopomyza xanthaspis (Loew, 1858) 

Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.L, 2 6 6 , 19.V.2001 , 
B. Merz leg. (MHNG). VS: Grächen, 1450 m a.s.L, 1 S, 5.vL1987, B. Merz leg. 
(MHNG); Leuk Pfynwald, 630 m a.s.L, 1 3, 6.vi.2001, B. Merz & B. Landry leg. 
(MHNG). 

Notes: A common species in Europe, ranging to the Mediterranean area 
(Croatia, Spain, Tunisia), Kazakhstan, Asian Russia and Japan. The larvae feed as leaf 
miner on Carex humilis. New for Switzerland. 

*Napomyza bellidis Griffiths, 1967 

Material examined: GE: Chancy, La Laire, 350 m a.s.L, 1 S, 18.iv.2003, 
B. Merz leg. (MHNG). 

Notes: This species is based on a type series reared from Bellis perennis in 
England and later recorded from further European countries (Austria, Belgium, 
England, Estonia, France, Germany, Hungary, Italy, Latvia, Lithuania, Maltese Islands, 
Portugal, Russia, Sweden and Ukraine) and Central and East Palaearctic (Uzbekistan, 
Japan). New for Switzerland. 

Napomyza cichorii Spencer, 1966 

Reference: Martinez, 1998: 271. 

Material examined: VS: Saas-Fee, 2400 m a.s.L, 1 S , 22.vii.1965, 0. Lomholdt 
leg. (ZMUC). 

Notes: This species was described from a pair reared from Cichorium intybus 
originating from the Netherlands and one male from an alpine meadow in Switzerland 
(VS: Orsières). Our male confirms the occurrence in the canton Valais. It is distributed 
mainly in temperate Europe, the Mediterranean region and penetrates to Armenia, East 
Siberia, Far East and Japan. The larvae feed in stems and roots of Cichorium intybus 
and C. endivia (Asteraceae). 

*Napomyza elegans (Meigen, 1830) 

Material examined: GR: Valbella, 1500 m a.s.L, 1 9 , 28.vii.1999, leg. B. Merz 
& Müller (MHNG). 

Notes: A European species which is more common in temperate and northern 
Europe. Host plants and early stages are unknown but flies are not infrequently caught 
on flowers of Valeriana officinalis and it is believed the larva may feed in the root of 
this plant. New for Switzerland. 

*Napomyza maritima von Tschirnhaus, 1981 

Material examined: TI: Biasca, 350 m a.s.L, 1 ô , 18.V.1991, B. Merz leg. 
(MHNG). 



AGROMYZIDAE IN SWITZERLAND 793 

Notes: This species was described from Germany and France and it is also 
recorded from the Czech Republic, Finland, Hungary, Kazakhstan, Russia, Slovenia 
and Uzbekistan. The larva feeds in stems of Artemisia maritima. This plant does not 
occur in Switzerland. Other species of Artemisia, like A. campestris and A. vulgaris, 
both common in Biasca, may be infested by this species. New for Switzerland. 

*Napomyza nigriceps van der Wulp, 1871 

Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 S , l.iv.2002, 1 S , 
18.iv.2003, B. Merz leg. (MHNG). 

Notes: The species is known from Western, Central and Northern Europe, but it 
is not recorded in the Mediterranean area. Its biology is unknown. New for 
Switzerland. 

*Napomyza tripolii Spencer, 1966 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 6, 21-31.viii.2000, 
pasture, MT, B. Merz leg. (CMB). 

Notes: The type series of this species was caught on Aster tripolium in England. 
Later records includes localities in Denmark, Germany, Ireland and Spain. The larva 
feeds in the stem of Aster tripolium. This plant does not occur in Switzerland. On the 
other hand, A. alpinus is a common plant in the collecting and may be host of N. 
tripolii. New for Switzerland. 

Phytoliriomyza arctica (Lundbeck, 1901) 

Reference: Martinez, 1998: 271. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 ó*, viii.2000, pasture, 
MT, B. Merz leg. (CMB); Rothenbrunnen, 600 m a.s.l., 1 o\ 25 .xi. 1992. VS: 
Baltschieder, 670 m a.s.l., 1 9 , 12.V.1991; Branson, Follatéres, 450-950 m a.s.l., 1 S , 
29.iii.2002; Leuk Pfynwald, 650 m a.s.l., 1 6 , 2.V.1997, B. Merz leg. (MHNG). 

Notes: Almost cosmopolitan species but not known from the Afrotropical 
Region; common in Europe. Early stages unknown, but once reported as stem miner 
on Sonchus, probably also on other Asteraceae. 

Phytoliriomyza melampyga (Loew, 1 869) 

Reference: Martinez, 1998: 271. 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 S, 25.viii.2001, 
B. Merz & Landry leg. (MHNG). 

Notes: This Holarctic species is common in temperate and northern Europe and 
also recorded from North America and Oriental Region (India). The larva forms a leaf 
mine on Impatiens particularly on /. noli-tangere and I. parviflora. 

* Phytoliriomyza perpusilla (Meigen, 1830) 

Material examined: VS: Branson, Follatéres, 450-950 m a.s.l., 1 â , 29.iii.2002, 
B. Merz leg. (MHNG). 

Notes: P. perpusilla is distributed everywhere in Europe including the 
Mediterranean area and penetrates to the Afrotropical Region (Cape Verde Is., Lesotho, 



794 M. CERNY 

South Africa). Host plant unknown but probably several genera of Asteraceae 
(Spencer, 1976). New for Switzerland. 

Phytomyza affinis Fallen, 1823 

References: Hendel, 1931-6: 334; Martinez, 1998: 271. 

Material examined: GR: Valbella Casoja, 1500 m a.s.l., 1 6, 19.vii.2000, 
B. Merz leg. (MHNG). 

Notes: A species known to occur in the West Palaearctic, in particular in 
temperate and northern Europe but also recorded in the Mediterranean area and the Far 
East (Kuril Islands). The larva feeds in seed-heads on Euphrasia. 

*Phytomyza albipennis Fallen, 1823 

Material examined: GR: S. Vittore, Monacello, 280 m a.s.l., 2 SS, 8.iv.l997, 
B. Merz leg. (MHNG). 

Notes: This species is recorded from Europe including the Mediterranean area 
(Croatia, Canary Islands, Italy, Spain and the former Yugoslavia). Host plant unknown 
but some specimens have been caught on Ranunculus and the larvae possibly feed as 
an internal stem-borer (Spencer, 1972b, 1990). New for Switzerland. 

Phytomyza angelicae Kaltenbach, 1872 

Reference: Martinez, 1998: 271. 

Material examined: JU: Bonfol, 450 m a.s.l., 1 cT, 30.iv.1990, B. Merz leg. 
(MHNG). 

Notes: A Holarctic species which is known from temperate and northern Europe 
(not recorded from southern Europe), Kazakhstan, Turkey and North America. The 
larva forms an upper surface blotch leaf mine on Angelica silve stris, A. archangelica, 
Laserpitium latifolium, infrequently also on Aegopodium podagraria. 

Phytomyza artemisivora Spencer, 1971 

Reference: Martinez, 1998: 271. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 S , viii.2000, pasture, 
MT, B. Merz leg. (CMB). 

Notes: This species was described from England, Denmark and Germany. 
P. artemisivora is distributed in the Palaearctic Region from temperate and northern 
Europe to Kyrgysztan and Japan. Not confirmed in the Mediterranean area. The larva 
forms a white linear leaf mine on Artemisia vulgaris. 

Phytomyza brischkei Hendel, 1922 

References: Hendel, 1931-6: 364; Martinez, 1998: 271. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 25 S S , viii.2000, 2 S S , 
2 1.-31. viii.2000, pasture, MT, B. Merz leg. (CMB); Lenzerheide, Sundroina, 1520 m 
a.s.L, 1 S , 17.V.1997, B. Merz leg. (MHNG). VS: Saas-Fee, 1800-2100 m a.s.l., 2 S S, 
26. vii. 1965, O. Lomholdt leg. (ZMUC). 

Notes: The species is only known from temperate Europe, no records are 
available from northern and southern Europe. Host plants are species of Trifolium, 
particularly T. repens, T. pratense, T.fragiferum and T. alpinum. 



AGROMYZIDAE IN SWITZERLAND 795 

*Phytomyza calthivora Hendel, 1934 

Material examined: GE: Versoix, Bois du Faisan, 400 m a.s.l., 1 6 , 6.iv.2002, 
B. Merz leg. (MHNG). 

Notes: A rare species of temperate and northern Europe which can be found at 
localities where Caltha palustris, the host plant, occurs. The larva forms a short and 
broad leaf mine. New for Switzerland. 

*Phytomyza calthophila Hering, 1931 

Material examined: GE: Versoix, Bois du Faisan, 400 m a.s.l., 1 â , 6.iv.2002, 
B. Merz leg. (MHNG). 

Notes: P. calthophila is more common in temperate and northern Europe than 
the preceding species, though both species may occur together at same sites on Caltha 
palustris. Compared with P. calthivora the leaf mine formed by the larva is long and 
narrow. New for Switzerland. 

Phytomyza chaerophylii Kaltenbach, 1856 

Reference: Martinez, 1998: 271. 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 8, 25.viii.2001, 
B. Merz & Landry leg. (MHNG). 

Notes: This species is known from Europe including the Mediterranean area and 
also recorded from Turkey and Japan. The larva forms a linear leaf mine on 
Chaerophyllum, Anthriscus sylvestris, less frequnetly on Conium maculatum, 
Conopodium majus, probably also on Carum, Daucus, Sison and Torilis. 

Phytomyza contìnua Hendel, 1920 

Reference: Martinez, 1998: 271. 

Material examined: ZH: Zürich, 420 m a.s.l., 1 9, 30.vi.1989, B. Merz leg. 
(MHNG). 

Notes: Very common in Europe and also recorded from China, Kamchatka and 
Japan. The larva feeds along the midrib of leaves of Cirsium and Carduus. 

*Phytomyza crassiseta Zetterstedt, 1860 

Material examined: GE: Bernex, Signal, 510 m a.s.l., 1 6 , 8.VÜ.2001, B. Merz 
leg. (MHNG). 

Notes: P. crassiseta has a typically dilated arista. This Holarctic species is 
common in Europe and also recorded from Russia (including the Asian part), Kuril 
Islands, Japan and the United States. The larva forms a linear mine, initially adjoining 
leaf margin on Veronica. New for Switzerland. 

*Phytomyza eumorpha Frey, 1946 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 S, 21.-31.vii.2000. 
pasture, MT, B. Merz leg. (CMB). 

Notes: This species was described from Finland and Spencer (1976) recorded it 
later from Sweden. Our locality is the first one in Central Europe. The biology of 
P. eumorpha is unknown. New for Switzerland. 



796 M. CERNY 

*Phytomyza evanescens Hendel, 1920 

Material examined: ZH: Zürich Oerlikon, 430 m a.s.L, 1 â , 23 . v. 1 992, B. Merz 
leg. (MHNG). 

Notes: Zlobin (1994) reclassified this species and placed it in Phytomyza where 
it belongs to the albipenni s-group. P. evanescens is characterized by a flap-like 
appendage on the epandrium. This Holarctic species is distributed in Europe from 
Island to the Mediterranean area (Dalmatia, Sicily and Spain), and also recorded from 
Tajikistan, Kuril Islands and North America. The larva feeds internally in the stem of 
Ranunculus acris and R. lanuginosus. New for Switzerland. 

Phytomyza fallaciosa Brischke, 1880 

Reference: Martinez, 1998: 271. 

Material examined: GR: Lenzerheide, 2000 m a.s.L, 1 â, viii.2000, pasture, 
MT, B. Merz leg. (CMB). 

Notes: P. fallaciosa was described from Poland. Its distribution area includes 
Europe and the easternmost Palaearctic (Kuril Islands). The larva forms a leaf mine on 
Ranunculus , particularly on R. repens, but also on R. lingua. 

* Phytomyza flavicornis Fallen, 1823 

Material examined: GE: Chancy, La Laire, 350 m a.s.L, 1 6 , 18.iv.2003. JU: 
Lucelle, 560 m a.s.L, 1 9 , 30.iv.1990. All B. Merz leg. (MHNG). 

Notes: This Holarctic species is common in Europe and North America. The 
larva feeds as an internal stem borer on Urtica dioica. New for Switzerland. 

Phytomyza glechomae Kaltenbach, 1862 

Reference: Martinez, 1998: 271. 

Material examined: GE: Chancy, La Laire, 350 m a.s.L, 2 â â, 18.iv.2003; 
Chancy Vers Vaux, 335 m a.s.L, 1 6 , l.vi.2002, B. Merz leg. (MHNG). 

Notes: P. glechomae is common in Europe, Japan and North America. The larva 
forms a distinctive leaf mine on Glechoma hederacea. 

* Phytomyza gymnostoma Loew, 1858 

Material examined: SZ: Brunni, 900 m a.s.L, 1 9 , 26 vi. 1990, B. Petersen leg. 
(ZMUC). VS: Leuk Pfynwald, 630 m a.s.L, 1 6, 21.iv.1998, B. Merz & Botta leg. 
(MHNG); Saas-Fee, 2000-2200 m a.s.L, 1 9 , 21.vii.1965, O. Lomholdt leg. (ZMUC). 

Notes: The species was described from Poland and later recorded from further 
European countries, Turkey and Turkmenistan. Recently, this large species is 
considered to be a pest on Allium sp. with a certain economic importance (Spencer, 
1973). New for Switzerland. 

*Phytomyza heracleana Hering, 1937 

Material examined: BE: Berner Alpen, Grindelwald env., 2500 m a.s.L, 1 â , 
1 9 , 16.vi.1997, B. Mocek leg. (CBM). 

Notes: P. heracleana was described from Germany and it is distributed mainly 
in countries of temperate Europe. Rarely it is also recorded from the Mediterranean 



AGROMYZIDAE IN SWITZERLAND 797 

area (Bulgaria and Italy). The larva forms a regular, interparenchymal blotch leaf mine 
on Heracleum sphondylium, H. sibiricum, and H. mante gazzianum. New for 
Switzerland. 

*Phytomyza hirsuta Spencer, 1976 

Material examined: GR: Lenzerheide, Piz Danis, 2250-2490 m a.s.l., 1 a, 
12. vii. 1996, B. Merz leg. (MHNG). 

Notes: This species was described from Finland and Norway. Later it was also 
recorded from Sweden. Recently this species was found in the High Tatra Mts in 
Slovakia (Cerny & Vaia, 2005). Our record represents the southernmost locality in 
Europe. New for Switzerland. 

*Phytomyza homogyneae Hering, 1927 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 S, 21-31. viii.2000, 
pasture, MT, B. Merz leg. (CMB). 

Notes: This mountain species was described from Germany, where it was 
reared from Homogyne alpina, where the larva forms a linear mine. The species is 
known from Central European mountains (recently found in the Sumava Mts, Czech 
Republic, Cerny & Vaia, 2005) but also from China, Japan and Taiwan. New for 
Switzerland. 

Phytomyza nigripennis Fallen, 1823 

References: Hendel, 1931-6: 439; Martinez, 1998: 272. 

Material examined: GE: Chancy, La Laire, 350 m a.s.l., 1 â, 24.iv.2003, 
B. Merz & F. Amiet leg. (MHNG). ZH: Zürich Ahmend, 450 m a.s.l., 1 9 , l.v.1995, 
B. Merz leg. (MHNG); Zürich Waldgarten, 1 S , 31 .hi. 1997, B. Merz leg. (MHNG). 

Notes: This species is characterized by conspicuously dark wings. It is 
distributed in temperate and northern Europe, with a few localities in Northern Italy 
and former Yugoslavia. Its host plant is not known, but it may attack Anemone 
nemorosa. The larva feeds probably in the stem or the root, early stages unknown 
(Spencer, 1976). 

Phytomyza notata Meigen, 1830 

Reference: Martinez, 1998: 272. 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 8 6 S , viii.2000, pasture, 
MT,B. Merz leg. (CMB). 

Notes: This species is known from temperate and northern Europe including 
northern Italy. The larva forms a short, broad, linear mine recorded on Ranunculus 
acer, R. auricomus, R. bulbosa and particularly on R. repens. 

*Phytomyza origani Hering, 1931 

Material examined: GE: Cartigny, Moulin de Vert, 470 m a.s.l., 1 â , 4.vi.2001 , 
M. Eggenberger & B. Merz leg. (MHNG). 

Notes: P. origani was described from Germany and later recorded from 
Bulgaria, the Czech Republic, Denmark, England, France, Hungary, Lithuania, Poland 



798 M. CERNY 

and Spain. The larva forms a typical leaf mine on Origanum vulgare (Spencer, 1976). 
New for Switzerland. 

* Phytomyza pauliloewii Hendel, 1920 

Material examined: VS: Branson, Follatères, 450-950 m a.s.L, 1 6 , 29.iii.2002, 
B. Merz leg. (MHNG). 

Notes: This species is only locally distributed in temperate Europe but also 
known from northern Europe (Finland and Sweden) and the Mediterranean area (Italy). 
The larva forms a small regular blotch leaf mine on Pimpinella spp. and Peucedanum 
oreoselinum. New for Switzerland. 

Phytomyza plantaginis Robineau-Desvoidy, 1851 

Reference: Martinez, 1998: 272. 

Material examined: VS: Leuk Pfynwald, 650 m a.s.L, 1 S , 23 .ix. 1992, B. Merz 
& Otto leg. (MHNG), 630 m a. s. 1., 1 S, 6.viii.l997, B. Merz leg. (MHNG). 

Notes: P. plantaginis is a common species of the West Palaerctic and North 
America (Canada, United States). Rarely also known from the East Palaearctic, 
Afrotropical, Oriental and Australasian Regions. The larva forms a narrow, white linear 
leaf mine on Plantago lanceolata and P. major. 

*Phytomyza platystoma (Hendel, 1920) 

Material examined: VS: Branson, Follatères, 450-950 m a.s.L, 3 SS, 
29.iii.2002, B. Merz leg. (MHNG). 

Notes: This alpine species was described as Napomyza platystoma but Spencer 
& Martinez (1987) transferred it to Phytomyza. It was described from Austria 
(Piestingtale, Nördliche Voralpen). This is the second record from Central Europe. Its 
biology is unknown. New for Switzerland. 

* Phytomyza pubic omis Hendel, 1920 

Material examined: GE: Avusy, Moulin de la Grave, 360 m a.s.L, 1 â, 
18.iv.2003, B. Merz leg. (MHNG). ZH: Zürich Ahmend, 440 m a.s.L, 1 S , 17.iv.1996, 
B. Merz leg. (MHNG). 

Notes: This species is distributed in temperate and northern Europe. No records 
are known from the Mediterranean area. The larva forms a short linear leaf mine on 
Aegopodium podagraria. New for Switzerland. 

* Phytomyza pullula Zetterstedt, 1848 

Material examined: ZH: Affoltern a. A., 600 m a.s.L, 1 â , 8.viii.l987, B. Merz 
leg. (MHNG). 

Notes: P. pullula belongs to the species with a Holarctic distribution. It is 
common in Europe and North America. The larva forms a narrow linear leaf mine on 
Achillea, Anthémis, Chrysanthemum vulgare, Matricaria, and most commonly on 
M. inodora. New for Switzerland. 

Phytomyza ranunculi (Schrank, 1803) 

References: Hendel, 1931-6: 467; Martinez, 1998: 272. 



AGROMYZIDAE IN SWITZERLAND 799 

Material examined: GR: Au s serf errera, 1300-1550 m a.s.l., 1 6 , 18.vi.1994, B. 
Merz leg. (MHNG); Lenzerheide, 2000 m a.s.l., 1 6 , 14. -21. vii .2000, pasture, MT, B. 
Merz leg. (CMB); Lenzerheide See, 1500 m a.s.l., 1 6, 15.vii.2000, B. Merz leg. 
(MHNG); Lenzerheide Sundroina, 1520 m a.s.l., 1 o\ 17.V.1997, B. Merz leg. 
(MHNG). TI: Monte San Giorgio, 600-1100 m a.s.l., 1 6, 5.vii.2001, B. Merz leg. 
(MHNG). ZH: Zürich Zürichberg, 600 m a.s.l., 1 S , 5.V.1993, B. Merz leg. (MHNG), 
1 6 , 8.V.1996, P. Walser leg. (MHNG). VD: Bonvillars en fauchant [= sweeping], 
630 m a.s.L, 1 9 , 25.iv.1988, D. Burckhardt leg. (MHNG). 

Notes: P. ranunculi is a common species mining Ranunculaceae in the Holarctic 
Region. 

*Phytomyza ranunculicola Hering, 1949 

Material examined: GE: Bernex, Chante-Merle, 415 m a.s.l., 1 ó\ 20.V.2002, B. 
Merz leg. (MHNG). 

Notes: The species was described from Germany. It is distributed in temperate 
Europe but it is probably absent from Scandinavia and the Mediterranean area. The 
larva forms a secondary blotch leaf mine on Ranunculus acer. New for Switzerland. 

*Phytomyza rapunculi Hendel, 1927 

Material examined: BE: Berner Alpen, Grindelwald env., 2500 m a.s.l., 1 â , 
1 9, 16.vi.1997, B. Mocek leg. (CBM). GR: Lenzerheide, 2000 m a.s.l., 1 o\ 
viri .2000, pasture, MT, B. Merz leg. (CMB). 

Notes: This European species was described from Austria and later recorded 
particularly in temperate and southern Europe. In Scandinavia it is known only from 
Sweden (Spencer, 1990). The larva forms a linear mine on Campanula rapunculoides, 
C . persici/olia and Phyteuma. New for Switzerland. 

*Phytomyza rhabdophora Griffiths, 1964 

Material examined: GR: Lenzerheide, 2000 m a.s.l., 1 â , viii.2000, pasture, 
MT, B. Merz leg. (CMB). Lenzerheide, Sundroina, 1550 m a.s.l., 1 o\ 18.vii.1997, 
B. Merz leg. (MHNG). VS: Saas-Fee, 1800 m a.s.l., 1 6 , 21 .vii. 1965, 0. Lomholdt leg. 
(ZMUC). 

Notes: The type locality is in Germany and the species is distributed in 
temperate Europe, but it is apparently missing in the Mediterranean area. Its biology is 
unknown, larvae feed probably on Leontodon (Tschirnhaus, 1969). New for 
Switzerland. 

*Phytomyza rostrata Hering, 1933 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 â , 2.vii.2001 , B. Merz 
leg. (MHNG). 

Notes: The species was described from Germany and it is distributed in 
temperate Europe, but apparently absent from the Mediterranean area. The larva forms 
a typical mine on Euphrasia, Melampyrum, Odontites and Rhinanthus (Spencer, 1976). 
New for Switzerland. 

Phytomyza rufipes Meigen 1830 

Reference: Martinez, 1998: 272. 



800 M. CERN Y 

Material examined: GE: Cartigny, Moulin de Vert, 360 m a.s.L, 1 S , l.v.1999, 
B. Merz leg. (MHNG). 

Notes: This pale species is common in the West Palaearctic and North America. 
The larva feeds inside the stem or midrib of larger leaves on Brassicaceae, mainly on 
Brassica. 

*Phytomyza sedi Kaltenbach, 1869 

Material examined: NE: St. Blaise, Les Riedes, 470 m a.s.L, 2 â 6 , 19.V.2001, 
B. Merz leg. (MHNG). 

Notes: This species was described from Boppard (Germany) and later recorded 
from Spain under the synonym Ph . catalaunica Spencer, 1960. Our record confirms the 
occurrence in Central Europe. Sedum is the host plant of this species. New for 
Switzerland. 

* Phytomyza soenderupi Hering, 1941 

Material examined: GE: Versoix, Bois du Faisan, 400 m a.s.L, 1 S , 6.iv.2002, 
B. Merz leg. (MHNG). 

Notes: This species was described from Denmark and later recorded from 
Belgium, the British Isles, the Czech Republic, Estonia, Germany and Norway. The 
larva feeds within the petiole of Caltha palustris. New for Switzerland. 

Phytomyza tenella Meigen, 1830 

Reference: Martinez, 1998: 272. 

Material examined: GR: Lenzerheide, 2000 m a.s.L, pasture, 2 â â , 23.viiL- 
9.ix.2000, MT, B. Merz leg. (CMB); Juf-Stallerberg, 2400 m a.s.L, 1 S, 18.viii.1991, 
B. Merz & A. Freidberg leg. (MHNG). 

Notes: Phytomyza tenella is distributed in Europe, China, Turkey, the United 
States and the Afrotropical Region (Ethiopia). The larva feeds in seed-heads of 
Pedicularis palustris and almost certainly other Pedicularis. 

'"Phytomyza varipes Mac quart, 1835 

Material examined: GR: Lenzerheide, Sundroina, 1550 m a.s.L, 1 â, 
18. vii. 1997, B. Merz leg. (MHNG). 

Notes: This rare species was described from Northern France. The larva feeds 
in seed-heads of Rhinanthus. New for Switzerland. 

Phytomyza vitalbae Kaltenbach, 1872 

Reference: Martinez, 1998: 272. 

Material examined: GR: Lenzerheide, 2000 m a.s.L, 1 â, 14.-21. vii.2000, 
pasture, MT. B. Merz leg. (CMB). VS: Leuk Pfynwald, 600 m a.s.L, 2 Sa, 
25.viii.2001, B. Merz & Landry leg. (MHNG). 

Notes: A species known from various European countries and also from Canada, 
South Africa, China, Nepal. Taiwan and Australia. The larva forms a long, narrow, 
upper surface leaf mine on Clematis vitalba, C. alpina and some other Clematis. 

Phytomyza wahlgreni Rydén, 1944 

References: Griffiths, 1964: 411; Spencer, 1976: 527; Martinez, 1998: 272. 



AGROMYZIDAE IN SWITZERLAND 801 

Material examined: AG: Thalheim, 550 m a.s.l., 1 6 , 20.V.1992, B. Merz leg. 
(MHNG). GR: Lenzerheide, 1600 m a.s.l., 1 6 , 8.x. 1991, B. Merz & M. Eggenberger 
leg. (MHNG); Lenzerheide, 2000 m a.s.l., 3 66, 14. -21. vii .2000, 2 66, 23.- 
31.viii.2000, pasture, MT, B. Merz leg. (CMB). NE: St. Blaise, Les Riedes, 470 m 
a.s.l., 1 6, 19.V.2001, B. Merz leg. (MHNG). VS: Saas-Fee, 2000 m a.s.l., 3 6 6, 
18.vii.1965, 2400 m a.s.l., 1 9, 19.vii.1965, 2200 m a.s.l., 2 9 9, 21. vii. 1965, 1800 m 
a.s.l., 1 9 , 24 .vii. 1965, O. Lomholdt leg. (ZMUC). 

Notes: A Holarctic species distributed in temperate and northern Europe. In the 
Mediterranean area known to occur in Italy. It is also recorded from the United States, 
China and the Kuril Islands. From Switzerland recorded by Griffiths (1964) for the first 
time (VS: Bérisal, under the synonym P. taraxacocecis Hering, 1949). The larva feeds 
in the midrib of leaves on Taraxacum spp. 

*Pseudonapomyza atra (Meigen, 1830) 

Material examined: FR: Mt. Vully, Bas Vully, 460-650 m a.s.l., 1 6 , 6.vi.2003, 
B. Merz & Amiet leg. (MHNG). GE: Bernex, Chante-Merle, 415 m a.s.l., 1 6, 
16.viii.2002, B. Merz leg. (MHNG); Cartigny, Moulin de Vert, 360 m a.s.l., 1 6, 
2.vi.2002, B. Merz leg. (MHNG); Chancy, La Laire, 350 m a.s.l., 1 6 , 9.VÜ.2002, B. 
Merz leg. (MHNG). VS: Visperterminen, Giw-Gebidemsee, 1900-2200 m a.s.l., 1 6, 
28.viii.2001 , B. Merz & Landry leg. (MHNG). ZH: Zürich-Hönggerberg, 600 m a.s.l., 
1 6, 26 .iv. 1993, B. Merz leg. (MHNG). 

Notes: Ps. atra is generally distributed in the Holarctic Region. It is common in 
northern and temperate Europe (Cerny 1992, 1998, 2005a). Spencer (1973) considered 
it to be rare in the Mediterranean area. The larva forms a short, narrow mine on 
Gramineae, feeding on many genera, e.g. Apera, Avena, Hordeum, Lolium, Phalaris, 
Poa, Secale and Triticum. New for Switzerland. 

*Pseudonapomyza errata Zlobin, 1993 

Material examined: GE: Russin, Les Baillets, 405 m a.s.l., 1 6 , 30.vi.2001, 
B. Merz leg. (MHNG). NE: St. Blaise, Les Riedes, 470 m a.s.l., 2 66, 19.V.2001, 
B. Merz leg. (MHNG). VS: Leuk Pfynwald, 600 m a.s.l., 1 6, 2.VÜ.2001; 
Visperterminen, 1400 m a.s.l., 1 6, 27 .vii. 1991, B. Merz leg. (MHNG). 

Notes: This species was described from Russia, Uzbekistan and Mongolia and 
it is also recorded from the Czech Republic and France. Its biology is unknown. New 
for Switzerland. 

*Pseudonapomyza eurasiatica Zlobin, 2003 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 6 , 6.vi.2001, 1 6, 
7. vi. 200 1, B. Merz & Landry leg. (MHNG). 

Notes: Recently described from the male holotype originating from Tajikistan 
and paratypes from Greece, Kazakhstan, Turkmenistan and Ukraine by Zlobin 
(2003b). This is the first record from Central Europe and represents the westernmost 
boundary of its distribution area. Biology is unknown. New for Switzerland. 

Pseudonapomyza europaea Spencer, 1973 

References: Papp, 1984: 313; Spencer, 1973: 270; Martinez, 1998: 272. 



802 M. CERNY 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 S , 25.V.1997, B. Merz 
leg . (MHNG) , 2 S S , 6 .vi .200 1 ; 1 S, 2 .vii .200 1 ; 1 S , 25 .viri .200 1 , B . Merz & Landry 
leg. (MHNG). 

Notes: Widely distributed in Europe and also recorded from Turkey and Japan 
(Cerny, 2005a). It is less common than Ps. atra. Spencer (1973) described this species 
from Italy (holotype), Austria, Macedonia, Serbia, Slovenia and Switzerland (VS: 
Stalden). Its biology is unknown. 

*Pseudonapomyza strobliana Spencer, 1973 

Material examined: GE: Russin, Les Baillets, 405 m a.s.l., 1 S , 8.viii.2002, B. 
Merz leg. (MHNG). VS: Leuk Pfynwald, 600 m a.s.l., 1 6, 6.vi.2001, 1 S, 
25.viii.2001, B. Merz & Landry leg. (MHNG). 

Notes: The species was described from Dalmatia and Austria. The center of its 
distribution area is probably in the Mediterranean area (Spain, Bulgaria, Turkey) but 
recently it was also recorded from the Czech Republic, France, Germany, Hungary, 
Sweden and Ukraine (Cerny, 1998, 2005a). Its biology is unknown. New for 
Switzerland. 

*Pseudonapomyza vota Spencer, 1973 

Material examined: VS: Leuk Pfynwald, 600 m a.s.l., 1 â , 7.vi.2001; Leuk 
Platten, 625 m a.s.l., 1 6 , 8.vi.2001, all B. Merz & Landry leg. (MHNG). 

Notes: This species was described from a single male originating from Spain. 
The distribution area of this species ranges from Spain to Israel, i.e. throughout the 
Mediterranean area. Biology of the species is unknown. New for Switzerland. 

CONCLUSIONS 

This review embraces new data on the fauna of mining flies or Agromyzidae 
occurring in Switzerland and adds 92 species to the 140 species listed by Martinez 
(1998) bringing the list of Swiss agromyzids to 232 species. The potential number of 
Swiss Agromyzidae species is probably much higher, given the different origins of the 
native fauna and the unusally diverse pattern of ecosystems. Such a conclusion seems 
to be confirmed by a comparison of species richness in neighbouring countries. Franz 
(1989) listed 270 species from Austria and Tschirnhaus (1999) 552 species from 
Germany. According to Spencer (1992) many species known from the Italian alpine 
valleys will probably also be found in the Swiss Alps. Süss (1999, 2001, 2002) 
recorded 224 species in Italy (173 from North Italy); of this number 112 species are 
common for both countries and 112 species are known exclusively from Italy. So far, 
no species of Hexomyza Enderlein, 1936, has been recorded from Switzerland, 
although 6 species are known in Europe and 5 of them live in Italy. Likewise 
Ptochomyza Hering, 1942, with three European species and Nemorimyza posticata 
(Meigen, 1830) known from adjoining countries, have not been found so far in 
Switzerland. Of 10 European species of Phytobia Lioy, 1864, only P. mallochi is 
known from Switzerland, while Melanagromyza Hendel, 1920, is represented by four 
species only (11 % of the European species) and Liriomyza Mik, 1894, by 18 species 
(13 %). There is thus good evidence that the list of Swiss agromyzids is far from final. 



AGROMYZIDAE IN SWITZERLAND 803 



ACKNOWLEDGEMENTS 

My sincerest thanks go to Miroslav Bartâk (Praha), Rudolf Meier 
(Copenhagen), Bernhard Merz (Genève) and Bohuslav Mocek (Hradec Krâlové) for 
allowing me to use the collections in their care and Rudolf Rozkosny (Brno, Czech 
Republic) for invaluable support during preparation of the manuscript. 

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Revue suisse de Zoologie 112 (4): 807-830; décembre 2005 



The millipede family Paradoxosomatidae in Paraguay, with 
descriptions of five new species (Diplopoda, Polydesmida) 

Sergei I. GOLOVATCH 

Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, 
Leninsky pr. 33, Moscow 119071 (V-71), Russia. E-mail: sgol@orc.ru 

The millipede family Paradoxosomatidae in Paraguay, with descrip- 
tions of five new species (Diplopoda, Polydesmida). - A review is pro- 
vided of the millipede family Paradoxosomatidae in Paraguay, with keys 
compiled to 14 genera and 63 species occurring in this country and/or 
adjacent areas. Five species are described as new: Catharosoma bilineatum 
sp. n., C. mahnerti sp. n., Broelemannopus minutus sp. n., Mestosoma sim- 
plex sp. n. and M. crassipes sp. n. 

Keywords: Diplopoda - Polydesmida - Paradoxosomatidae - taxonomy - 
Paraguay. 

INTRODUCTION 

The family Paradoxosomatidae is among the largest among the Diplopoda, with 
nearly 200 genera currently accepted as valid. The American fauna is highly peculiar 
due to the absence of indigenous paradoxosomatids north of Costa Rica. In South 
America two major centres of diversification are distinguishable, one in the area of 
southern Brazil, Paraguay and northern Argentina, the other in Peru, northern Bolivia 
and possibly Ecuador. The vast regions of northern Brazil, Guiana, Venezuela and 
Colombia appear to harbour very few species (Jeekel, 1968, 2002). 

The Neotropical fauna itself is strongly dominated by relatively few (about a 
dozen) genera of the endemic tribe Catharosomatini. Of these genera, Mestosoma 
Silvestri, 1897 is certainly the largest and particularly widespread, with about 75 
species occurring throughout South America and reaching both Costa Rica and the 
island of Dominica (introduced?) in the north. Catharosoma Silvestri, 1913 is the 
second largest genus, its 13 species being known from Brazil, Paraguay and Argentina 
(Hoffman, 1980, 1999). The remaining genera are mono- to oligotypic. 

The present paper provides a review of, and keys to, the bulk of the paradoxo- 
somatids occurring in the southern Neotropical diversification centre, based both on all 
available literature sources and on the important collection of Paradoxosomatidae from 
Paraguay housed in the Muséum d'histoire naturelle, Geneva (MHNG). Five new 
species from three genera of Catharosomatini have been revealed there, all described 
below and thus considerably enriching our knowledge of the fauna of Paraguay. A few 
paratypes from the material have been retained for the collection of the Zoological 
Museum, State University of Moscow (ZMUM), Russia. 



Manuscript accepted 07.02.2005 



808 S.I. GOLOVATCH 



FAUNISTIC REVIEW 



Since detailed reviews of the history of research on Neotropical Paradoxo- 
somatidae are available and still fully relevant (Jeekel, 1963, 1968), only a short 
account appears necessary here concerning the species list of the particular region 
involved. Virtually all later contributions (e.g. Hoffman, 1977, 1999; Golovatch, 1992; 
Jeekel, 2002; Golovatch et al., 2003) are irrelevant in the present context, as they only 
or chiefly treat more northern faunas. 

The following Paradoxosomatidae from Paraguay and/or the adjacent parts of 
Bolivia, northern Argentina and southern Brazil have hitherto been recognized as valid: 

Broelemannopus Verhoeff, 1938 

Broelemannopus escaramucensis (Schubart, 1944) - Brazil (Säo Paulo) 

(Schubart, 1944, 1952) 

B. glabratus (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b) 
B. ibitiensis (Schubart, 1945) - Brazil (Säo Paulo) (Schubart, 1945a) 

B. pirassunungensis (Schubart, 1944) - Brazil (Sao Paulo) (Schubart, 1944, 
1945a, 1952) 

Catharosoma Silvestri, 1897 

Catharosoma curitibense Schubart, 1953 - Brazil (Parana) (Schubart, 1953) 

C. digitale Schubart, 1953 - Brazil (Parana) (Schubart, 1953) 
C. hoffmani Kraus, 1956 - Paraguay (Kraus, 1956) 

C. intermedium (Carl, 1902) - Brazil (Rio Grande do Sul) (Carl, 1902; Attems, 

1914, 1937) 

C. mesorphinum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 

1937; Schubart, 1953) 

C. mesoxanthum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 

1937; Schubart, 1953) 

C. mixtum Kraus, 1956 - Brazil (Santa Catarina) (Kraus, 1956) 

C. myrmekurum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 

1937; Schubart, 1953) 

C.palmatum Schubart, 1953 - Brazil (Parana) (Schubart, 1953) 

C. palustre Schubart, 1943 - Brazil (Mato Grosso) (Schubart, 1943) 

C. paraguayense (Silvestri, 1895) - Paraguay, Argentina and Brazil (Santa 

Catarina and Mato Grosso) (Silvestri, 1895b, 1902; Attems, 1898, 1914, 1937; 

Schubart, 1953; Kraus, 1956; Jeekel, 1965) («South America», according to Mauriès, 

1998) 

C.peraccae Silvestri, 1902 - Paraguay (Silvestri, 1902; Attems, 1914, 1937) 
C. taeniatum (Brolemann, 1929) - Brazil (Santa Catarina) (Brolemann, 1929; 

Attems, 1937; Schubart, 1953) 

Chondro morpha Silvestri, 1897 

Chondromorpha xanthotricha (Attems, 1898) - pantropical, introduced (e.g. 
Jeekel, 1963) 



PARADOXOSOMATIDAE IN PARAGUAY 809 

Gonodrepanoides Schubart, 1945 

Gonodrepanoides travassosi Schubart, 1945 - Brazil (Rio de Janeiro) 
(Schubart, 1945b) 

Gonodrepanum Attems, 1914 

Gonodrepanum drepanephoron (Attems, 1898) - Brazil (Säo Paulo) and 
Argentina (Attems, 1898, 1901, 1914, 1937; Brölemann, 1902b; Mauriès, 1998) 
G.falciferum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) 
G.flavolineatum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) 
G.furcatum Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) 
G. grajahuense Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) 
G. levisetum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 1898, 1914, 
1937; Schubart, 1953; Mauriès, 1998) 

G. levisetum var. coniferum (Attems, 1898) - Brazil (Santa Catarina) (Attems, 
1898, 1914, 1937; Schubart, 1953; Mauriès, 1998) 

G. torresae Schubart, 1945 - Brazil (Rio de Janeiro) (Schubart, 1945b) 

Habrodesmoides Attems, 1943 

Habrode smoide s costalimai (Schubart, 1945) - Brazil (Rio de Janeiro) 
(Schubart, 1945b) 

H. perturbans Attems, 1943 - Brazil (Rio de Janeiro) (Attems, 1943) 

Mestosoma Silvestri, 1897 

Mestosoma alticola (Attems, 1931) - Bolivia (Cochabamba) and Peru (Attems, 

1931, 1937; Kraus, 1956) 

M. balzami (Silvestri, 1895) - Bolivia (Yungas) (Silvestri, 1895a; Attems, 1914, 

1937) 

M. bicolor Silvestri, 1898 - Paraguay and Brazil (Mato Grosso) (Silvestri, 1898, 

1902; Attems, 1899, 1914, 1937; Jeekel, 1965) 

M. boliviae (Chamberlin, 1957) - Bolivia (near Potosi) (Chamberlin, 1957) 
M. borellii (Silvestri, 1895) - Argentina (Tucuman) and Paraguay (Silvestri, 

1895b; Attems, 1914, 1937; Mauriès, 1998) 

M. camer ani (Silvestri, 1895) - Argentina (Chaco) (Silvestri, 1895b; Attems, 

1914, 1937; Jeekel, 1965; Mauriès, 1998) 

M. carioca (Schubart, 1945) - Brazil (Rio de Janeiro) (Schubart, 1945b) 

M. derelictum (Silvestri, 1895) - Bolivia (Misiones Mosetenes) (Silvestri, 

1895a; Attems, 1914, 1937) 

M. differens Kraus, 1956 - Bolivia (Cochabamba) (Kraus, 1956) 

M. femorale (Schubart, 1943) - Brazil (Mato Grosso) (Schubart, 1943) 

M. kalliston (Attems, 1898) - Brazil (Rio Grande do Sul) (Attems, 1898, 1914, 

1937) 

M. luctuosum Silvestri, 1897 - Bolivia (Caiza) (Silvestri, 1897; Attems, 1914, 

1937) 



810 S.I. GOLOVATCH 

M. lugubre Silvestri, 1897 - Argentina (Buenos Aires) and Uruguay (Silvestri, 

1897, 1902; Attems, 1914, 1937; Mauriès, 1998) 

M. montanum (Silvestri, 1895) - Bolivia (Yungas) (Silvestri, 1895a; Attems, 

1914, 1937) 

M.perfidum (Schubart, 1943) - Brazil (Sào Paulo) (Schubart, 1943) 

M. pseudomorphum (Silvestri, 1895) - Paraguay (Silvestri, 1895b; Cari, 1902; 

Attems, 1914, 1937; Jeekel, 1965) 

M.pulvillatum (Attems, 1898) - Paraguay (Attems, 1898, 1901, 1914, 1937) 
M. salvadorii (Silvestri, 1895) - Argentina (Salta), Bolivia (Chaco) and 

Paraguay (Silvestri, 1895b, 1902; Attems, 1914, 1937; Hoffman, 1977; also Costa 

Rica, according to Mauriès, 1998) 

M. schindlerì Kraus, 1956 - Bolivia (Silhuencas) (Kraus, 1956) 

M. tricuspis (Verhoeff, 1938) - Paraguay (Verhoeff, 1938) 

M. truncatum (Schubart, 1943) - Brazil (Mato Grosso) (Schubart, 1943) 

M. vittatum (Attems, 1898) - Paraguay (Attems, 1898, 1914, 1937) 

Mogyella Schubart, 1944 

Mogyella nana Schubart, 1944 - Brazil (Säo Paulo) (Schubart, 1944, 1952) 

Mogyosoma Schubart, 1944 

Mogyosoma hamatum Schubart, 1944 - Brazil (Sào Paulo) (Schubart, 1944) 

Ologonosoma Silvestri, 1898 

Ologonosoma iguassuense (Schubart, 1953) - Brazil (Parana) (Schubart, 1953; 
Mauriès, 1998) 

O. sanctum (Silvestri, 1895) - Paraguay (Silvestri, 1895a; Attems, 1914, 1937; 
Mauriès, 1998) 

Orthomorpha Bollman, 1893 

Orthomorpha coarctata (De Saussure. 1860) - pantropical, introduced (e.g. 
Jeekel, 1963) 

Oxidus Cook, 1911 

Oxidus gracilis (C. L. Koch, 1847) - subcosmopolitan, introduced (e.g. Jeekel, 
1963) 

Promestosoma Silvestri, 1898 

Promestosoma boggianii Silvestri, 1898 - Paraguay (Silvestri, 1898; Attems, 
1899, 1914, 1937; Jeekel, 1965; Mauriès, 1998) and Brazil (Mato Grosso and Mato 
Grosso do Sul) (original data) 

Pseudogonodrepanum Schubart. 1945 

Pseudogonodrepanum scitum Schubart, 1945 - Brazil (Rio de Janeiro) 
(Schubart, 1945b) 



PARADOXOSOMATIDAE IN PARAGUAY 811 

A few more species, i.e. Mestosoma laetum Silvestri, 1897 (Bolivia, San 
Francisco - Silvestri, 1897), «Strongylosoma» nitidum Brölemann, 1902 (Brazil, 
Cubatäo, Säo Paulo - Brölemann, 1902a), «Strongylosoma» pustulatum Brölemann, 
1902 (Brazil, Säo Paulo - Brölemann, 1902b) and Catharosoma bromelicola Schubart, 
1945 (Brazil, Rio de Janeiro - Schubart, 1945b), the male characters of which are not 
or are insufficiently known, still remain incertae sedis; none can be included in a key 
(Jeekel, 1963). 

The same concerns Iulidesmus Silvestri, 1895, with its type-species /. typicus 
Silvestri, 1895 known only from a female holotype from Bolivia (Yungas) (Silvestri, 
1895a). Jeekel (1963) correctly excluded Iulidesmus from Paradoxosomatidae on the 
basis of a later redescription of /. typicus from Chilean material provided by Silvestri 
(1905). But Hoffman (1980), rightly doubting the conspecificity of the Bolivian and 
Chilean samples, believes that the holotype of/, typicus represents a Mestosoma. If so, 
then Mestosoma becomes a junior synonym of Iulidesmus . Only topotypic material in 
connection with revisionary work can shed additional light on the identity of all of the 
enigmatic taxa mentioned above. 

DESCRIPTIONS OF NEW SPECIES 

Catharosoma bilineatum sp. n. Figs 1-9 

Material: Holotype S (MHNG), Paraguay, Alto Parana Prov., Forestry Centre 
(C.F.A.P.), Puerto Presidente Stroessner, plot 7, 18.03.1983, leg. P. Berner & C. Dlouhy. - 
Paratypes: 1 9 (MHNG), same locality, together with holotype; 1 ó\3 9 9 (MHNG), 1 6,1 $ 
(ZMUM), same locality, around Pindo trunk, plot 3, 24.03.1983, leg. C. Dlouhy; 1 9 (MHNG), 
same locality, soil sample, plot 4 (Monte natural), Winkler extraction, 10.03.1983, leg. C. 
Dlouhy; 1 9 (MHNG), same plot 4 (Monte natural), 10.03.1983, leg. P. Berner & C. Dlouhy; 1 
9 (MHNG), same locality, soil sample, plot 2 (Monte natural), 29.02.1983, leg. C. Dlouhy; 1 6 
(MHNG), same locality, soil sample, plot 8 (Monte natural), 18.03.1983, leg. P. Berner & C. 
Dlouhy. 

Name: To emphasize the presence of two dark paramedian stripes divided by a similar- 
ly wide but light axial stripe. 

Diagnosis: Differs from congeners by the characteristic coloration, the strong- 
ly reduced paraterga visible only on segments 2-4, the presence of relatively short an- 
tennae, the deeply divided sternal lobe present between coxae 6 of the S , the subunci- 
form tip of the hypoproct, the bifid tip of the epiproct, coupled with certain details of 
solenophore structure. 

Description: Length 21-27 mm (6 , 9), width of midbody metazona 1.9-2.1 
(6) or 2.1-2.8 mm ( 9 ); 9 usually larger and somewhat broader than 6 . Holotype ca 
23 mm long and 2.0 mm wide. Coloration in alcohol pale yellowish brown to brown, 
with a characteristic pair of brown to dark brown paramedian stripes divided by an 
equally wide axial stripe of background coloration; legs and sterna yellowish gray, 
slightly paler than background coloration; tip of antennae pallid, penultimate anten- 
nomere dark brown; posterior half of body always somewhat paler than anterior one. 

Body subcylindrical, not moniliform. Postcollar constriction faint, width of 
head = 5-16 > collum = 4 > 2 = 3; on segments 17-20 trunk gradually and gently ta- 
pering toward telson both in width and in height. Antennae relatively short, slightly 
davate, in situ reaching beyond segment 2 dorsally (Fig. 1), a little shorter in 9. 



812 



S. I. GOLOVATCH 




Figs 1-9 
Catharosoma bilineatum sp. n.. S holotype: 1) anterior body portion, lateral view; 2) telson, lat- 
eral view; 3) sternal structures between coxae 4-7 (coxae 4 on top); 4) sternal structures between 
midbody coxae; 5) leg 12; 6-9) left gonopod, medial, ventral, lateral and ventromedial views, re- 
spectively. Scale bars 1.0 (1-5) and 0.25 mm (6-9). 



Paraterga strongly reduced, only on segment 2 fully developed as low keels with a 
caudal tooth, discernible on segments 3 and 4 as arcuated sulci (Fig. 1), on following 
segments totally missing. Tegument smooth, shining, only rear parts of metazona often 
faintly rugulose. Limbus thin, caudal margin entire. Metaterga fully devoid of a trans- 
verse sulcus; setae medium- sized, often abraded, arranged 2+2 in a transverse line 
behind a shallow stricture between pro- and metazona. Ozopores lateral, opening level 
to metatergal surface ca 1/3 metazonital length away from caudal edge. Pleurosternal 
carinae shaped like narrow keels with a caudal spinule (Fig. 1) traceable until segment 
16 ($) or 17 (cT), on following segments wanting. Epiproct (Fig. 2) rather long, coni- 
form, bifid, in â a little longer than in 9 . Hypoproct (Fig. 2) subtriangular, tip pointed, 
unciform, directed ventrad; 1+1 paramedian setae at caudal edge considerably 
separated from each other. Paraterga. pleurosternal carinae and hypoproct uncus in 6 
a little more strongly developed than in 9 . 

Sterna densely setose. Sternum between coxae 3 of 6 with a paramedian, nearly 
contiguous pair of bunches of setae. Sternum between coxae 4 of cT with a prominent, 
linguiform, laterally setose outgrowth directed antero ventrad (Fig. 3); a similar but 



PARADOXOSOMATIDAE IN PARAGUAY 813 

much lower outgrowth between coxae 5 of â (Fig. 3); a more or less prominent and 
deeply divided outgrowth between coxae 6 of â poorly delimited caudally against a 
similar but less prominent bulge with peculiar paramedian bunches of setae between 
coxae 7 of 6 (Fig. 3). Postgonopodial sterna with shorter (between anterior coxae) or 
longer (between posterior coxae), coniform, paramedian spines developed a little bet- 
ter in 6 (Fig. 4) than in 9 . 

Legs without tarsal brushes but densely setose ventrally; setation gradually 
thinning out toward telson; tibiae swollen ventrally and supporting characteristic 
brushes (Fig. 5) between leg-pair 9 until two last pairs. Legs in 6 a little longer than 
in 9 , as usual becoming a little longer and slenderer toward telson. Each coxa 2 of â 
with a prominent, somewhat sinuate, distoventral spine carrying a gonopore at base. 

Gonopods (Figs 6-9) highly complex. Coxite elongate, subcylindrical, setose 
disto ventrally; cannula normal. Telopodite strongly unciform, with a hypertrophied, as 
usual densely setose pre femoral part somewhat longer than femorite. A few undu- 
lations on ventrocaudal face and a few longitudinal ridges anterodorsally at base of 
solenophore. Solenophore coiled, ventrally with both a slightly folded base of lamina 
medialis (m) and a subtriangular parabasal lobe (p) similarly well-developed; p 
followed by a spine (k) supporting the tip of a flagelliform solenomere at base of a 
hyaline lobule (h), this lobule supporting the penultimate 1/4 extent of both lamina 
lateralis and lamina medialis; distalmost 1/4 extent of both laminae slender, unciform, 
free of support. 

Remarks: Judging from the colour pattern and gonopod structure, this new 
species seems to be especially close to C. mesoxanthum, but both differ in size, in 
structure of the process between coxae 6 of the â , and in direction of the apical 
papillae on the epiproct (see also key below). 

Catharosoma mahnerti sp. n. Figs 10-17 

Material: Holotype 6 (MHNG), Paraguay, Prov. Concepción, Estancia Laguna Negra, 
15 km E of Paso Barreto, dead wood, 13.10.1985, leg. Expédition zoologique du Muséum de 
Genève. - Paratypes: 1 6,2 9 9 (MHNG), 1 6, 1 $ (ZMUM), same locality, together with 
holotype; 1 6 (MHNG), Prov. Concepción, near mouth of Ao River, Trementina (Estancia 
Laguna Negra), forest, 13.10.1985, leg. Expédition zoologique du Muséum de Genève. 

Name: Honours Dr Volker Mahnert, who entrusted me this valuable material for study. 

Diagnosis: Differs from congeners by the submoniliform body, the longer 
antennae, the absence of a distinct colour pattern and of an unciform hypoproct, the 
presence of two paramedian pilose ridges between coxae 3 of the 6 , and in certain 
details of gonopod structure. 

Description: Length 20-23 mm (â , 9 ), width of midbody metazona 1.7 (â) to 
2.3-2.5 mm ( 9 ); 9 usually larger and somewhat broader than 6 . Holotype ca 20 mm 
long and 1 .7 mm wide. Coloration in alcohol uniformly light brown to red-brown; only 
penultimate antennomere rather dark brown, tip of antenna pallid; legs paler, yellow- 
brown to brown. 

Body subcylindrical, submoniliform. Postcollar constriction faint, width of 
head = 5-16 > collum = 4 > 2 = 3; on segments 17-20 trunk gradually and gently 
tapering toward telson both in width and in height. Antennae medium-sized, slender, in 
situ reaching beyond segment 3 dorsally (Fig. 10), a little shorter in 9. Paraterga 



814 



S.I. GOLOVATCH 




14 15 



Figs 10-17 
Catharosoma mahnerti sp. n., o* paratype: 10) anterior body portion, lateral view; 11) metater- 
gum 10, dorsal view; 12) telson, lateral view; 13) hypoproct, ventral view; 14) sternal structures 
between coxae 2-7 (legs 2 at left); 15) sternal structures between midbody coxae; 16) leg 15; 17) 
right gonopod, medial view. Scale bars 1.0 (10-16) and 0.2 mm (17). 



strongly reduced, only on segment 2 fully developed as low keels devoid of a caudal 
tooth, discernible on segments 3 and 4 as arcuated lines, on segment 5 as modest 
swellings (Fig. 10), on following segments totally missing. Tegument smooth, shining, 
only metazona at places faintly rugulose. Limbus thin, caudal margin entire. Metaterga 
fully devoid of a transverse sulcus; setae short, often abraded, arranged 3+3 on seg- 
ments 2 and 3. on following segmens 2+2 in a transverse line behind a rather deep 
stricture between pro- and metazona (Fig. 11). Ozopores lateral, opening level to 
metatergal surface ca 1/3 metazonital length away from caudal edge (Fig. 10). 
Pleurosternal carinae like narrow keels with a caudal spinule (Fig. 10) traceable until 
segment 16 (9) or 17-18 (<3), onward wanting. Epiproct (Fig. 12) rather long, coni- 
form, faintly emarginate at tip in dorsal view, in â a little longer than in 9 . Hypoproct 



PARADOXOSOMATIDAE IN PARAGUAY 815 

(Figs 12, 13) subtriangular, tip narrowly rounded, nearly pointed, straight; 1 + 1 para- 
median setae at caudal edge poorly separated. Paraterga, pleurosternal carinae and 
hypoproct a little more strongly developed in â than in 9 . 

Sternal structure (Figs 14, 15) much like in C. bilineatum sp. n., but with two 
paramedian pilose ridges between coxae 3 of S and paramedian bunches of setae on 
poorly developed knobs between coxae 6 of S . 

Legs without tarsal brushes but densely setose ventrally; setation gradually 
thinning out toward telson; tibiae swollen ventrally and supporting characteristic 
brushes (Fig. 16) between leg-pair 7 and two last pairs. Legs in S a little longer than 
in 9 , as usual becoming a little longer and slenderer toward telson. Each coxa 2 of â 
with a strong, disto ventral, sigmoid, apically pointed process carrying a gonopore at 
base (Fig. 14). 

Gonopods (Fig. 17) highly complex, much like in C. bilineatum n. sp., but cox- 
ite more elongate, telopodite circular and elongate, while solenophore ventrally with a 
much larger, hyaline lobe/base of lamina medialis (m), an apically strongly unciform 
parabasal lobe (p) followed by a spiniform process supporting a flagelliform 
solenomere at base of a hyaline lobule (h), this lobule supporting the penultimate 1/4 
extent of both lamina lateralis and lamina medialis; distalmost 1/4 extent of both 
laminae likewise slender, unciform, free of support. 

Remarks: Judging from the particularly elongate gonopod telopodite, which 
includes the femorite, this new species seems to be especially close to C. curitibense 
and C. intermedium, but the circular telopodite and the shape of its individual elements 
in C. mahnerti sp. n. are quite characteristic. 

At first Jeekel (1963) merged Catharosoma and Mestosoma together, but later 
(1968) he separated them on the account of the apparently more apomorphic character 
states observed in Mestosoma species. This viewpoint has also been shared by 
Hoffman (1980). 

Broelemannopus minutus sp. n. Figs 18-25 

Material: Holotype 6 (MHNG), Paraguay, Alto Parana Prov., Forestry School, 12 km S 
of Puerto Presidente Stroessner, soil sample 1 , 04.1983, leg. C. Dlouhy. - Paratypes: 20 6 6 , 22 
9 9,2 juv. (MHNG), 2 6 6 , 2 9 9 (ZMUM), same locality, together with holotype; 1 6 , 1 9 
(MHNG), same locality, soil sample 8, 04.1984; 2 6 6 , 1 9 (MHNG), same locality, soil sample 
24, 06.1984; 3 66 (MHNG), same locality, soil sample 10, 05.1984; 19,3 juv. (MHNG), same 
locality, soil sample 22, 25.06.1984; 1 6, 1 "9 (MHNG), same locality, soil sample 25, 
25.06.1984; 1 6 (MHNG), same locality, soil sample 7, 04.1984; 1 6 , 3 juv. (MHNG), same lo- 
cality, soil sample 23, 25.06.1984; 2 66 (MHNG), same locality, soil sample 3, 21.05.1984; 1 
6,1 9,1 juv. (MHNG), same locality, soil sample 13, 04.1984; 1 6 , 1 9 , 1 juv. (ZMUM), same 
locality, soil sample 30, 04.1984; 14 66, 17 9 9 (MHNG), same locality, soil sample 12, 
04.06.1984, all leg. C. Dlouhy. 

Name: To emphasize the small body size. 

Diagnosis: Differs from congeners by the particularly small size, the especially 
strongly reduced paraterga visible only on segment 2, the lack of transverse metatergal 
sulci, the absence of a distinct colour pattern and of an unciform hypoproct, and in 
certain details of solenophore structure. 

Description: Length 7-9 mm (6 , 9), width of midbody metazona 0.6-0.7 (6) 
to 0.8-0.9 mm ( 9 ); 9 usually larger and somewhat broader than 6 . Holotype ca 8 mm 



816 



S.I. GOLOVATCH 




Figs 18-25 
Broelemannopus minutes sp. n., 6 paratype: 18) anterior body portion, lateral view; 19) caudal 
body portion, lateral view; 20) sternal structures between coxae 3-7 (coxae 3 on top); 21) ster- 
nal structures between midbody coxae; 22) leg 10; 23-25) right gonopod, medial, lateral and an- 
terodorsal views, respectively. Scale bars 0.5 (18-19), 0.25 (20-22) and 0.1 mm (23-25). 



long and 0.7 mm wide. Coloration in alcohol uniformly brown to red-brown, only 
juveniles, antennae and legs paler, yellow-brown to brown; tip of antenna pallid. 

Body subcylindrical, not moniliform. Postcollar constriction faint, width of 
head > collum >2>3<4<5-16:on segments 17-20 trunk gradually and gently 
tapering toward telson both in width and in height. Antennae medium-sized, slender, in 
situ reaching the end of segment 2 dorsally (Fig. 18), a little shorter in 9 . Paraterga 
strongly reduced, only present as low lines devoid of a caudal tooth on segment 2 
(Fig. 18), on following segments totally missing. Tegument smooth, shining. Limbus 
thin, caudal margin entire. Metaterga fully devoid of a transverse sulcus; setae 
relatively long, sometimes abraded, arranged 2+2 on segment 2, on following seg- 
ments 1 + 1 in a transverse line behind a shallow stricture between pro- and metazona 
(Fig. 18). Ozopores lateral, opening level to metatergal surface ca 1/3 metazonital 
length away from caudal edge. Pleurosternal carinae like narrow arcuated lines (Fig. 
18) traceable until segment 16 ( 9 ) or 17 (<3), on following segments wanting. Epiproct 
(Fig. 19) rather long, coniform, in 6 a little longer than in 9 , tip with a stong claw- 
shaped uncus directed dorsocaudally. Hypoproct (Fig. 19) subtriangular, tip narrowly 
rounded, nearly pointed, straight; 1 + 1 very long paramedian setae at caudal edge 
poorly separated from each other. Paraterga. pleurosternal carinae, epi- and hypoproct 
in 6 a little more strongly developed than in 9 . 



PARADOXOSOMATIDAE IN PARAGUAY 817 

Stema mostly sparsely setose. Sternum between coxae 3 of â furnished with 
several transverse rows of relatively small, subcontiguous, scale-like structures on a 
boss (Fig. 20). Sterna between coxae 4 and 5 of S with a very prominent, linguiform, 
laterally setose outgrowth directed anteroventrad (Fig. 20); a paramadian pair of 
bunches of setae between coxae 5 and 7 of S (Fig. 20); a central, compact and thicker 
bunch of setae on a swelling between coxae 6 of $ (Fig. 20). Postgonopodial sterna 
without modifications (Fig. 21). 

Legs without tarsal brushes but densely setose ventrally, without modifications 
(Fig. 22); setation gradually thinning out toward telson; legs in S a little longer than 
in $ , as usual becoming a little longer and slenderer toward telson. Each coxa 2 of â 
nearly unmodified, with a small distoventral cone surmounted by a gonopore. 

Gonopods (Figs 23-25) not particularly complex. Solenophore ventrally with a 
distinct but not hypertrophied, rounded, hyaline lobe/base of lamina medialis (m); 
another simple, parabasal lobe (p) supporting the tip of a flagelliform solenomere at 
base of a smaller hyaline lobule (h), the latter supporting the subterminal part of both 
lamina lateralis and lamina medialis; distalmost extent of both laminae very short and 
rather broad, subunciform and subtruncate. 

Remarks: Judging from the particularly small size, the similarly uncigerous 
epiproct, the paraterga present on segment 2 only, as well as from several other traits, 
including those of the gonopod, the new species seems to be especially close to 
B. ibitiensis, from Monte Alegre, Säo Paulo State, Brazil (Schubart, 1945a). The 
distinctions of B. minutus lie in the total absence of sternal cones on postgonopodial 
segments of the 6 , as well as the presence of a smaller and differently armed lobe 
between coxae 3 of the S , and of a smaller lobe p on the solenophore (see also key 
below). B. ibitiensis has only tentatively been placed in Broelemannopus (see Jeekel, 
1963) but, since this species shows especially close affinities with B. minutus sp. n., 
there can no longer be any doubt that both are congeners best to be assigned to 
Broelemannopus . 

In general the genus Broelemannopus is so poorly distinguished from 
Mestosoma that at first Jeekel (1963) merged these two genera together and treated the 
former as only a species group of the latter. However, later (1968) he separated them 
on the basis of certain apomorphies observed in both (see key below), a view which is 
also shared by Hoffman (1980). 

Mestosoma simplex sp. n. Figs 26-32 

Material: Holotype 6 (MHNG), Paraguay, Prov. Cordillera, 5 km N of Emboscada, Rio 
Piribebuy, forest with bromeliads, dead wood and leaves, 05.04.1985, leg. Expédition 
zoologique du Muséum de Genève. - Paratype: 1 juv. (MHNG), same locality, together with 
holotype. 

Name: To emphasize the only slightly modified legs of the S and the relatively simple 
gonopod structure . 

Diagnosis: Differs from congeners by the relatively small size, the only slight- 
ly modified legs of the â , and the relatively simple solenophore structure. 

Description: Length of holotype ca 12 mm, width of midbody metazona 
1.5 mm. Coloration in alcohol uniformly light red-brown, anterior body portion 



818 



S. I. GOLOVATCH 




Figs 26-32 
Mestosoma simplex sp. n., 6 holotype: 26) anterior body portion, lateral view; 27) left halves of 
segments 10 and 1 1 , dorsal view; 28) sternal structures between coxae 4 and 5 (coxae 4 on top); 
29) leg 15; 30-32) right gonopod, medial, submedial and lateral views, respectively. Scale bar 
0.6 (26-29) and 0.3 mm (30-32). 



slightly more intensely reddish; penultimate antennomere darker brown, tip of antenna 
pallid; legs paler, yellow-brown. 

Body strongylosomoid, not moniliform, with poorly developed but evident 
paraterga. Postcollar constriction apparent, width of head = collum > 2 = 4 « 5-16; on 
segments 17-20 trunk gradually and gently tapering toward telson both in width and in 
height. Antennae medium-sized, slightly davate, in situ reaching beyond segment 2 
dorsally (Fig. 26), a little shorter in juvenile. Paraterga 2 and 5-19 delimited by an 
evident sulcus not only dorsally but also ventrocaudally (Fig. 26), all set low, 
considerably larger on pore-bearing segments than on poreless ones (Fig. 27), only on 
segments 18 and 19 slightly projecting beyond rear tergal contour. Tegument smooth 
and shining. Limbus thin, caudal margin entire. Metaterga 5-17 with a faint but 



PARADOXOSOMATIDAE IN PARAGUAY 819 

apparent transverse sulcus far from reaching base of paratergum, the sulcus being very 
slightly sinuate anteromedially; axial impression on metaterga interrupted in the mid- 
dle, barely visible; tergal setae untraceable (Figs 26, 27). Stricture between pro- and 
metazona thin and shallow (Figs 26, 27). Ozopores lateral, lying on paraterga at ca 1/4 
metazonital length away from caudal edge (Figs 26, 27). Pleurosternal carinae like 
narrow keels delimited by a sulcus dorsally, devoid of a caudal spinule (Fig. 26), trace- 
able until segment 17, on further segments wanting. Epiproct rather long, coniform, 
narrowly truncate at tip in dorsal view, in S a little longer than in juvenile. Hypoproct 
roundly triangular, tip evidently rounded, straight; 1+1 paramedian setae at caudal edge 
rather strongly separated from each other. 

Sterna modestly setose, mainly unmodified but both with an evident, deeply 
emarginate, setose, linguiform outgrowth directed anteroventrally between coxae 4 of 
ô and with a paramedian pair of bunches of setae between coxae 5 of 6 (Fig. 28). 

Legs only slightly modified, in 3 with dense tarsal brushes starting from leg- 
pair 1 and with dense tibial brushes starting from leg-pair 2 (Fig. 29), both these 
brushes thinning out toward telson but absent only from last two leg-pairs. Each coxa 
2 of S with a low distoventral cone bearing a gonopore. 

Gonopods (Figs 30-32) relatively simple. Coxite subcylindrical, not particularly 
elongate, sparsely setose disto ventrally. Telopodite subfalcate, a densely setose 
prefemoral part normal in shape, much shorter than femori te. Solenophore unciform, 
somewhat constricted at base, relatively stout and simple; lobe m at base of both 
solenomere and lamina medialis highly inconspicuous, ledge-shaped, lamina lateralis 
slightly better developed than lamina medialis, tips of both subtruncate. 

Remarks: Due to the basally strongly constricted solenophore, M. simplex sp. n. 
joins group III in the sense of Jeekel (1963), which has hitherto been known to 
comprise only three species, one each from the Brazilian states of Pernambuco, 
Alagoas and Säo Paulo. However, the new species differs in its smaller size, in the 
absence of femoral and some sternal modifications in the S , and in the particularly 
simple gonopod structure. 

Mestosoma crassipes sp. n. Figs 33^2 

Material: Holotype S (MHNG), Paraguay, Prov. Conception, Ao. Tagatya-mi, small 
grove, near "gué", sifted litter, 23.10.1985, leg. Expédition zoologique du Muséum de Genève. 
Name: To emphasize the modified and inflated legs of the 6 . 

Diagnosis: Differs from congeners by the relatively small size, the conspicu- 
ously modified legs of the S , and by the relatively complex gonopod structure. 

Description: Length of holotype ca 12 mm, width of midbody metazona 
1.3 mm. Coloration in alcohol uniformly light brown; antennae brown, penultimate an- 
tennomere darker brown; tip of antenna pallid; legs slightly paler. 

Body strongylosomoid, not moniliform, much like in M. simplex sp. n. Post- 
collar constriction apparent, width of head ? collum = 4>2 = 3<5< 6(7)- 16; on seg- 
ments 17-20 trunk gradually and gently tapering toward telson both in width and in 
height. Antennae medium-sized, slightly davate, in situ reaching beyond segment 2 
dorsally (Fig. 33). All paraterga set low, delimited by an evident sulcus not only 
dorsally but also ventrally to ventrocaudally (Fig. 33), on segment 19 particularly flat 



820 



S.I. GOLOVATCH 




Figs 33-42 
Mestosoma crassipes sp. n., 6 holotype: 33) anterior body portion, lateral view; 34) left half of 
metatergum 10, dorsal view; 35) caudal body portion, lateral view; 36) sternal structures be- 
tween coxae 3 and 4 (coxae 3 on top); 37) leg 7; 38) leg 9; 39-42) left gonopod, submedial, sub- 
lateral, ventrolateral and anterodorsal views, respectively. Scale bars 1 .0 (33-35), 0.5 (36-38) and 
0.3 mm (39-42). 



in lateral view, considerably larger on pore-bearing segments than on poreless ones 
(Figs 33, 34), never projecting beyond rear tergal contour. Tegument smooth and shin- 
ing. Limbus thin, caudal margin entire. Metaterga 5-18 with a faint but apparent trans- 
verse sulcus far from reaching base of paratergum, the sulcus being slightly sinuate an- 
teromedially; axial impression on metaterga wanting; tergal setae untraceable (Figs 33, 
34). Stricture between pro- and metazona thin and shallow (Figs 33, 34). Ozopores lat- 
eral, lying on paraterga at ca 1/4 metazonital length away from caudal edge (Figs 33, 
34). Pleurotergal carinae like narrow keels delimited by a sulcus dorsally, devoid of a 
caudal spinule (Fig. 33), traceable until segment 17, on following segments wanting. 
Epiproct rather long (Fig. 35), coniform, narrowly truncate at tip in dorsal view. 
Hypoproct triangular, tip pointed, straight; 1+1 paramedian setae at caudal edge widely 
separated from each other (Fig. 35). 

Sterna mainly modestly setose, postgonopodial ones unmodified, pregonopodi- 
al ones with a paramedian pair of bunches of setae on small cones between coxae 3 and 



PARADOXOSOMATIDAE IN PARAGUAY 821 

4 of S (Fig. 36); stema between coxae 5-7 of â concave and with some particularly 
long setae (like in Fig. 36). 

Legs modified, with dense tarsal brushes starting from leg-pair 1 and with dense 
tibial brushes starting from leg-pair 2 (Figs 37, 38), both these brushes thinning out to- 
ward telson and virtually absent only from last two leg-pairs. Femora 4-6 conspicu- 
ously inflated and each with an evident distoventral tooth. Legs 7 (Fig. 37) with a dis- 
toventral cone on each coxa and a little less strongly incrassate femora, the tooth situ- 
ated more ventro-apically. Femora 9-11 (Fig. 38) nearly normal, ventral knobs in dis- 
tal one-third gradually disappearing toward leg-pair 12. Each coxa 2 with a low dis- 
toventral cone bearing a gonopore. 

Gonopods (Figs 40-42) somewhat more complex than in the previous congener, 
especially so due to a longitudinal sulcus on dorsal face of femorite, a digitiform 
process (m), not a ledge- shaped structure lying at base of lamina lateralis, as well as a 
short, unciform and pointed solenophore tip. 

Remarks: This new species seems to be particularly close to M. alticola and to 
a few other congeners possessing a small but evident process (m) at base of both the 
solenophore and solenomere (see Attems, 1937). The colour pattern in M. crassipes sp. 
n. resembles that of M. femorale, yet the gonopod structure is quite different (see 
Schubart, 1943). Due to the presence of two small tubercles between coxae 3 of the S 
and an attenuated tip of the solenophore, the new species is similar to M. perfidum, but 
the latter species is larger (17-18 mm long and 1.9-2.0 mm wide versus 12 and 
1.3 mm), and their gonopods are very different as well. 

IDENTIFICATION KEYS 

Key to tribes and genera of Paradoxosomatidae occurring in Paraguay and 
adjacent areas 

1 Paraterga strongly developed, wing-like. (Introduced, normally synan- 

thropic species) 2 

Paraterga poorly developed to missing (Figs 1, 10, 18, 26, 33) (Tribe 
Catharosomatini, autochthonous species) 4 

2 Metaterga densely papillate and setose; a bituberculate process between 
coxae 4 of â ; femora 4-7 of 6 each with a ventro-parabasal knob; gono- 
pod femorite stout, solenophore with a large basal prong ventrally 

Tribe Sulciferini: Chondromorpha (C. xanthotricha) 

Metaterga not papillate, at most poorly setose; sternum between coxae 4 

of S without process; legs of â without such femoral tubercles; gono- 
femorite not so stout, almost as long as to longer than solenophore 3 

3 Caudal corners of paraterga surpassing rear tergal contour only on cau- 
dalmost segments (16-19); gonofemorite relatively short, distinctly 
broadened distally; solenophore bifid, at base with a long apical process 

as well as a prong and a lobe ventrally . . Tribe Sulciferini: Oxidus (O. gracilis) 
Caudal corners of paraterga surpassing rear tergal contour on all body 
segments; gonopod telopodite very slender and simple, rod-shaped; go- 
nofemorite slightly longer than solenophore, latter without outgrowths 

at base but with a minute lobule at tip 

Tribe Orthomorphini: Orthomorpha (O. coarctata) 



822 S.I. GOLOVATCH 

4 Gonopod devoid of a solenophore, solenomere free 5 

Gonopod with a solenophore sheathing and supporting a fiagelliform 
solenomere 9 

5 Gonopod telopodite extremely simple, subfalcate to subcircular, devoid 

of any outgrowths at base of a subflagelliform solenomere . . . Gonodrepanum 
Gonopod telopodite more complex, with 1-2 processes or dilatations in 
femoral or postfemoral region 6 

6 Gonofemorite with a large, medial, lobuliform dilatation; solenomere 

with a small tooth near tip Mogyella (M. nana) 

Gonofemorite without a large dilatation but sometimes with a tooth dis- 
tally or parabasally; 1-2 dilatations or processes at base of solenomere, 
latter sometimes stout and thick 7 

7 Body small, 0.7 mm wide. Gonopod telopodite sub falcate, much like in 
Gonodrepanum; femori te with a small ventro-parabasal tooth; a small 
but evident, elongated, subtriangular dilatation/lobe fused ventrally at 

base with a fiagelliform solenomere Pseudogonodrepanum (P. scitum) 

Body medium-sized, >1.0 mm wide. Gonopod telopodite mostly sub- 
erect, usually with two independent teeth/outgrowths at base of soleno- 
mere 8 

8 Entire gonopod telopodite suberect, with an evident process at base of a 
rather thick, sometimes bifid solenomere; a setose, linguiform, sternal 

process between coxae 4 of ó Habrodesmoides 

Only gonopod femorite elongated and suberect; solenomere subflagelli- 

and unciform, acuminate, at base with two relatively small teeth/out- 
growths; sternal process between coxae 4 of S either missing or repre- 
sented by two paramedian knobs Ologonosoma 

9 Solenophore simple, subflagelliform, acuminate, supporting a similarly 

fiagelliform solenomere 10 

Solenophore complex, sometimes with a process or lobe at base, usually 
consisting of several more or less folded lobes/laminae, almost entirely 
sheathing a fiagelliform solenomere 11 

10 Body width 0.7 mm. Prefemoral (setose) part of gonopod much shorter 
than acropodite; femorite with a conspicuous lamella on medial side; 
both solenophore and solenomere somewhat reduced, about half as long 

as femorite Mogyosoma (M. hamatum) 

Body width 1.1-1.5 mm. Prefemoral portion of gonopod elongated, 
nearly as long as acropodite; femorite without a lobe on medial side; 

both solenophore and solenomere longer than femorite 

Gonodrepanoides (G. travassosi) 

1 1 Solenophore relatively simple but with a large lateral branch/process at 

base Promestosoma (P. boggianii) 

Solenophore more complex, usually consisting of several lobes but 
devoid of a large lateral branch at base 12 

12 Sternal cones usually present (Figs 4 & 15); each coxa 2 of S with a 
strong distoventral process carrying a gonopore (Figs 1 & 14); tibiae of 



PARADOXOSOMATIDAE IN PARAGUAY 823 

S usually conspicuously inflated (Figs 5 & 16). Prefemoral (densely 
setose) portion of gonopod hypertrophied, about as long as femorite 

Catharosoma 

Sternal cones usually absent; normally each coxa 2 of 6 with a small 
cone carrying a gonopore; tibiae of <S not inflated. Prefemoral portion of 

gonopod normal, shorter than femorite 13 

13 Tarsal brushes often absent even in S ; sternite between coxa 3 of 6 with 
a conspicuous comb- or tongue-shaped structure (Fig. 20); sternite 
between coxae 5 of S with a particularly large process directed antero- 
ventrally and covering a boss or protuberance if any between coxae 4 

Broelemannopus 

Tarsal brushes in S mostly present; sternite between coxa 3 of S usually 
without protuberances; a particularly large process directed anteroven- 
trally and placed between coxae 4, not 5, of â Me sto soma 

Key to Broelemannopus species of Paraguay and adjacent areas 

1 Body of adult < 10 mm long and < 1 .0 mm wide; epiproct topped with 
a prominent claw-shaped uncus directed slightly dorsad (Fig. 19); cones 

absent at least between front coxae of postgonopodial sterna 2 

Body > 14 mm long and > 1 .6 mm wide; epiproct without central uncus 

at tip, latter usually bifid; sternal cones in â present between both pairs 

of legs of each segment; Brazil 3 

2 Lobe between coxae 4 and 5 of â linguiform, entire (Fig. 20); sternal 
cones on midbody segments of â totally absent (Fig. 21); solenophore 

lobe p small (Figs 23-25); Paraguay B. minutus 

Lobe between coxae 4 and 5 of â divided distally; sternal cones in S 
present between posterior coxae of segments 8-18; solenophore lobe p 
large; Säo Paulo State, Brazil B. ibitiensis 

3 Body 1.6-2.2 mm (<?) or 1.6-2.4 mm wide (9). Paraterga traceable as 
low arcuated sulci on segments 2-4, on following segments only as 
slight impressions visible near caudal margin; pleurosternal carinae 

traceable until segment 14 B . pirassunungensis 

Body 2.3-3.3 mm wide (cT, 9). Each paratergite traceable as a low 
arcuated crest at least on segment 2; pleurosternal carinae visible until 
segment 15 4 

4 Sternal lobe between coxae 3 of â comb-shaped, entire distally; coxae 
2-6 of cT each with a disto ventral coniform process; central protuberance 
between coxae 7 of â absent; distal end of solenophore not divided 

B. escaramucensis 

Sternal lobe between coxae 3 of S linguiform, deeply incised distally; 

only coxae 2 and 3 of 6 each with a coniform process; central protu- 
berance between coxae 7 of 6 present; distal end of solenophore deeply 
divided into two large lobes, p being one of these B. glabratus 



824 S. I. GOLOVATCH 



Key to Catharosoma species 



1 Sternum between coxae 6 of â with a single, undivided process or 

swelling 2 

Sternum between coxae 6 of â either with a deeply divided process or 

with two distinct paramedian processes, or with two paramedian knobs 

with bunches of long setae 5 

2 Hypoproct tip unciform; gonopod f emorite with a prominent, subtrian- 

gular, distal lobe C. digitale 

Hypoproct tip straight; gonof emorite without a prominent, subtrian- 
gular, distal lobe 3 

3 Body width about 2.2 mm; no sternal modifications except pilosity be- 
tween coxae 5 of ct; tip of solenophore simple, like a broad and sub- 
acuminate lobe C. mixtum 

Body width 3.0-3.2 mm; a swelling and/or a paramedian pair of knobs 
between coxae 5 of â ; tip of solenophore more complex 4 

4 Paraterga 2 like swellings, paraterga 3 and 4 sulciform; light subtrian- 
gular spots on posterior parts of proterga and on anterior parts of 
metaterga against a dark background; apical piece of solenophore short, 

acuminate C. peraccae 

Paraterga 2-4 like low crests; background coloration pale yellowish with 

a wide castaneous axial stripe; apex of solenophore very broad and of 
rather irregular shape C. mesorphinum 

5 Even paraterga 2 expressed as sulci; no sternal modifications except 

pilosity between coxae 5 of S C. myrmekurum 

Crest-like paraterga present at least on segment 2 (Figs 1 & 10), often 
ridge/crest-like even on segments 2-4; sternal modifications present 
between coxae 5 of o" 6 

6 Paraterga 2-4 distinctly crest-like, onward sulciform on segments 5,7,9 

and 10; a distinct bilobed process present between coxae 3 of â C . palmatum 
Paraterga even more poorly developed, totally untraceable on segments 
behind 4 th 7 

7 Body about 12 mm long and 1.3 mm wide; coxae 2 of 6 produced 
distally into a small mammiform process; gonofemorite with a medio- 

parabasal digitiform process C. curitibense 

Body width > 1 .7 mm; coxae 2 of â usually produced into a more or less 
prominent process (Fig. 14); gonofemorite devoid of such a process 8 

8 Hypoproct unciform, its tip directed ventrad 9 

Hypoproct tip straight 12 

9 Dorsum rather dark, red-brown, with two yellowish paramedian stripes; 

pleurosternal carinae visible only until segment 15 C. taeniatum 

Dorsum pale but with two dark, brownish paramedian stripes; pleu- 
rosternal carinae visible at least until segment 16 10 

10 Body of â 1.9-2.1 mm wide; a large and deeply incised process present 
between coxae 6 of â (Fig. 3) C. bilineatum 



PARADOXOSOMATIDAE IN PARAGUAY 825 

Body width > 2.6 mm; only a pair of small paramedian knobs between 
coxae 6 of (J 11 

11 Apical papillae on epiproct acuminate, elongate and directed distodor- 

sad; gonofemorite strongly broadened distad C. mesoxanthum 

Apical papillae on epiproct small, simple, inconspicuous; gonofemorite 
slender C. intermedium 

12 A paramedian pair of low setigerous crests present between coxae 3 of 

o"; gonopod telopodite elongate and circular (Fig. 17) C. mahnerti 

At most a paramedian pair of bunches of setae present between coxae 3 

of S ; gonopod telopodite falcate and stout 13 

13 A single protuberance between coxae 5 of â and a paramedian pair of 

conical processes between coxae 6 of â C. palustre 

Two setigerous knobs on a swelling between coxae 5 of â and either a 
swelling or a paramedian pair of spinules between coxae 6 of 6 14 

14 Body width 2.0-2.3 (<?) to 2.5 mm ($); lobe p of solenophore ancori- 

form and bifid C. paraguayense 

Body width 2.6 mm (o*); tip of lobe p of solenophore blunt, devoid of 

teeth C. hoffmani 

Key to Gonodrepanum species 

1 Coloration of adults normally castaneous brown with a wide, uninter- 
rupted, axial, contrastingly creamy stripe; Rio de Janeiro, Brazil 2 

Coloration uniformly castaneous to black, usually devoid of a contrast- 
ingly pallid stripe, or axial stripe/line thin and interrupted; Argentina 
and/or Brazil 3 

2 Smaller, only known from 9: 16-18 mm long and 1.8-2.0 mm wide; 

clear sulci in place of paraterga 2-4 G.flavolineatum 

9 18-21 mm long and 2.2-2.4 mm wide, S 15-16 mm long and 1.5-1.6 

mm wide; paraterga on segment 2 like crests, on segments 3 and 4 like 
clear sulci, on following segments like striae; paramedian pairs of 
setigerous protuberances present between coxae 3-5 of S G. torresae 

3 Body particularly slender: 6 20 mm long and 1 .2 mm wide. Dorsum and 
sides uniformly blackish; paraterga 2-4 crest-like, on following 

segments sulciform; sternal cones absent; Brazil and Argentina 

G. drepanephoron 

Body neither so long (< 14 mm even in 9 ) nor so slender. Coloration of 
adults usually castaneous, sometimes yellowish; paraterga represented 

by sulci at most; sternal cones present; Brazil 4 

4 Epiproct topped with two long, claw-shaped, diverging unci directed 

dorsocaudad; a pale axial stripe/line absent 5 

Epiproct without such unci; a pale, axial, interrupted stripe/line usually 

present 6 

Body 12-14 mm long and 1.3-1.7 mm wide; a small ventral tubercle 
present only on femora 3 and 4 of 6 ; sternal protuberance between coxae 



826 S.I. GOLOVATCH 

3 of S bifid, stema between coxae 4 and 5 devoid of protuberances; 

solenomere subcircular G. grajahuense 

Body 9-12 mm long and 1.0-1.3 mm wide; a distoventral tubercle 
present/traceable on femora 3-9 of S; median sternal protuberances 
present between coxae 3-5 of â ; solenomere regularly and gently sub- 
falcate G.furcatum 

6 Body about 14 mm long and 1.3-1.5 mm wide. Metatergal sulci absent; 

femora of S not modified; a large median process with two knobs 
behind and between coxae 4 of â; solenomere relatively short and 

suberect; Santa Catarina, Brazil G. levisetum 

(G. levisetum var. coniferum seems to only represent a colour form dis- 
tinguished by a yellowish body devoid of a pattern) 
Body 11-12.5 mm long and 1.1 mm wide. Metatergal sulci light but 
present; a pair of diverging processes between coxae 5 of â ; solenomere 
relatively long, subcircular; Rio de Janeiro, Brazil G . falcìferum 

Key to Habrodesmojdes species of Paraguay and adjacent areas 

1 Paraterga replaced by a low arcuated ridge on segment 2, by sulci on 

segments 3 and 4; pleurosternal carinae traceable until segment 18; a 
small protuberance present between coxae 5 of 6; solenomere retrorse, 

with a slender, subfalcate, simple process at base H. costalimai 

Segments 2-18 with only slight sulci/striae in place of paraterga; pleuro- 
sternal carinae present on segments 2-7; sterni te between coxae 5 of â 
devoid of a protuberance; solenomere directed distoventrad, process at 
its base massive, stout and branched H. perturbans 

Key to Mestosoma species of Paraguay and adjacent areas 

1 Sterna between coxae (3)4-6(7) of 6 each with a more or less distinct 

median process or protuberance; sternal cones in â absent 2 

At least some of these sterna devoid of a protuberance; sternal cones in 

6 often present 5 

2 Body width > 2.3 mm; metatergal sulcus present on segments subse- 
quent to 5 th ; no single sternal process between coxae 6 and 7 of S 3 

Body width < 1.5 mm (â): metatergal sulci absent; a single, prominent, 
sternal process between coxae 6 and 7 of â ; Bolivia 4 

3 Coloration uniformly brownish; sternal processes or protuberances 

present between coxae 3-6 of 6; Rio de Janeiro, Brazil M. carioca 

With a wide pale axial stripe against a dark olive-brown background; 
sternal process present between coxae 4 of S , only protuberances or 
swellings between coxae 5 and 6 in â ; Bolivia M. schindleri 

4 Body width 1.0 mm (<?); colour pattern in adults indistinct, body uni- 
formly yellowish; femora 3-5 of S each with a ventral tubercle . . . M. alticola 
Body width about 1.5 mm (8)\ a wide, pale, axial stripe against a cho- 
colate-brown background; femora in â apparently without ventral 
tubercles M. boliviae 



PARADOXOSOMATIDAE IN PARAGUAY 827 

5 Sterna of segment 6 in â with a pair of processes at anterior border, 
sterna of segment 5 of S devoid of processes; neither tibial nor tarsal 

brushes present M. differens 

Sterna of segment 6 in â without processes, those of segment 5 of â 

with or without processes; tibial and tarsal brushes in S normally present ... 6 

6 Colour pattern relatively distinct, usually entire dorsum or axial stripe 
pale to yellowish, sometimes this stripe divided into two paramedian 
stripes or broken into series of spots, more or less strongly contrasting 

with a dark, normally brown background coloration of the sides 7 

Colour pattern indistinct, normally dorsum and sides uniformly brown .... 14 

Sternal process between coxae 4 of a present, conspicuous 8 

Sternal process between coxae 4 of 6 absent to very small 10 

8 Body width < 2.0 mm. Dorsum with axial stripe(s), background 
coloration brown; gonofemorite not very broad, solenophore evidently 

longer than 1/2 femorite 9 

Body width > 2.0 mm. Dorsum with an axial series of yellowish spots, 
background coloration dark (red-)brown to black; gonofemorite 
conspicuously broadened, solenophore scarcely half as long as femorite; 
Paraguay 10 

9 Dorsum with two yellowish paramedian stripes; tibial and tarsal brushes 
in 6 present; a bilobate process present between coxae 4 of â ; Paraguay 

M. vittatum 

Dorsum with two series of paramedian spots; tibial and tarsal brushes in 

S absent; a lobe-shaped process present between coxae 4 of â ; Bolivia 
M. montanum 

10 Solenophore with 2-3 prongs/spines subapically 11 

End of solenophore subtriangular, often rounded 12 

11 Legs uniformly blackish; a pair of paramedian knobs between coxae 3 
of 8; femora 4-7 in S each with a ventral tubercle, each coxa 7 of 6 

with a distoventral process M. tricuspis 

Legs brown; a process present between coxae 4 of S ; apparently neither 
femora nor coxae of S with ventral tubercles/processes . . M. pseudomorphum 

12 Body width about 3.0 mm. Dorsum with a pale, sometimes yellowish 
axial stripe; solenophore strongly curved proximad, its end almost in 

contact with base of femorite M. salvadorii 

Body width about 2.0 mm. Dorsum with an axial series of conspicuous 
yellowish spots; solenophore not so strongly curved 13 

13 Median spots situated on proterga and in anterior parts of metaterga; 
solenophore in medial view almost twice as slender as distofemoral part 

M. canterani 

Median spots lying only in middle of metaterga; solenophore in medial 

view nearly as broad as distofemoral part M. kalliston 

14 A single process present between coxae 4 of â 15 

Either two cones/tubercles or nothing between coxae 4 in â 17 

15 Body width about 2.0 mm. Sternal process between coxae 4 of â short 

and subquadrate; solenophore bifid apically M. balzami 



828 s - I- GOLOVATCH 

Body width 1 .3-1 .8 mm. Shape of sternal process between coxae 4 in 3 

and of solenophore different 16 

16 Metatergal sulci distinct on segment 5 and following segments; sternal 
process between coxae 4 of S short and conical; gonofemorite much 

broader than solenophore M. lugubre 

Metatergal sulci absent; sternal process between coxae 4 of S long and 
rectangular; gonofemorite considerably slenderer than solenophore M. borellii 

17 Solenophore base conspicuously constricted (Figs 30-32) M. simplex 

Solenophore base broader 18 

18 Body length about 12 mm, width 1 .3 mm. Metatergal sulcus visible also 
on segment 18; pleurosternal carinae present until segment 17; a disto- 
ventral tubercle present on femora 4-12 of 6 , and a process on coxae 9 
of 6 (Fig. 38); a characteristic process (m) at base of solenophore, tip 

of solenophore pointed (Figs 39-42) M. crassipes 

Body length at least 17 mm, width 1.8 mm. Metatergal sulci either 
absent or at most traceable only until segment 17; distofemoral tubercles 
present until leg 10 of 6 at most; coxae 9 of d normal; no process at 
base of solenophore, tip of solenophore more or less rounded 19 

19 Body length 30-34 mm, width 3.8-4.2 mm; tip of solenophore very 

broadly subtruncate M. truncatum 

Body length < 29 mm, width < 3.5 mm; tip of solenomere not subtruncate . 20 

20 Faint sternal cones traceable between coxae 3 and behind segment 7 of 

S ; distoventral tubercles present only on femora 9 and 10 of â . . M. perfidum 
Noteworthy sternal modifications absent; distoventral tubercles usually 
present only until femora 7 of 6 21 

21 Tip of solenophore bilobate, rather deeply emarginate/notched in the 

middle, forming more or less equal, rounded lobes 22 

Tip of solenophore different 23 

22 Coloration blackish, legs red-brown; both lobes of solenophore tip sub- 
equal; Bolivia M. derelictum 

Coloration dark brown, legs light brown; proximal lobe of solenophore 

tip somewhat smaller than distal one; Paraguay M . pulvillatum 

23 Tip of solenophore subacuminate, narrowly rounded; Bolivia . . . M. luctuosum 
Tip of solenophore broadly rounded; Brazil and/or Paraguay 4 

24 Body coloration dark, legs and venter contrastingly pale; pleurosternal 
carinae traceable until segment 15; a distoventral tubercle present only 

on femora 7 of â ; ventral brushes on tibiae of â absent M. bicolor 

Body coloration brown, legs and venter only slightly paler than remain- 
ing body; pleurosternal carinae traceable until segment 16; distoventral 
tubercles present on femora 4-7 of 6 ; ventral brushes present both on 
tibiae and tarsi in Ô M. femorale 

Key to Ologonosoma species 

1 Sternal process between coxae 4 of â missing; solenomere retrorse; 

Brazil (Parana) O. iguassuense 



PARADOXOSOMATIDAE IN PARAGUAY 829 

Sternal process between coxae 4 of â represented by two paramedian 
knobs; solenomere directed distoventrad; Paraguay O. sanctum 

ACKNOWLEDGEMENTS 

I am most grateful to Dr Volker Mahnert (MHNG), who entrusted me this valu- 
able material for study. Peter Schwendinger (MHNG) kindly edited an advanced draft. 

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Revue suisse de Zoologie 112 (4): 831-848; décembre 2005 



On the systematic position of Podopterocus Banks and Dinopsocus 
Banks, with a revised diagnosis of the genus Sigmatoneura 
Enderlein (Psocodea: 'Psocoptera': Psocidae) 

K. YOSHIZAWA 1 , C. LIENHARD 2 & A. B. IDRIS 3 

1 Systematic Entomology, Graduate School of Agriculture, Hokkaido University, 
Sapporo, 060-8589 Japan. E-mail: psocid@res.agr.hokudai.ac.jp 

2 Muséum d'histoire naturelle, c.p. 6434, CH-1211 Genève 6, Switzerland. 

3 Center for Insect Systematics, Faculty of Sciences and Technology, Uni versiti 
Kebangsaan Malaysia, 43600 Bangi, Malaysia. 



On the systematic position of Podopterocus Banks and Dinopsocus 
Banks, with a revised diagnosis of the genus Sigmatoneura Enderlein 
(Psocodea: 'Psocoptera': Psocidae). - The genus Podopterocus Banks has 
been known only from male specimens and has been characterized by the 
thickened first antennal fiagellomere and the expanded hind tibia. The genus 
Dinopsocus Banks has been characterized by the thickened first antennal 
fiagellomere. Detailed morphological examinations and brief molecular- 
based identification (only for one species) suggest that both taxa are 
congeneric. Based on characters of female terminalia and fore wing 
venation, both taxa are furthermore considered to be closely related to the 
genus Sigmatoneura Enderlein. The discovery of two new species and 
observations on Sigmatoneura spp. reveal the limited significance of the 
diagnostic characters of Podopterocus and Dinopsocus. Therefore we con- 
sider both Podopterocus and Dinopsocus as junior synonyms of Sigma- 
toneura. Monophyly of the genus Sigmatoneura, including Podopterocus, 
Dinopsocus , and the subgenus Longifolia Li, is well supported by the 
unique fore wing venation of females and by sexually dimorphic fore wing 
venation and coloration. Two new species, Sigmatoneura kakisayap sp. n. 
and Sigmatoneura lemahsayap sp. n., which would be classified under 
Podopterocus or Dinopsocus by the previous generic definitions, are des- 
cribed. Sigmatoneura longicornis comb, n., the type species of Podo- 
pterocus, is redescribed and transferred to Sigmatoneura and the female of 
this species is described for the first time. Dinopsocus atratus, the type 
species of Dinopsocus , is synonymized with S. longicornis. Sigmatoneura 
semicolorata comb. n. is redescribed and transferred from Dinopsocus to 
Sigmatoneura. 

Keywords: Psocodea - Psocidae - Sigmatoneura - Podopterocus - 
Dinopsocus - systematics - Malaysia - Indonesia - Brunei - Singapore. 



Manuscript accepted 01 .09.2005 



832 K. YOSHIZAWA ET AL. 



INTRODUCTION 



Podopterocus Banks, 1920 and Dinopsocus Banks, 1920 are enigmatic taxa of 
the family Psocidae. The former genus has been known only from one species, 
Podopterocus longicornis Banks, 1920, and only from male specimens (see New, 
1975) characterized by highly specialized, broadened hind tibiae (Figs ID, 3C) and a 
thickened first antennal flagellomere (Fig. ID). Podopterocus was once treated as a 
subgenus of Eremopsocus McLachlan, 1866 by Roesler (1944) but is now regarded as 
a separate genus (Mockford, 1975) and is assigned to the tribe Cerastipsocini of the 
subfamily Psocinae (Lienhard & Smithers, 2002). The genus Dinopsocus on the other 
hand contains two species, which are also characterized by a thickened first antennal 
flagellomere. As Podopterocus, Dinopsocus was once treated as a subgenus of 
Eremopsocus but is now regarded as a separate genus and assigned to the tribe 
Cerastipsocini (Lienhard & Smithers, 2002). However, a detailed examination of 
phylogenetically relevant characters of these genera has not yet been carried out. 
Therefore their exact taxonomic status and systematic position remained unclear. In 
addition, New (1978) tentatively suggested that Podopterocus and Dinopsocus may 
represent the same taxon, closely related to the genus Sigmatoneura Enderlein, 1908, 
a member of the tribe Metylophorini (Lienhard & Smithers, 2002). 

Recently we obtained some specimens that can be identified as Podopterocus 
and/or Dinopsocus collected in Brunei, Indonesia (Sumatra), Singapore and Malaysia 
(peninsula and Sabah). By comparing these specimens and after having examined the 
type material of D. atratus, the type species of the genus Dinopsocus, we concluded 
that these two genera are actually congeneric and show extreme sexual dimorphism. 
Furthermore, extensive examination of specimens of Sigmatoneura revealed that the 
diagnostic characters for Podopterocus and Dinopsocus are not clear-cut but rather 
continuous or homoplastic. In the following, we synonymize Dinopsocus and 
Podopterocus with Sigmatoneura and discuss their systematic position based on a 
morphological examination of this material. We also describe two new species that 
would be assigned to Podopterocus and/or Dinopsocus in the previous sense, based on 
specimens from Malaysia and Sumatra. 

Methods of morphological examination follow Yoshizawa (2002). Specimens 
stored in 70% or 99.5% ethanol were used (except for dry preserved Dinopsocus 
types). Depositories of specimens are abbreviated as follow: MCZ - Museum of 
Comparative Zoology, Harvard University, USA (type specimens of Dinopsocus); 
MHNG - Muséum d'histoire naturelle, Genève, Switzerland; SEHU - Hokkaido 
University Insect Collection, Sapporo, Japan; UKM - Center for Insect Systematics, 
Universiti Kebangsaan Malaysia, Bangi, Malaysia. 

SYSTEMATICS 

Sigmatoneura Enderlein 

Sigmatoneura Enderlein, 1908: 761. Type species: Cerastipsocus subcostalis Enderlein, 1903. 
Podopterocus Banks, 1920: 308, syn. n.; Mockford, 1975: 251. Type species: Podopterocus 

longicornis Banks. 1920. Eremopsocus {Podopterocus): Roesler, 1944: 147. 
Dinopsocus Banks, 1920: 307, syn. n.: Mockford, 1975: 251. Type species: Dinopsocus atratus 

Banks, 1920. Eremopsocus (Dinopsocus): Roesler, 1944: 147. 



PODOPTEROCUS, DINOPSOCUS AND S1GMAT0NEURA 833 

Revised diagnosis. Large psocids, fore wing length about 6 mm in male and 
about 8 mm in female. Antenna very long, first flagellomere normal or sometimes thi- 
ckened in both sexes (Fig. 1AB vs Fig. 1CDE). Male fore wing hyaline or sometimes 
with pale pigmentation or blackish brown markings, female fore wing blackish brown 
(Fig. 2); male fore wing venation normal, female fore wing R4+5 strongly sinuate; Rs 
and M fused for a short distance, meeting at a point, or connected by a cross vein of 
variable length in both sexes. Male hind tibia usually weakly flattened and somewhat 
enlarged, rarely paddle-shaped, on each side with a flat wing-like expansion (Fig. 3); 
female hind tibia of normal cylindrical shape (Fig. 1BE). 

Sigmatoneura kakisayap Yoshizawa & Lienhard, sp. n. Figs 1AB, 2 AB, 3 A, 4, 5 

Podopterocus sp. Johnson, Yoshizawa & Smith, 2004: 1774. 

Podopterocus sp. KY240. GenBank (online database for gene sequences): accession number for 

1 8S rDNA of holotype male is AY630557 . 
Podopterocus sp. KY329. GenBank (online database for gene sequences): accession number for 

1 8S rDNA of paratype female is DQ 1 1 6946 . 

Holotype. Male. MALAYSIA (peninsula), Gunung Berembun, Cameron Highlands, 
Pahang, 14. vii. 2003, leg. H. Kojima et al. (canopy fogging) (UKM). 

Paratopes. MALAYSIA (peninsula): 2 males, same data as for holotype (SEHU and 
UKM); 1 male, from the type locality, 15. vii. 2003, leg. H. Kojima et al. (canopy fogging) 
(UKM); 1 female, Gunung Jasar, Cameron Highlands, Pahang, 14. iii. 2003, leg. K. Yoshizawa 
(beating dead branches) (SEHU); 1 female, from the type locality, 15. vii. 2003, leg. 
N. Takahashi (sweeping branches) (UKM). MALAYSIA (Sabah - West Coast Residency): 
1 male (heavily damaged, lacking antennae and hind legs), Mt. Kinabalu, 1500 m, Liwagu Trail 
Section 2, 30. iv. 1987, leg D. Burckhardt & I. Lobi (MHNG); 1 female, Mt. Kinabalu, 1750- 
1850 m, Liwagu Trail, 20. iii. 1983, leg. C. Lienhard (MHNG). INDONESIA (Sumatra): 1 male, 
Pematang Siantar, 1. vii. 1984 - 13. iv. 1985, leg. E. W. Diehl (MHNG). 

Description. MALE (Fig. 1A). Head black, with narrow pale markings on frons 
along internal margins of eyes; gena white. Appendices blackish brown; first flagello- 
mere not thickened. Eye small, IO/D=2.7. Thorax black, except for white membranous 
regions, propleuron, posterior margin of mesoscutum and metascutellum. Fore wing 
(Fig. 2 A) hyaline, with blackish brown tinge along apical margin between veins Rl and 
Ml, veins and pterostigma black; venation normal, Sc reaching C, Rs-M cross vein 
long, first section of CuAl almost perpendicular to the anterior wing margin, CuAl+M 
connection long. Hind wing hyaline, with brownish tinge posteroproximally, veins 
black. Legs (Fig. 1A) blackish brown, distal ends of femora and basal ends of tibiae of 
front and middle legs brown, trochanter and basal end of tibia of hind leg white; hind 
leg long, hind tibia (Fig. 3 A) with symmetrical wing-like expansions along distal 2/3. 
Abdomen whitish in ground color, dorsally black except for 7th and 8th segments, gray 
in distal ventral region. 

Terminalia. Dorsal shelf of clunium absent (Fig. 4AB), dorsal region of clunium 
with round shallow concavity in the middle. Epiproct (Fig. 4B) without membranous 
region anteromedially. Paraproct as in Fig. 4 AB. Hypandrium (Fig. 4C) covered with 
wrinkles, anterolaterally fused with clunium, anteromedially with semicircular mem- 
branous region, posteriorly with roughly trapezoidal projection and with longitudinal 
crest medially. Phallosome (Fig. 4D): parameres very weakly sclerotized; aedeagus 
gradually narrowing to truncated posterior end, posterior margin slightly rounded and 



834 



K. YOSHIZAWA£T AL- 




FIO. 1 

Habitus of Sigmatoneura spp. (C dried specimen, others in alcohol). A: S. kakisayap sp. n., male 
paratype; B: S. kakisayap sp. n., female paratype; C: S. semicolorata, male holotype; D: S. lon- 
gicornis, male from Brunei; E: S. longicornis, female from Singapore. 

with tiny denticules laterally; phallobase gradually tapering anteriorly, with very short 
projection (anteriorly). 

Length (in mm): body 4.3-4.5; fore wing 5.8-6.1; hind wing 3.6-3.8. 

FEMALE. Almost as in male, except as follows. Head (Fig. IB): white mar- 
kings along eyes broad, expanded from antennal sockets to vertex. First and second 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 



835 






Fig. 2 
Fore wings of Sigmatoneura spp. A: S. kakisayap sp. n., male holotype; B: S. kakisayap sp. n., 
female paratype; C: S. semicolorata, male holotype; D: S. lemahsayap sp. n., male holotype; E: 
S. lemahsayap sp. n., female paratype; F: S. longicornis, male from Brunei; G: S. longicornis, 
female from Singapore. 



segments of maxillary palpus white. IO/D=2.9. Thorax: laterally with broad longitu- 
dinal white band. Mesoscutellum yellowish white. Fore wing (Fig. 2B) blackish 
brown, with white region along first section of Cu Al and along posterior wing margin 
between veins CuA2 and Ani; veins and pterostigma black, except the following 



836 K - YOSHIZAWA ET AL. 

yellowish white veins: basal section of Rs, basal 2/3 of R2+3, basal 3/4 of R4+5, Rs- 
M cross vein, M, basal parts of M branches, and CuA; R4+5 strongly sinuate in the 
middle, Rs-M cross vein long, first section of CuAl directed anterodistally, CuAl+M 
connection short. Hind wing pale brown, its veins black. Legs (Fig. IB): hind leg nor- 
mal, hind tibia white in distal 2/3. 

Genitalia. Subgenital plate (Fig. 5A): egg guide pointed apically, with transver- 
sal crest basally; body of subgenital plate wide, connected with egg guide by narrow 
sclerite, anteromedian part broadly membranous. Gonapophyses (Fig. 5B): ventral 
valve long; dorsal valve widely membranous, without distal process; external valve 
short and wide, densely covered with long setae along posterior margin. 

Length (in mm): body 4.6-5.1; fore wing 7.9-8.4; hind wing 5.2-5.3. 

Distribution. Malaysia (peninsula and Sabah), Indonesia (Sumatra). 

Etymology. The specific epithet, a noun in apposition, is a combination of two 
Malayan words, kaki (= leg) and sayap (= wing). The synonymized generic name 
Podopterocus is a combination of the Greek words for «leg» and «wing». 

Remarks. This new species can be distinguished from the other species of the 
genus Sigmatoneura by the long hind tibia with dorso-ventrally symmetrical wing- 
like expansions in the male (Fig. 3 A, cf. 3BC) and by the morphology of the subgenital 
plate in the female. 

Males and females of this species (Fig. 1 AB) are significantly different in fore 
wing coloration and venation, and in hind leg morphology. Therefore male-female 
conspecifity was also checked by analysing a 957 bp fragment of the 18S rDNA gene 
sequence (GenBank accession numbers: AY630557 for holotype male and DQ1 16946 
for one female paratype from Gunung Jasar). No substitution has been identified 
between the gene sequences of the male holotype and the female paratype, which 
supports conspecifity. 

Sigmatoneura semicolorata (Banks) comb. n. Figs IC, 2C, 6 

Dinopsocus semicoloratus Banks, 1920: 307. 

Eremopsocus (Dinopsocus) semicoloratus: Smithers, 1967: 98. 

Specimen examined. Holotype male. PHILIPPINES, Luzon, Makiling, leg. C. F. Baker 
(MCZ: 10801). 

Redescription (coloration in dried condition). MALE (Fig. 1C). Head blackish 
brown. Appendices blackish brown; first flagellomere thickened. Eye small, IO/D=2.4. 
Thorax blackish brown. Fore wing (Fig. 2C) hyaline, basal 1/3 and apical part blackish 
brown, anterior part of apical marking extending proximally to apical part of Rs and 
CuAl+M connection, veins and pterostigma black; venation normal, Sc reaching C, Rs 
and M fused for a short distance, first section of CuAl directed anterodistally, 
CuAl+M connection short. Hind wing hyaline, basal 1/3 brown, veins black. Legs 
(Fig. 1C) blackish brown; hind tibia almost normal, very slightly flattened apically. 
Abdomen blackish brown, with whitish longitudinal band laterally. 

Terminalia. Dorsal shelf of clunium absent (Fig. 6AB). Epiproct (Fig. 6B) with 
narrow membranous region anteromedially, and with slight transversal swelling 
medially. Paraproct as in Fig. 6AB. Hypandrium (Fig. 6C) with smooth surface pos- 
teriorly, lateral region with some weak wrinkles, anterolaterally fused with clunium, 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 



837 




Fig. 3 
Male hind tibiae of Sigmatoneura spp. A: S. kakisayap sp. n., male holotype; B: S. lemahsayap 
sp. n., male holotype; C: S. longicornis, male from Brunei. 



anteromedially with triangular membranous region, posteriorly with roughly trapezoi- 
dal projection. Phallosome (Fig. 6D) relatively long, opened basally; parameres rather 
well sclerotized; aedeagus gradually narrowing to truncated posterior end, posterior 
margin slightly concave and with tiny denticules posterolaterally; phallobase tapering 
anteriorly, without projection. 

Length (in mm): body 3.8; fore wing 5.9; hind wing 4.1. 

Distribution. Philippines (Luzon). 

Remarks. S. semicolorata can be distinguished from the other species of the 
genus Sigmatoneura by the unique male fore wing markings. S. kakisayap sp. n., des- 
cribed above, has also hyaline fore wings with some dark brown markings, but it 
differs from S. semicolorata by the absence of a thickened first flagellomere and by the 
greatly expanded hind tibia. The presence of a thickened first flagellomere in S. semi- 
colorata indicates some relationships with the two species described below, but colo- 
ration of fore wing in male is remarkably different. The female of S. semicolorata is so 
far unknown. 



Sigmatoneura lemahsayap Yoshizawa & Lienhard, sp. n. Figs 2DE, 3B, 7, 8 A 

(?) Dinopsocus atratus Banks sensu New, 1978: 45 (see remarks below). 
Not Dinopsocus atratus Banks, 1920: 307 (see S. longicornis, below). 

Holotype. Male. INDONESIA (N-Sumatra), «Holzweg 2», 10 km NE Prapat, 1050 m, 
98°57' E, 2°44' N, 9. i. 1988, leg. E. W. Diehl (MHNG). 



838 



K. YOSHIZAWA£T AL. 




Fig. 4 
Male terminalia (holotype) of Si gmatoneur a kakisayap sp. n. A: terminalia, lateral view; B: epi- 
proct and left paraproct (trichobothrial field not figured), dorsal view; C: hypandrium, ventral 
view; D: phallosome, ventral view. 



Paratopes. INDONESIA (N-Sumatra): 1 female, same locality as for holotype, 27. xi. 
1989, leg. E. W. Diehl (MHNG); 1 female, Huta Padang, 99°14' E, 2°45' N, 400 m, 2. ii. 1991, 
leg. H. Malicky (MHNG). 

Description. MALE. (Male holotype in bad condition, coloration of body, 
except for wings, almost indistinguishable: i.e., just uniformly pale brown without any 
distinct marking). First flagellomere thickened. Eye small, IO/D=2.6. Fore wing 
(Fig. 2D) pale brown, veins black; venation normal. Sc ending in cell c, Rs-M cross 
vein short, first section of CuAl directed anterodistally, CuAl+M connection short. 
Hind wing pale brown, basal 1/3 darker, veins brown. Hind tibia (Fig. 3B) with very 
weakly developed asymmetrical expansions along distal half, ventral expansion 
stronger (confirmed for both hind legs). 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 



839 




Fig. 5 
Female genitalia (paratype) of Sigmatoneura kakisayap sp. n. A: subgenital plate, showing struc- 
ture (left) and color pattern (right); B: gonapophyses. 

Terminalia. Dorsal shelf of clunium absent (Fig. 7A), dorsal region of clunium 
with round shallow concavity in the middle. Epiproct (Fig. 7 A) with membranous 
region anteromedially. Paraproct as in Fig. 7A. Hypandrium (Fig. 7B) with smooth sur- 
face, only laterally with few wrinkles, anterolaterally fused with clunium, antero- 
medially with broad membranous region, posteriorly with semicircular projection. 
Phallosome (Fig. 7C): parameres very weakly sclerotized; aedeagus gradually 
narrowing to truncated posterior end, posterior margin slightly rounded and with tiny 
denticules; phallobase gradually tapering anteriorly, with short projection (anteriorly). 

Length (in mm): body 4.6; fore wing 6.2; hind wing 4.3. 

FEMALE. Head brown; gena white. Appendices brown; first and second seg- 
ments of maxillary palpus paler. First flagellomere thickened. Eye black, IO/D=3.0. 
Thorax brown, laterally with broad longitudinal white band; meso- and metascutellum 
paler. Fore wing (Fig. 2E) brown, with white region along first section of Cu Al and 
along posterior wing margin between veins CuA2 and Ani; veins and pterostigma 
black, except the following yellowish white veins: basal section of Rs, basal 2/3 of 
R2+3, basal 3/4 of R4+5, Rs-M cross vein, M, basal parts of M branches, and CuA; 
R4+5 strongly sinuate in the middle, Rs-M cross vein long, first section of CuAl 
directed anterodistally, CuAl+M connection short. Hind wing pale brown, with darker 
portion anteroproximally; veins black. Legs brown; trochanters and base of femora 
paler; hind leg of normal cylindrical shape. 

Genitalia. Subgenital plate (Fig. 8A): egg guide with a pair of very shallow 
trenches anterolaterally, truncated apically, apical margin folded upwards and some 
apical setae arising from dorsal surface of egg guide; body of subgenital plate wide, 
connected with egg guide by narrow sclerite, anteromedian part broadly membranous. 
Gonapophyses as in S. kakisayap sp. n. (see description above). 



840 



K. YOSHIZAWAErAL. 




Fig. 6 
Male terminalia (holotype) of Sigmatoneura semicolorata. A: terminalia, lateral view; B: epi- 
proct and left paraproct (trichobothrial field not figured), dorsal view; C: hypandrium, ventral 
view; D: phallosome, ventral view. 



Length (in mm): body 4.2-5.0; fore wing 7.2-7.6; hind wing 5.2-5.3. 

Distribution. Indonesia (Sumatra). 

Etymology. The specific epithet, a noun in apposition, is a combination of two 
Indonesian words, lemah (= weak) and sayap (= wing). The synonymized genus 
Podopterocus was characterized by the wing-like hind tibia of males, but in this species 
the hind tibia is only weakly expanded in males. 

Remarks. S. lemahsayap sp. n. is most similar to S. longicornis , redescribed 
below, because of the thickened first flagellomere in both sexes and the widely pig- 
mented fore wing in males. However, these species can be clearly distinguished by the 
shape of the male hind tibia. In having a less expanded hind tibia, S. lemahsayap is 
similar to most Sigmatoneura species, but it differs from them by its thickened first 
flagellomere. 

The female from Kuala Lumpur described by New (1978) as Dinopsocus atra- 
tus shows an egg guide which is strikingly similar to that of S. lemahsayap (cf. Fig. 8 A 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 



841 




Fig. 7 
Male terminalia (holotype) of Sigmatoneura lemahsayap sp. n. A: epiproct and left paraproct; B: 
hypandrium, ventral view; C: phallosome, ventral view. 



and New, 1978: fig. 6). Therefore we consider it as very probable that New's specimen 
does not belong to S. longicornis (= D. atratus, see below) but to S. lemahsayap. The 
specimen from Kuala Lumpur should be re-examined to check this hypothesis. 



Sigmatoneura longicornis (Banks) comb. n. 



Figs 1DE,2FG,3C,8B.9 



Podopterocus longicornis Banks, 1920: 308. 

Eremopsocus {Podopterocus) longicornis: Roesler, 1944: 147; New, 1975: 245 (redescription of 
male, see remarks below). 



842 K . YOSHIZAWA ET AL. 

Dìnopsocus atratus Banks, 1920: 307, syn. n. 

(?) Not Dìnopsocus atratus Banks sensu New, 1978: 45 (see remarks on S. lemahsayap, above). 

Eremopsocus {Dìnopsocus) atratus: Roesler, 1944: 147. 

Specimens examined. Lectotype female of Dìnopsocus atratus (present designation), 
[MALAYSIA (Sabah):] Borneo, Sandakan, leg. C. F. Baker (MCZ: 10800); paralectotype female 
of D. atratus (present designation), PHILIPPINES [Luzon], Mt. Banahao, leg. C. F. Baker 
(MCZ: 10800); non-type female of D. atratus mentioned in the original description, SINGA- 
PORE, leg. C. F. Baker (MCZ). 

BRUNEI: 1 male (reared from nymph) and 4 nymphs, Bru-88/30 (Bêlait District), 
Sungai Liang, Arboretum of Forest Service, primary forest (mixed dipterocarp forest), alt. about 
50m,21.xi. 1988, leg. C. Lienhard (MHNG); 1 male, Bru-88/42, same locality and collector, alt. 
20-50 m, 25. xi. 1988 (SEHU). MALAYSIA (Sabah): 1 female, Sepilok, 24 km W of Sandakan, 
v-viii, 1983, leg. S. Nagai (MHNG). SINGAPORE: 1 female, Bukit Timah Nature Reserve, 16. 
xi. 1981, leg. D. H. Murphy (MHNG); 1 female, same locality, no date, leg. D. H. Murphy 
(MHNG). 

Diagnosis. The male of this species is easily distinguishable from any other 
psocid by the unique structure of its hind tibia, which is distinctly paddle-shaped due 
to a conspicuous, flattened, wing-like expansion on each side; these expansions are 
dorso- ventrally asymmetrical, the ventral expansion is broadest at basal 1/3 and the 
dorsal expansion broadest at distal 1/3 (Fig. 3C). The female of this species is very 
similar to that of S. lemahsayap sp. n., described above, but can be distinguished by the 
shape of egg guide (Fig. 8B). 

Length of male (Bru-88/42, in mm): body 4.5; fore wing 5.8; hind wing 4.1 . 

Redescription of male terminalia. Clunial dorsal shelf absent (Fig. 9AB). 
Epiproct (Fig. 9B) with membranous region anteromedially. Paraproct as in Fig. 9AB. 
Hypandrium (Fig. 9C) with smooth surface, only mediolaterally with few wrinkles; 
anterolaterally fused with clunium; anteromedially with broad membranous region; 
posteriorly with semicircular projection. Phallosome (Fig. 9D): parameres very weakly 
sclerotized; aedeagus narrowing to truncated posterior end, posterior margin slightly 
concave; phallobase gradually tapering anteriorly, with very short projection (an- 
teriorly). 

Description of female. Very similar to the previous species, S. lemahsayap 
sp. n., but smaller. Apical margin of egg guide of subgenital plate truncated in S. le- 
mahsayap but arched in S. longicornis (Fig. 8AB). 

Length (in mm): body 4.5-4.8; fore wing 6.6-6.7; hind wing 4.6-4.8. 

Distribution. Singapore (Banks, 1920; New, 1975; present study), Malaysia 
(Sabah [Banks, 1920 and present study]; peninsula [Banks, 1938]), Brunei (present 
study), Philippines (Banks, 1920). NOTE: Soehardjan (1958) lists Dinopsocus atratus 
for Sarawak, probably because of an erroneous interpretation of Banks' original state- 
ment («Borneo: Sandakan»), and Endang et al. (2002) erroneously list it for Indonesia. 

Remarks. S. longicornis , the type species of Podopterocus, has been described 
from a male collected in Singapore (sex not mentioned in the original description, but 
the holotype of Podopterocus longicornis must be a male because of having a broadly 
expanded hind tibia). Banks' description and figures (Banks, 1920: pi. 1, fig. 4, fore 
wing and hind tibia) closely correspond to our specimens from Brunei, and we are 
convinced that they belong to the same species. S. longicornis has briefly been redes- 
cribed by New (1975) on the basis of three males from Singapore. The only significant 
difference between New's description and our observations on the Brunei males 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 



843 




Fig. 8 
Female subgenital plates. A: Sigmatoneura lemahsayap sp. n. (paratype); B: Sigmatoneura lon- 
gicornis (specimen from Singapore). 

concerns the «broad rugose apex» of the phallosome mentioned by New, which 
contrasts with the slightly concave, smooth apical margin of the phallosome in the 
Brunei material. The differences concerning the shape of the expanded hind tibia and 
of the phallobase between our Figs 3C and 9D and New's figures 8 and 11, are pro- 
bably due to deformations by slide-mounting (phallosome) and to some teratological 
phenomena or optical distortion (hind leg). In our opinion, it is not very likely that 



844 



K. YOSfflZAWA£rAL. 




Fig. 9 
Male terminalia of Sigmatoneura longicornis (specimen from Brunei). A: terminalia, lateral 
view; B: epiproct and left paraproct (trichobothrial field not figured), dorsal view; C: hypan- 
drium, ventral view; D: phallosome. ventral view. 



New's specimens belong to an additional species closely related to S. longicornis . 
However, this possibility can only be excluded with certainty after a re-examination of 
New's material. Concerning shape of hind tibia, our Brunei males correspond more 
closely to Banks' than to New's figure. In our opinion this justifies the assumption that 
these specimens are conspecific with the male holotype of S. longicornis . 

Male and female specimens of the present material assigned to S. longicornis 
are superficially highly divergent and have not been collected simultaneously. There- 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 845 

fore the proposed male-female combination remains somewhat tentative. However, 
three females examined above were collected in Singapore, type locality of S. longi- 
cornis. Their morphological features do not contradict the present observations concer- 
ning sexual dimorphism in the genus Sigmatoneura (see Discussion, below). The only 
other Sigmatoneura species with a thickened first flagellomere, from which female ge- 
nital morphology has been described, is S. lemahsayap sp. n., which is distinguished 
by the shape of its egg guide (Fig. 8 AB). 

DISCUSSION 

In the present study we examined some psocid species that have been or that 
can be assigned to the genera Podopterocus and/or Dinopsocus which are here placed 
in the synonymy of Sigmatoneura. Although a greatly expanded hind tibia has been 
recognized as the most prominent diagnostic character of Podopterocus, our present 
examination revealed that in some species the hind tibia is sexually dimorphic in shape. 
Since Podopterocus has been diagnosed by such a dimorphic character, the genus has 
been known only from males. The genus Dinopsocus on the other hand has been 
characterized by the thickened first flagellomere, but this character is also present in 
males and females of Podopterocus longicornis, the type species of Podopterocus (Fig. 
IDE). Furthermore, the lectotype of Dinopsocus atratus (type species of Dinopsocus) 
is considered to be a female of P. longicornis. As already suggested by New (1978), it 
is now evident that females of Podopterocus have been assigned to the genus Dino- 
psocus because of this extreme sexual dimorphism. Male-female combination of these 
highly sexually dimorphic psocids was confirmed genetically only for Sigmatoneura 
kakisayap sp. n., described and discussed above, the female of which is not of 
«Dinopsocus type» (i.e., lacking a thickened first flagellomere). However, the result of 
the molecular male-female match clearly indicates that the expanded hind tibia is a 
secondary sexual character only developed in males, while corresponding females have 
a normal cylindrical hind tibia. Similar sexual dimorphism is also known in some 
psocids of the family Caeciliusidae. For example, in the flavidus group of the genus 
Valenzuela Navas (see Mockford, 1993; Lienhard, 1998) and in the genus Phymo- 
caecilius Li (see Li Fasheng, 2002) a more or less conspicuous swelling of the fore ti- 
bia (and sometimes middle tibia) has been observed in males, never in females. Based 
on these observations, we concluded that the species described under Podopterocus 
and Dinopsocus are actually congeners. The above mentioned molecular-based match 
of a «Podopterocus type» male with a «Sigmatoneura type» female definitively 
confirmed close relationships between these taxa, as already suggested by New (1978). 
In the following we discuss the arguments in favor of synonymizing Podopterocus, 
Dinopsocus and Sigmatoneura instead of assigning subgeneric status to Podo- 
pterocus I Dinopsocus within the genus Sigmatoneura s. 1. 

Mosaic distribution of the diagnostic characters of Podopterocus and Dino- 
psocus is evident. S. longicornis, S. lemahsayap sp. n. and S. semicolorata belong to 
the «Dinopsocus type» in having a thickened first flagellomere in both sexes. In 
contrast, S. longicornis and S. kakisayap are clearly of the «Podopterocus type» in 
having a greatly expanded paddle-shaped hind tibia in males. S. lemahsayap has a 
slightly but distinctly expanded hind tibia and therefore is close to «Podopterocus 



846 K . YOSHIZAWA ET AL. 

type». Checking our material of Sigmatoneura s. str. (i. e., species without thickened 
first flagellomere and without significant expansion of male hind tibia: e. g., S. kolbei), 
we observed a general tendency for very slight expansion and flattening of the hind 
tibia in males. 

The apomorphic presence of a long Rs-M cross vein in the fore wing was 
observed in all possible Podopterocus and/or Dinopsocus species (except for S. semi- 
colorata). However, an Rs-M cross vein was also observed in some Sigmatoneura s. 
str. species, showing a certain variability from almost fused at one point to as long as 
in S. kakisayap. All known species of the subgenus Sigmatoneura (Longifolia) Li, 2002 
also have a long Rs-M cross vein (Li Fasheng, 2002). Therefore, this character state 
also cannot be used to diagnose Podopterocus and/or Dinopsocus. 

The thickened first flagellomere is shared by three of four species examined 
above (S. longicornis, S. lemahsayap and S. semicolorata), but a secondary reversal of 
the character state in S. kakisayap is also possible (see also discussion below). New 
(1978) mentioned that the subcosta of Dinopsocus is long and approaching R, whereas 
that of Sigmatoneura is short and approaching the costa, and he tentatively considered 
this character as diagnostic for these genera. However, in all specimens examined here, 
the subcosta is short and approaching the costa (Fig. 2). No other apomorphic charac- 
ters clearly characterizing Podopterocus and/or Dinopsocus could be found. Judging 
from these observations, we concluded that the differences between Sigmatoneura s. 
str. and Podopterocus! Dinopsocus are continuous or highly homoplastic and that it is 
impossible to clearly define the latter genera. Thus, synonymizing these genera with 
Sigmatoneura is the most reasonable and practical solution. Monophyly of 
Sigmatoneura, including Podopterocus, Dinopsocus and the subgenus Longifolia, is 
well supported by the following unique autapomorphies: female fore wing blackish 
brown in ground color, with veins R2+3, R4+5 and first section of Cu Al white, and 
with vein R4+5 strongly sinuate; sexual dimorphism in fore wing coloration and 
venation. Molecular data (18S rDNA: Johnson et al., 2004) also show that 
Sigmatoneura. including Podopterocus! Dinopsocus (at least S. kakisayap), form a 
monophyletic group. 

Among the species examined above, S. longicornis , S. lemahsayap and S. semi- 
colorata share an apomorphic character state, the thickened first flagellomere, whereas 
S. kakisayap has normal antennae in both sexes. In contrast, S. longicornis and S. kaki- 
sayap share an apomorphic condition, the well expanded hind tibia, whereas in S. 
lemahsayap and S. semicolorata the male hind tibia is only slightly expanded. As 
already mentioned above, there are conflicts in the distribution of apomorphic 
character states among these species. Well expanded paddle-shaped hind tibia in males 
might have evolved independently in S. longicornis and S. kakisayap, because the ex- 
pansion in S. kakisayap is symmetrical, whereas that in S. longicornis and S. lemah- 
sayap is clearly asymmetrical, although weakly developed in the latter species. 

The subgenus Longifolia was recognized in the genus Sigmatoneura by Li 
Fasheng (2002). This subgenus shares the characteristic sexual dimorphism in the fore 
wing coloration and venation with the subgenus Sigmatoneura. Longifolia is characte- 
rized by the elongate egg guide of the female subgenital plate (autapomorphic for the 
subgenus) and thus monophyly of the subgenus can be confirmed. However, no 



PODOPTEROCUS, DINOPSOCUS AND SIGMATONEURA 847 

apomorphy supporting the monophyly of the subgenus Sigmatoneura is known. 
Although we do not propose here synonymy of Sigmatoneura and Longifolia, we 
would like to point out the fact that Li's treatment probably established a paraphyletic 
taxon (the subgenus Sigmatoneura). 

Podopterocus and Dinopsocus have previously been assigned to the tribe 
Cerastipsocini, whereas Sigmatoneura has been classified under the tribe 
Metylophorini sensu Mockford (1993) (Smithers, 1980; Lienhard & Smithers, 2002). 
Placement of Sigmatoneura into Metylophorini is based on the shape of the gonapo- 
physes, i.e., absence of the distal process of the dorsal valve, which is considered to 
be an autapomorphy for the tribe. Females of possible Podopterocus and Dinopsocus 
examined here also have this character state (Fig. 5B); therefore placement of 
Sigmatoneura, including Podopterocus and Dinopsocus , into Metylophorini is jus- 
tified. Li Fasheng (2002) established the subfamily Sigmatoneurinae for the genus 
Sigmatoneura, and assigned the tribes Cerastipsocini and Metylophorini (sensu 
Mockford, 1993) to a different subfamily (Cerastipsocinae). As mentioned above, the 
genus Sigmatoneura shares an apomorphic character with the other genera of 
Metylophorini, and Li's subdivision of the family Psocidae has to be rejected. 

ACKNOWLEDGEMENTS 

We thank P. Perkins and S. Cover for the loan of the type specimens of 
Dinopsocus spp. KY thanks H. Kojima, S. Nomura, N. Takahashi, M.Y. Ruslan, 
Y.F. Ng, M. Fauzi and J. Saiful Firdaus for assistance in the field. KY's collecting trip 
to Malaysia was supported by the Japan Society for the Promotion of Science (JSPS) 
grant 14255016 to O. Yata. This study was partly supported by the JSPS grant 
15770052 to KY. CL thanks B. Hauser for organizing joint collecting trips to Sabah 
(1983) and Brunei (1988), H. Malicky, S. Nagai, D. H. Murphy and the late E. W. Diehl 
for the generous gift of psocids collected by them, and P. Schwendinger for technical 
assistance and for critical reading of the manuscript. 

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Harvard College 64(3): 299-362, 7 pis. 
Banks, N. 1938. Further Neuropteroid insects from Malaya. Journal of the Federated Malay 

States Museum 18: 220-235, 2 pis. 
Endang Sri Kentjonowati, Thornton, I. W. B. & New, T. R. 2002. The Psocidae (Insecta: 

Psocoptera) of Java and the eastern islands of Indonesia. Invertebrate Systematics 16(1): 

107-176. 

Enderlein, G. 1903. Die Copeognathen des indo-australischen Faunengebietes. Annales histo- 

rico-naturales Musei nationalis Hungarici 1: 179-344. 
Enderlein, G. 1908. Die Copeognathenfauna der Insel Formosa. Zoologischer Anzeiger 33: 

759-779. 
Johnson, K. P., Yoshizawa, K. & Smith, V. S. 2004. Multiple origins of parasitism in lice. 

Proceedings of the Royal Society of London (B) 271: 1771-1776. 
Li Fasheng 2002. Psocoptera of China (two volumes). Science Press, Beijing. (In Chinese with 

English summary.) 
Lienhard, C. 1998. Psocoptères euro-méditerranéens. Faune de France 83: XX+517 pp. 



K. YOSHIZAWA ETAL. 



LffiNHARD, C. & Smithers, C. N. 2002. Psocoptera (Insecta): World Catalogue and Bibliography. 
Instrumenta Biodiversitatis V, Genève: XLI+745 pp. 

McLachlan, R.1866. New genera and species of Psocidae. Transactions of the Entomological 

Society of London (Third Series) 5: 345-352. 
Mockford, E. L. 1975. Genus Eremopsocus McLachlan: distinction from Cerastipsocus Kolbe 

and review of species (Psocoptera: Psocidae). Psyche 82(2): 244-258. 
Mockford, E. L. 1993. North American Psocoptera (Insecta). Flora and Fauna Handbook 10: 

XVIII+455 pp. Sandhill Crane Press, Gainesville, Florida. 
New, T. R. 1975. Psocidae (Psocoptera) from Malaysia and Singapore. Oriental Insects 9: 

243-259. 
New, T. R. 1978. Notes on some Oriental genera of Cerastipsocinae (Psocoptera: Psocidae). 

Oriental Insects 12: 43-48. 
Roesler, R. 1944. Die Gattungen der Copeognathen. Stettiner Entomologische Zeitung 105: 

117-166. 
Smithers, C. N. 1967. A catalogue of the Psocoptera of the world. Australian Zoologist 14(1): 

1-145. 
Smithers, C. N. 1980. The position of Sigmatoneura Enderlein (Psocoptera: Psocidae). 

Australian Entomological Magazine 6(5): 93. 
Soehardjan, M. 1958. First contribution to a study of Copeognatha (Corrodentia) of the 

Indonesian Archipelago. Idea 11(1): 25-33. 
Yoshizawa, K. 2002. Phylogeny and higher classification of suborder Psocomorpha (Insecta: 

Psocodea: 'Psocoptera'). Zoological Journal of the Linnean Society 136: 371-400. 



Revue suisse de Zoologie 112 (4): 849-857; décembre 2005 



Review of the species of Platypalpus Macquart from Guangxi, China 
(Diptera, Hybotidae, Tachydromiinae) 

Ding YANG 1 2 & Bernhard MERZ 3 

1 Department of Entomology, China Agricultural University, Beijing 100094, China. 
E-mail: dingyang@cau.edu.cn 

2 Key Lab of Insect Evolution & Environmental Changes, Capital Normal University, 
Beijing 100037, China. 

3 Département d'entomologie, Muséum d'histoire naturelle, C. P. 6434, 
CH-1211 Genève 6, Switzerland. E-mail: bernhard.merz@mhn.ville-ge.ch 



Review of the species of Platypalpus Macquart from Guangxi, China 
(Diptera, Hybotidae, Tachydromiinae). - The species of the genus Platy - 
palpus Macquart from Guangxi (Southern China) are reviewed. The 
following 3 species are described, illustrated and discussed: P. alamaculatus 
sp. n., P. maoershanensis sp. n., P. xanthodes sp. n. A key to the species of 
Platypalpus from Guangxi is presented. 

Keywords: Hybotidae - Tachydromiinae - Platypalpus - Guangxi - new 
species. 

INTRODUCTION 

Platypalpus Macquart is one of the largest genera of the subfamily Tachy- 
dromiinae (Hybotidae). It can be easily recognized by the following characters: eyes 
narrowly separated on face, mid leg raptorial, and anal cell present (Chvâla, 1975; 
Grootaert & Chvâla, 1992). The genus has a worldwide distribution and its center of 
diversity seems to be the Western Palaearctic region with some 250 described species 
(Chvâla & Kovalev, 1989; Grootaert & Chvâla, 1992). In contrast, only some 31 
species were described from the Oriental region up to 1975 (Smith, 1975). Since then, 
25 species from China have been added by Yang, D. & Yang, C. (1989, 1990, 1992, 
1997), Yang, C. & Yang, D. (1992), Saigusa & Yang (2002), Yang et al (2002). 
Therefore, 33 species of Platypalpus have been reported from China. The major 
references dealing with Platypalpus are as follows: Brunetti, 1920; Frey, 1943; Smith, 
1965; Chvâla, 1975, 1989; Grootaert, 1984; Grootaert & Chvâla, 1992. 

The genus Platypalpus from Guangxi (Southern China) is poorly studied with 
only Platypalpus guangxiensis Yang & Yang, 1992 known from this province. In the 
present paper, three species are described as new to science, based on the specimens 
collected by Ms Shuwen An and Mr Xingyue Liu by sweep net and light trap in 2003. 
A key to all four species from Guangxi is presented. 



Manuscript accepted 01.02.2005 



850 D. YANG & B. MERZ 

MATERIALAND METHODS 

The specimens on which this study is based on are deposited in the following 
collections: 

CAU = Insect collection of China Agricultural University, Beijing, China 
MHNG = Muséum d'histoire naturelle, Genève, Switzerland 
In order to allow an easy comparison of descriptions of various authors, the ter- 
minology of morphological terms follows generally Grootaert & Chvâla (1992), except 
for crossveins which are written in capitals. 

The following abbreviations for bristles are used: acr-acrostichial, ad-antero- 
dorsal, av-antero ventral, dc-dorsocentral, h-humeral, oc-ocellar, npl-notopleural, pd- 
posterodorsal, psa-postalar, pv-postero ventral, sa-supraalar, sc-scutellar, vt- vertical. 

KEY TO THE SPECIES OF PLATYPALPUS FROM GUANGXI 

1 Thorax mainly yellow; wing without dark spots; mid femur entirely yellow . . 2 
Thorax black; wing with brown spot between Costa and R-M (Fig. 1); 

mid femur black with yellow tip alamaculatus sp. n. 

2 Mesonotum entirely yellow 3 

Mesonotum yellow with rather large black mid-longitudinal spot 
maoershanensis sp. n. 

3 Scutellum yellow; apical projection of right genital lamella with wide 

obtuse apex (Fig. 11) xanthodes sp. n. 

Scutellum black; apical projection of right genital lamella with acute 

apex guangxiensis Yang, C. & Yang, D., 1992 

SPECIES TREATMENT 

1 . Platypalpus alamaculatus sp. n. Figs 1-5 

Material 

Holotype 6 , CHINA: Guangxi: Maoershan National Nature Reserve, 1100-1600 m, 
29 .VI .2003, Xingyue Liu (CAU). 

Paratypes: 56 a, 79 9, same data as holotype (CAU); 5ââ, 109 9, Guangxi: 
Maoershan National Nature Reserve, Hongjunting, 1600 m, light trap, 28.VI.2003, Xingyue Liu 
(CAU); 3 ó*ó\ 39 9, Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, 
light trap, 30.VI.2003, Xingyue Liu (MHNG); 4cî Ô , 1 9 , Guangxi: Maoershan National Nature 
Reserve, Sanjiangyuan, 1900 m, light trap, 30.VI 2003, Shuwen An (CAU). 

Etymology 

The specific name refers to the maculate wing. 

Diagnosis 

Mainly black species; head with 2 vt; wing with a brown spot between C and 
R-M (Fig. 1); legs yellow, but mid and hind coxae black; mid femur black with yellow 
tip; mid femur with row of pv bristles; mid tibia with long acute apical spur bearing 1 
curved apical hair (Fig. 2). 

Description 

Male. Body length 2.6-2.8 mm, wing length 2.9-3.1 mm. 



PLATÌPALPUS FROM GUANGXI 



851 




Figs 1-5 
Platypalpus alamaculatus sp. n, male. 1, vving; 2, apex of mid tibia; 3, right genital lamella; 
4, genitalia, dorsal view; 5, left genital lamella. 



Head, including occiput and clypeus black, gray microtrichose. Eyes narrowly 
separated on face; frons wider than face. Jowls in lateral view very narrow, almost 
invisible. Hairs on head pale, bristles black. Ocellar tubercle with 1 oc and 2 posterior 
hairs; 2 vt (curved inward vt short). Antenna black; pedicel with a circle of apical hairs; 
first flagellomere short conical, about 1.3 times longer than wide, with some short pale 
apical hairs; arista 4.5-4.7 times as long as first flagellomere, black, short pubescent. 
Proboscis blackish with blackish bristles; palpus brown to blackish with 3 long pale 
bristles (apical bristle long). 

Thorax black, grey microtrichose. Hairs on thorax pale, bristles blackish; 
1 short weak h curved medially, 2 npl (anterior npl short), acr in about 4 irregular rows, 
1 short psa, long dc and prese absent, scutellum with 2 pairs of sc, basal sc one third 
as long as apical sc. Legs yellow; fore coxa yellow, mid coxa and posterior half of hind 
coxa black; mid trochanter black; fore and hind femora yellow, mid femur black with 
yellow tip; tarsomeres 3-5 of all legs gradually becoming darker towards tip. Hairs on 
legs blackish, bristles black. Coxae with pale bristles and hairs. Fore femur 1.3 times 
and mid femur 2.0 times as wide as hind femur; fore femur with one row of short av 
and one row of long pv (4 apical av the longest); mid femur with two rows of very short 
black v spine -like bristles (but ventral bristles on basal quarter longer), and with one 
row of rather long brownish yellow pv; fore tibia with 3 ad, apically with 1 ad; mid 
tibia (Fig. 2) with a row of short weak black ventral bristles and long acute apical spur 



852 D. YANG & B. MERZ 

(distinctly longer than thickness of mid tibia) bearing 1 curved apical hair. Wing 
(Fig. 1) hyaline with a brown spot from tip of cell c running to R-M; veins dark brown, 
R 4+5 and M slightly convergent apically; cross veins almost contiguous. Squama 
yellow with brownish margin, with pale hairs. Halter yellow with brownish yellow 
base. 

Abdomen subshining black, very thinly grey microtrichose. Hairs on abdomen 
pale. Male genitalia (Figs 3-5): Left genital lamella rather narrow in dorsal view, its 
apical projection finger-like; right genital lamella rather wide in dorsal view, its apical 
projection rather small and subtriangular; both cerei equally long, right cercus slightly 
narrower. 

Female. Body length 2.3-2.5 mm, wing length 2.8-3.0 mm. Similar to male, but 
av and pv on fore femur shorter except 3 bristle-like apical av. 

Distribution 

China (Guangxi). 

Remarks 

The new species belongs to the P. pallidiventris-cursitans species-group 
(Chvâla, 1989). It can be easily separated from other known species from China by the 
colouration of legs and wing pattern. 

2. Platypalpus guangxiensis Yang, C. & Yang, D. 

Platypalpus guangxiensis Yang, C. & Yang, D., 1992: 46. Type locality: Guangxi (Tianlin). 

Diagnosis 

Thorax yellow but scutellum black. Abdomen yellow, but tergites medially and 
apically brownish. Apical projection of right genital lamella spine-like, left cercus 
spine-like and curved inwards apically. 

Distribution 

China (Guangxi, Sichuan, Hubei). 

3. Platypalpus maoershanensis sp. n. Figs 6-9 

Material 

Holotype â , CHINA: Guangxi: Maoershan National Nature Reserve, 2100 m, light trap, 
5. VII. 2003. Shuwen An (CAU). 

Paratype: 1 9 , Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, 
light trap, 30.VI.2003, Shuwen An (CAU). 

Etymology 

The species is named after the type locality. 

Diagnosis 

Mainly yellow species with 2 vt. Mesonotum subshiny with one rather large, 
black mid-longitudinal spot; h, dc and row of pv on mid femur absent. Legs yellow 
with dark brown tarsomere 5. Antenna yellow; first flagellomere short oval. 



PLATYPALPUS FROM GUANGXI 



853 




Figs 6-9 
Platypalpus maoershanensis sp. n, male. 6, apex of mid tibia; 7, right genital lamella; 8, geni- 
talia, dorsal view (long apical bristles on left genital lamella omitted, cf. Fig. 9); 9, left genital 
lamella. 



Description 

Male. Body length 2.7 mm, wing length 3.3 mm. 

Head black, grey microtrichose. Eyes narrowly separated on face; face slightly 
narrower than frons. Hairs and bristles on head pale. Ocellar tubercle with 2 oc and 
2 posterior hairs; 2 vt. Antenna pale yellow; scape without dorsal hairs; pedicel with a 
circle of pale apical hairs; first flagellomere short oval, 1.2 times longer than wide, with 
short pale apical hairs; arista 4.2 times as long as first flagellomere, black, short pu- 
bescent. Proboscis yellow with blackish tip, with sparse black hairs; palpus yellow 
with 4 pale bristles (of which apical bristle is rather long). 



854 D- YANG & B. MERZ 

Thorax yellow, thinly pale grey microtrichose; mesonotum subshiny with one 
rather large mid-longitudinal black spot; scutellum, metano turn and lateropleuron (= 
anatergite & katatergite) (except lateral portion) black. Bristles and hairs on thorax 
pale; h absent, 2 npl (anterior npl short, hair-like), 1 long psa, short, sparse acr nearly 
6-seriate, dc absent, 1 prese (shorter and weaker than psa) in front of scutellum, scutel- 
lum with 2 pairs of sc (basal pair 1/3 as long as apical pair). Legs yellow with dark 
brown tarsomere 5. Bristles and hairs on legs pale; fore coxa with 4 dorsal apical bris- 
tles, mid coxa with 4 dorsal apical bristles, hind coxa with 3 dorsal apical bristles. Fore 
femur weakly and mid femur distinctly thickened, fore femur 1.1 times and mid femur 
1.7 times as wide as hind femur; fore femur apically with 1 dorsal bristle, 1 av and 1 
pv; mid femur with two rows of spine-like ventral bristles (posterior row slightly 
longer than anterior row, but basal ventral bristle rather long and mostly brownish yel- 
low), without row of pv; mid tibia with a row of short blackish ventral bristles and fin- 
ger-like apical spur which is longer than thickness of tibia. Wing hyaline; veins brown- 
ish yellow, R4 + 5 and M weakly convergent apically. Squama yellow with pale hairs. 
Halter yellow. 

Abdomen yellow, thinly pale grey microtrichose; tergites 1-5 and hypopygium 
dark brown. Hairs on abdomen pale. Male genitalia (Figs 7-9): Left genital lamella 
rather elongate in dorsal view, its apical projection wide and obtuse, apically with long 
bristles; right genital lamella rather wide in dorsal view, its apical projection very short 
and with wide apex; left cercus with acute apex, right cercus with wide rounded apex. 

Female. Body length 2.8 mm, wing length 3.1 mm. Similar to male. 

Distribution 

China (Guangxi). 

Remarks 

This new species cannot be placed in any of the species group defined by 
Chvâla (1989). It is similar to Platypalpus hamulatus Yang & Yang, 1989, from Tibet, 
but the mid-longitudinal spot on mesonotum is uniformly wide. In P. hamulatus, the 
mid-longitudinal spot on mesonotum is widened towards the scutellum (Yang, D. & 
Yang, C, 1989). 

4. Platypalpus xanthodes sp. n. Figs 10-13 

Material 

Holotype S , CHINA: Guangxi: Maoershan National Nature Reserve, 1100-1600 m, 
29.VI.2003, Xingyue Liu (CAU). 

Paratypes: Id 19, same data as holotype (CAU); 3 9 9 , Guangxi: Maoershan National 
Nature Reserve. Hongjunting, 1600 m, light trap, 28.VI.2003, Xingyue Liu (CAU); 1 6 , 3 9 9 , 
Guangxi: Maoershan National Nature Reserve, Sanjiangyuan, 1900 m, light trap, 30.VI.2003, 
Xingyue Liu (CAU): ló\ 3 9 9, Guangxi: Maoershan National Nature Reserve, 350 m, light 
trap, 3.VÜ.2003, Shuwen An (MNHG). 

Etymology 

The specific name refers to the yellow thorax and abdomen. It is an invariable 
epithet. 



PLATYPALPUS FROM GUANGXI 



855 




Figs 10-13 
Platypalpus xanthodes sp. n, male. 10, apex of mid tibia; 11, apical projection of right genital 
lamella; 12, genitalia, dorsal view (postgonite [?] drawn as contour behind the cerei); 13, left 
genital lamella. 



Diagnosis 

A mainly subshining yellow species with only the head capsule black; 2 vt; first 
flagellomere about as long as wide, arista much longer; 4 irregular rows of acr; mid 
femur without pv; mid tibia with an obtuse apical spur about as long as diameter of 
tibia; male cerei very asymmetrical; male left genital lamella with long bristles. 

Description 

Male. Body length 2.7-3.0 mm, wing length 2.9-3.1 mm. 

Head black, grey microtrichose, but frons and face orange-brown micro- 
trichose. Eyes narrowly separated on face; face narrower than frons. Bristles and hairs 
on head pale. Ocellar tubercle with 1 oc and 2 very short posterior hairs. Jowls almost 
invisible in lateral view; 2 vt. Antenna yellow; scape without dorsal hairs; pedicel with 
a circle of mostly pale apical hairs; first flagellomere short oval, 1.1 times as long as 



856 D- YANG & B. MERZ 

wide, with some hairs; arista very long, 3.5-3.7 times as long as first flagellomere, 
brown, short pubescent. Proboscis yellow with brown hairs; clypeus yellow, mictro- 
trichose; palpus yellow with 3 long pale bristles, the apical one much longer than the 
others. 

Thorax yellow, thinly grey microtrichose; mesonotum subshiny, without darker 
stripe; katepisternum with indistinct shining yellow spot medially; meron with a small 
longitudinal, black spot. Hairs on thorax pale, bristles brownish yellow; 1 short hair- 
like h curved inward, 2 npl (anterior npl short and hair-like), acr in 4-6 irregular rows, 
dc not longer than acr, 1 long psa, 1 weak prese in front of scutellum, scutellum with 
2 pairs of sc (basal pair very short, 1/3 of apical pair). Legs yellow, but tarsomere 5 
slightly brownish apically. Hairs on legs pale. Fore femur weakly and mid femur dis- 
tinctly thickened, fore femur 1.2 times and mid femur 1.7 times as wide as hind femur; 
mid femur with two rows of short black ventral bristles (posterior row longer than 
anterior row, but ventral bristles on basal quarter long and brownish yellow), without 
row of pv; mid tibia (Fig. 10) with a row of blackish ventral bristles, finger-like apical 
spur about as long as diameter of tibia. Wing hyaline; veins brownish yellow; 
crossveins contiguous; R^^ and M parallel apically. Squama yellow with pale hairs. 
Halter yellow. 

Abdomen yellow, pale grey microtrichose; hypopygium mostly yellow. Hairs 
on abdomen pale. Male genitalia (Figs. 11-13): Left genital lamella rather narrow, its 
apical projection wide and obtuse apically, and with long apical brisües; right genital 
lamella slightly wide in dorsal view, its apical projection short with wide and obtuse 
apex; left cercus long with spine-like apex, right cercus shorter with obtuse apex. 

Female. Body length 2.7-3.1 mm, wing length 2.8-3.2 mm. Similar to male. 

Distribution 

China (Guangxi). 

Remarks 

The new species belongs to the P. longicornis species-group of Chvâla (1989). 
It is very similar to Piaty palpus guangxìensìs Yang & Yang from Guangxi, Sichuan and 
Hubei, but can be separated from the latter by the yellow scutellum and right surstylus 
with wide obtuse apex. In guangxiensis , the scutellum is black, and the right surstylus 
has an acute apex (Yang, C. & Yang, D., 1992). 

CONCLUSIONS 

Guangxi belongs to South China which is considered to be part of the Oriental 
region with a subtropical and tropical climate. Although Platypalpus is basically a 
genus of temperate regions, it is noteworthy that 4 species are known to occur in this 
province. However, they were all found at higher altitude (above 1000 m, only one 
record of P. xanthodes from 350 m). Among them the three new species are endemic 
to Guangxi so far. The only already known species has a much wider distribution with 
records extending to Sichuan and Hubei in Central China region. Taking into account 
the bad collecting effort in Guangxi it is well possible that more species will eventually 
be found in this province. 



PLATYPALPUS FROM GUANGXI 857 



ACKNOWLEDGEMENTS 

Our sincere thanks are due to Ms Shuwen An and Mr Xingyue Liu (China 
Agricultural L T niversity, Beijing) for collecting specimens and for their help during the 
study. The research was funded by the National Natural Science Foundation of China 
(No. 30070 100_No. 30225009). 

REFERENCES 

Brunetti, E. 1920. Diptera Brachycera Vol. 1. The Fauna of British India, including Ceylon & 

Burma. London, 401 pp. 
Chväla, M. 1975. Lhe Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. I. 

Fauna Entomologica Scandinavia 3: 1-336. 
Chväla, M. 1989. Monograph of northern and central European species of Platypalpus 

(Diptera, Hybotidae), with data on the occurrence in Czechoslovakia. Acta Universitatis 

Carolinae - Biologica 32: 209-376. 
Chväla, M. & Kovaley, V. G. 1989. Family Hybotidae (pp. 174-227). In: Soós, A. & Papp, L. 

(eds). Catalogue of Palaearctic Diptera. Vol. 6. Akadémiai Kiadó, Budapest, 435 pp. 
Frey, R. 1943. Übersicht der paläarktischen Arten der Gattung Platypalpus Macq. (=Coryneta 

Meig.). (Dipt., Empididae). Notulae Entomologicae 23: 1-19. 
Grootaert, P. 1984. Fwo new species of Platypalpus Macquart (Diptera, Empidoidea, Hybo- 
tidae) from Papua New Guinea. Bulletin et Annales de la Société Royale d'Entomologie 

de Belgique 120: 283-290. 
Grootaert, P. & Chväla, M. 1992. Monograph of the genus Platypalpus (Diptera: Empidoidea, 

Hybotidae) of the Mediterranean region and the Canary Islands. Acta Universitatis 

Carolinae - Biologica 36: 3-226. 
Saigusa, L. & Yang, D. 2002. Empididae (Diptera) from Funiu Mountains, Henan, China (I). 

Studia Dipterologica 9 (2): 519-543. 
Smith, K. G. V 1965. Diptera from Nepal: Empididae. Bulletin of the British Museum (Natural 

History), Entomology 17 (2): 61-112. 
Smith, K. G. V. 1975. Family Empididae (pp. 185-211). In: Delfinado, M. D. & Hardy, D. E. 

(eds). A Catalog of the Diptera of the Oriental Region. Vol. 2. The University Press of 

Hawaii, Honolulu, 459 pp. 
Yang, C. & Yang, D. 1992. Three new species of Empididae from Guangxi - Diptera: Brachy- 
cera. Journal of the Guangxi Academy of Sciences 8 (1): 44-48. 
Yang, D. & Yang, C. 1989. The dance flies of Xizang (II) (Diptera: Empididae). Acta Agri - 

culturae Universitatis Pekinensis 15 (4): 415-424. 
Yang, D. & Yang, C. 1990. Eleven new species of the subfamily Tachydrominae from Yunnan 

(Diptera: Empididae). Zoological Research 11 (1): 63-72. 
Yang, D. & Yang, C. 1992. Diptera: Empididae (pp. 1089-1097). In: Chen, S. (ed.). Insects of 

the Hengduan Mountains Region. Vol. 2. Science Press, Beijing, 1547 pp. 
Yang, D. & Yang, C. 1997. Diptera: Empididae (pp. 1469-1476). In: Yang, X. (ed.). Insects of 

the Three Gorge Reservoir Area of Yangtze River. Chongqing Publishing House, 

Chongqing, 1847 pp. 
Yang, D., An, S. & Gao, C. 2002. New species of Empididae from Henan (Diptera) (pp. 30-38). 

In: Shen, X. & Shi, Z. (eds). The Fauna and Taxonomy of Insects in Henan. Vol. 5. 

China Agricultural Scientech Press, Beijing, 453 pp. 



Revue suisse de Zoologie 112 (4): 859-867; décembre 2005 



A contribution to the knowledge of the Chyromyidae (Diptera) 
of Italy with description of a new species of Aphaniosoma Becker 

Martin J. EBEJER 

Hendre, Colwinston, Cowbridge, South Glamorgan CF71 7NL, United Kingdom. 

E-mail: martinebejer@evemail.net 



A contribution to the knowledge of the Chyromyidae (Diptera) of Italy 
with description of a new species of Aphaniosoma Becker. - The family 
Chryomyidae is reported from Sicily (eleven species) and from Sardinia 
(nine species). Additional records of species for mainland Italy are also 
given. A new species of Aphaniosoma is described from France, Sicily and 
Sardinia. A total of eleven species are new to Italy: Chyromya miladae, 
Gymnochiromyia fallax, G.flavella, G. inermis, G. mihalyii, Aphaniosoma 
collini, A. claridgei, A. grisescens, A. micromacro, A. proximum, and A. bi- 
falcatum sp. n. A list is given of all the seventeen species so far recorded for 
Italy and its larger islands, together with their data and their distribution in 
the West Palaearctic. 

Keywords: Diptera - Chyromyidae - Aphaniosoma bifalcatum sp. n. - 
Italy - Sardinia - Sicily - France. 

INTRODUCTION 

The family Chyromyidae of Italy has not been well studied and there are no 
published records from Sardinia or from Sicily. In the Checklist of the Diptera of Italy 
(Canzoneri et al., 1995) only three species are listed, namely: Chyromya flava 
(Linnaeus, 1758), C. oppidana (Scopoli, 1763) and Aphaniosoma nigrohirtum 
(Hendel, 1933). All occur in the north of Italy. The two species of Chyromya, according 
to the Checklist, also occur in the south. The Catalogue of Palaearctic Diptera (Soós, 
1984) gives only A. nigrohirtum as specifically occurring in Italy. Thus, both works 
overlooked or did not accept the record, based on a single specimen deposited in the 
Museum of Natural History in Vienna, of A. approximatum Becker, which was given 
in Die Fliegen der Paläarktischen Region (Czerny, 1927). However, this specimen was 
re-examined and it belonged not to approximatum, but to a new species, egregium 
Ebejer (1998b); in the same paper, A. melitensis Ebejer and A. propinquans Collin were 
also recorded from Italy. 

Recent studies of this family of Diptera (Ebejer, 1998a, b; Carles-Tolrâ, 2001) 
have revealed that there are many more species in the Mediterranean. Thus, the list 
given in the Catalogue of Palaearctic Diptera (Soós, 1984) is now very out of date. 
During a brief visit to Sicily in the spring of 1999, eleven species of Chyromyidae were 



Manuscript accepted 26.11.2004 



860 M - J - EBEJER 

encountered. Dr Merz found nine species in Sardinia in the spring of 2002 and Dr Gatt 
collected three species in Tuscany in the summer of 2003 . They are listed in this article 
with their data and summarised in Table 1 . Eleven of these species are new records for 
Italy. 

Table 1: The Italian species of Chyromyidae. (The north and south Italian regions are 
according to the Checklist of the Diptera of Italy.) 

North Italy South Italy Sardinia Sicily 

1 . Chyromya flava (Linnaeus) 

2. Chyromya miladae Andersson 

3. Chyromya oppidana (Scopoli) 

4. Gymnochiromyia fallax Ebejer 

5. Gymnochiromyia flave Ha (Zetterstedt) 

6. Gymnochiromyia inermis (Collin) 

7. Gymnochiromyia mihalyii Soós 

8. Aphaniosoma bifalcatum sp. nov. 

9. Aphaniosoma claridgei Ebejer 

10. Aphaniosoma col lini Lyneborg 

11. Aphaniosoma egregium Ebejer 

12. Aphaniosoma grise scens Ebejer 

13. Aphaniosoma melitensis Ebejer 

14. Aphaniosoma micromacro Carles-Tolrâ 

15. Aphaniosoma nigrohirtum Hendel 

16. Aphaniosoma propinquans Collin 

17. Aphaniosoma proximum Ebejer 



MATERIAL AND METHODS 

Specimens were collected by hand-net sweeping only. All the material, which is 
cited in this article, is dry mounted and deposited in the personal collections of the 
collectors or their Institutions [Dr Deeming - National Museum of Wales, Cardiff, UK 
(NMWC); Dr P. Gatt - Rabat, Malta; Dr B. Merz - Museum of Natural History, 
Geneva, Switzerland (MHNG); and in the author's collection (MJE)]. 

In the course of this study a number of specimens similar to A. claridgei Ebejer 
could not be identified with certainty. Detailed examination showed these to belong to 
a new species and this is described below. 

For the indentification to species, the reader is referred to the following litera- 
ture: for the genus Chyromya - Andersson, 1971, 1976; for Gymnochiromyia - Soós, 
1979, Ebejer, 1998a; and for Aphaniosoma - Collin, 1949; Ebejer, 1993, 1998b, Carles- 
Tolrâ, 2001. 

The nomenclature of the hypopygium that is used in this article takes into 
account the terminology given in the Contributions to a Manual of Palaearctic Diptera 
(Sinclair, 2000). The term gonostylus is retained for the time being, since there are un- 
certainties of homologies with regard to this structure. It is so identified because it 
articulates with the posterior end of the hypandrium and in many Aphaniosoma it bears 
sensory setulae (absent in the species). The term corresponds to terminology used in 
recent literature on the Chyromyidae (Ebejer. 1998a, b). 



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CHYROMYIDAE OF ITALY 861 



RESULTS 

Chyromya flava (Linnaeus, 1758) 

Material examined. ITALY: Sicily: 33 S , 2 9 9 , Catania, Randazzo, Monte Spagnolo, 
1200 m, grassland, edge of woodland of Acer, Castanea, Pinus, 6.vi.l999, J.C. Deeming; 1 2 , 
same data, but M.J. Ebejer; 1 6 , 8 9 9 , same data, but B. Merz; 1 8 and 1 9 , Messina, Nebrodi, 
Cesare, 1150 m, on Tilia, 8.vi.l999, J.C. Deeming; 1 9 , Messina, Nebrodi, Troina, Elia River, 
1000 m, 8.vi.l999, M.J. Ebejer; 8c? o\ 129 9 , Catania, Randazzo, Lago di Gurrida, Quercus, 
Populus, 870 m, 11. vi. 1999, M.J. Ebejer; 3<?o\59 9, same data, but B. Merz. 

This is one of the commonest and most widespread species in the family. It is 
also known from North America and North Africa. It has been observed in good num- 
bers on and beneath leaves of isolated broad-leaved trees, especially Tilia and Populus, 
growing at the edge of open woodland. New to Sicily. 

Chyromya miladae Andersson, 1976 

Material examined. ITALY: Sicily: 1 9 , Catania, Randazzo, Lago di Gurrida, Quercus, 
Populus, 870 m, 11. vi. 1999, M.J. Ebejer. 

Described from the Czech Republic and later recorded from Slovakia, this 
species was recently listed as occurring in Britain (Chandler, 1998). This uncommon 
species is probably associated with broad-leaved woodland. New to Sicily. 

Chyromya oppidana (Scopoli, 1763) 

Material examined. ITALY: Sicily: 1 9 , Catania, Randazzo, Lago di Gurrida, Quercus, 
Populus, 870 m, ll.vi.1999, M.J. Ebejer. 

This species is as frequent as flava and it has the same wide distribution and 
habits. New to Sicily. 

Gymnochiromyia fallax Ebejer, 1998 

Material examined: ITALY: Sardinia: 59 9, Nuoro reg., 120 m, 8 km E of Oliena, 
Hotel su Gologone, 1 6. vi .2002, B. Merz & M. Eggenberger. 

This species is typically found in saltmarshes and coastal regions. It was 
described from Malta, but it is also found in Mallorca. New to Sardinia. 

Gymnochiromyia flavella (Zetterstedt, 1848) 

Material examined: ITALY: Sardinia: 19, Baunei reg., Santa Maria Navarrese, 
18.vi.2002, B. Merz & M. Eggenberger. 

Widespread in Europe and probably all the Palaearctic including North Africa. 
New to Sardinia. 

Gymnochiromyia inermis (Collin, 1933) 

Material examined: ITALY: Sardinia: 1 9 , Villacidro reg., 600 m, Nuxeddu Mts., 
Montimannu forest, 12. vi .2002, B. Merz & M. Eggenberger; 19, Nuoro reg., Monte Albo 
1120 m, Punta Catirina, 16.vi.2002, B. Merz & M. Eggenberger. Sicily: 2c? 6 , 3 9 9 , Messina, 
Nebrodi, Troina, Lago D'Ancipa, mixed woodland, 8.vi.l999, M.J. Ebejer; 19 same data, 
B. Merz; 1 9 , Catania, Randazzo, Monte Spagnolo, 1200 m, grassland, Acer, Castanea, Pinus, 
6.vi.l999, M.J. Ebejer; 19, same data, but B. Merz; 1 9, Messina, Bronte, Monte Minardo, 
Quercus forest, 345 m, 11 .vi. 1999, M.J. Ebejer; 1 9 , same data, but J.C. Deeming. 



862 M - J- EBEJER 

When Collin (1933) described this species, he gave a number of examples of 
birds' nests from where it was reared. Deeming (1998) described the puparium and 
cephalopharyngeal skeleton. It is a common species and it is the most likely member 
of the genus to be found at some altitude and in broad-leaved woodland. New to 
Sardinia and Sicily. 

Gymnochiromyia mihalyii Soós, 1979 

Material examined: ITALY: Sardinia: 1 o\ 5 2 2, Nuoro reg., Monte Albo 1120 m, 
Punta Catirina, 16.vi.2002, B. Merz & M. Eggenberger; ló\ Dorgali, Rio Flumineddu, Gola di 
Gorropu, 15.vi.2002, B. Merz & M. Eggenberger. Sicily: 1 2 , Catania, Etna, 3 km NW of Milo, 
Castanea, Corylus, 1000 m, 5.vi.l999, M.J. Ebejer. 

Carles-Tolrâ (1992) recorded this species from Spain and Merz (1997) recorded 
it from Switzerland. The present author has specimens from Britain, Turkey and 
Greece. It is probably widespread in several Mediterranean countries. Unlike inermis, 
it seems to prefer open countryside including dunes and cultivated fields. New to 
Sardinia and Sicily. 

Aphaniosoma bifalcatum sp. n. Figs 1-3 

Material examined: Holotype: S , ITALY: Sardinia: Dorgali reg., 350 m, Cala Luna 
South of Cala Gonone, 1 7. vi. 2002, B. Merz & M. Eggenberger (MHNG). Paratypes: lo", 22 2, 
same data and depository; lo*, same data, but in MJE; le?, Bosa reg., m, Bosa Marina, 
20.vi.2002, B. Merz & M. Eggenberger (MHNG). Sicily: 1 ó\ 12, Siracusa, Noto, Vendicali, 
coastal marshes and dunes, M.J. Ebejer (MJE). FRANCE: 26 6 , 42 2 , Montpellier, Lotte salt- 
marsh, 23.viii.2000, J.C. Deeming (NMWC). 

A dark brownish black species of the approximatum Becker group characterized 
by a pair of long frontal setae anterior to the ocellar triangle and long narrow dark 
surstyli. 

Holotype, male. Head: yellow, ocellar triangle black, occiput deep black except 
for narrow, yellow postocular margin; frons at vertex about half width of head; frons, 
antenna, gena and mouth parts all yellow; third antennal segment with distinct setulae 
anteriorly, arista black except for yellow basal segments; gena about half as high as 
eye; chaetotaxy: a pair of very long frontal setae present on frons in front of ocellar 
triangle and 8 pairs of pale setulae scattered on frons, 2 long orbitals and 2 shorter ones 
in front of these, 1 internal and 1 external vertical. 

Thorax: black with dark grey dusting becoming brownish in prescutellar de- 
pression and on scutellum; scutellum with pale apical margin; humerus and notopleur- 
al depression clear yellow; pleura all dark brownish black except for narrow yellow 
margins to sutures; metanotum black; chaetotaxy: 1 humeral, 1 posthumeral, 2 noto- 
pleural, the anterior, strongly developed - about twice as long as posterior; 2+5 dorso- 
centrals, but only the prescutellar strong, 2+4 acrostichals, 3 intra-alar, 1 post-alar and 
supra- alar, 2 pairs of marginal scutellars; 1 mesopleural with 4 short setulae in 
vertical row below this, 1 sternopleural at posterior upper corner, and 4 setulae in 
vertical row at anterior part; stigmatical setula present. Wing: broad, hyaline with 
anterior cross vein pale yellow and other veins all brownish, becoming darker towards 
apex; distance between cross veins about 1.2 times length of posterior cross vein; 
distance between R 2+ 3 and R 4+5 on costa is half that between R 4+5 and M 1+2 . Haltère: 



CHYROMYIDAE OF ITALY 



863 




gon 



bac sci 



Figs 1-3 
Aphaniosoma bifalcatum sp. n., male terminalia. 1, hypopygium lateral aspect; 2, pregenital 
sterilite; 3, hypopygium posterior aspect; abbreviations: bac sci = bacilliform sclerite, ep = 
epandrium, gon = gonostylus, hyp = hypandrium, surs = surstylus, t6 = sixth tergite; scale bar 
= 0.1mm. 



864 M - J - EBEJER 

yellow. Legs: all yellow except for 5 th tarsal segment of all legs - this being broadly 
infiascateci around margin; hind coxa simple and brownish at base. 

Abdomen: tergites black with narrow pale posterior margins; epandrium with 
shiny pale brown spot at base near cerei, which are relatively long, yellow and pale- 
haired. 

Hypopygium (Figs 1-3): typical of the approximatum Becker - creperum Collin 
group, having an internalised and reduced epandrium with prominent dark surstyli 
visible externally; distiphallus poorly sclerotized and relatively amorphous; bacilliform 
sclerite and gonostylus prominent, darkly pigmented and heavily sclerotized; gono- 
stylus articulates with posterior arm of hypandrium very close to where this articulates 
with inferior margin of epandrium. 

Female. As in male, but without secondary sexual characters; apex of abdomen 
with tergite 7 strongly curved on sides to form a broad and short conical cavity with 
the cerei deep within; apical 2 sternites, in middle third, heavily sclerotized and dark 
shiny brown pigmented. 

Variation. The males from France and the one from Bosa in Sardinia have a 
greyish yellow scutellum and a dusky yellow prescutellar area; the grey mesonotal 
pattern fades posteriorly except in the middle where it reaches the scutellar margin and, 
laterally, the yellow area extends forwards along the intra-alar line; the pleura have the 
sutures more broadly yellow and the hind marginal bands of the tergites are broader, 
especially laterally. The colour variations are present also in the females from France. 
A male paratype from France was dissected and is identical to the paratype from 
Sardinia, which is illustrated in the figure. 

Similar species. The nominate form resembles the dark species of the claridgei 
Ebejer group, whereas the paler form resembles grisescens Ebejer and proximum 
Ebejer. If the surstyli are properly extruded, then bifalcatum can be recognised because 
these structures are sinuous and the apex is somewhat spatulate. 

Etymology. The curved surstylus together with the sinuate gonostylus gives an 
appearance of two sickles (Latin singular: falx) on each side of the postabdomen. 

Aphaniosoma claridgei Ebejer, 1995 

Material examined: ITALY: Toscana: 3 6 6 , 7 9 9 , Parco Nazionale della Maremma, 
Collelungo, Bonifica Canal, 2.ix.2003, P. Gatt; 26 6, 159 9, plus 89 9 in alcohol, Parco 
Nazionale della Maremma, Bocca D'Ombrone estuary, saltmarsh, 2.ix.2003, P. Gatt; 3 S â , 4 9 9 , 
Parco Nazionale della Maremma, Collelungo. beach and dunes, 2.ix.2003, P. Gatt. Sardinia: 
3c? <? , 1 9 , Dorgali reg., 350 m, Cala Luna South of Cala Gonone, 17.vi.2002, B. Merz & M. 
Eggenberger; 3 e? e? , 3 9 9 , Bosa reg., m, Bosa Marina, 20.vi.2002, B. Merz & M. Eggenberger; 
13c? <?, 39 9, Cuglieri Reg., Sinis Peninsula, m, Capo Mannu, 21.vi.2002, B. Merz & M. 
Eggenberger, Sicily: 6c? c?, 39 9, Siracusa, Noto, Vendicari, coastal marshes and dunes, M.J. 
Ebejer; llc?c?, 169 9 , same data, but B. Merz; 2c?c?, 59 9 , same data, but J.C. Deeming. 

A typical salt marsh species with a preference for coastal areas, it is widespread 
in the Mediterranean. It is often the dominant species of the genus in such habitats. 
There are many similar species in this group. New to Italy: Sardinia and Sicily. 

Aphaniosoma collini Lyneborg, 1973 

Material examined: ITALY: Sardinia: 2c? c? , Bosa reg., Bosa Marina, m, 20.vi.2002, 
B. Merz & M. Eggenberger. 



CHYROMYIDAE OF ITALY 865 

This species was described from Spain. It is common in Tunisia, Mallorca, 
France and Malta and may be one of the more widespread species in the group with 
projections on the hind trochanter in the male. New to Sardinia. 

Aphaniosoma egregium Ebejer, 1998 

Material examined: ITALY: Sicily: 1 6 , Siracusa, Noto, Vendicali, coastal marshes and 
dunes, J.C. Deeming (MJE). 

This is a distinctive, but one of the most uncommon species of Aphaniosoma. 
Other than the types, very few specimens of this species have been seen. New to Sicily. 

Aphaniosoma grisescens Ebejer, 1998 

Material examined: ITALY: Sardinia: 20 S , Dorgali reg., 350 m, Cala Luna South of 
Cala Gonone, 17.vi.2002, B. Merz & M. Eggenberger; 1 6 , Baunei reg., Santa Maria Navarrese, 
18.vi.2002, B. Merz & M. Eggenberger; 26 6 , 1 9 , Cuglieri Reg., Sinis Peninsula, m, Capo 
Mannu, 21.vi.2002, B. Merz & M. Eggenberger; 36 6, Bosa reg., m, Bosa Marina, 20.vi.2002, 
B. Merz & M. Eggenberger. 

Described from Tunisia, it is also known from Malta. Not a common species, 
but it can easily be confused with claridgei and proximum. New to Sardinia. 

Aphaniosoma melitensis Ebejer, 1993 

Material examined: ITALY: Toscana: 16 , 29 9, Parco Nazionale della Maremma, 
Collelungo, Bonifica Canal, 2.ix.2003, P. Gatt; 1 9 , Parco Nazionale della Maremma, Bocca 
D'Ombrone estuary, saltmarsh, 2.ix.2003, P Gatt. Sardinia: 26 6 , 8 9 9 , Dorgali reg., 350 m, 
Cala Luna South of Cala Gonone, 17.vi.2002, B. Merz & M. Eggenberger; 29 9, Baunei reg., 
Santa Maria Navarrese, 1 8. vi. 2002, B. Merz & M. Eggenberger; 8ó* 6 ,1 9 , Bosa reg., m, Bosa 
Marina, 20.vi.2002, B. Merz & M. Eggenberger; 16 , 19, Cuglieri Reg., Sinis Peninsula, m, 
Capo Mannu, 21.vi.2002, B. Merz & M. Eggenberger. Sicily: 36 6, 3 9 9, Siracusa, Noto, 
Vendicali , coastal marshes and dunes , M . J . Ebejer; 2 6 6 , 16 9 9 , same data , but B . Merz ; 2 6 6 , 
8 9 9, same data, but J.C. Deeming. 

This is another common species known from several European countries where 
it inhabits flushed meadows, marshes and dunes. It is a polymorphic species with small 
very dark forms appearing quite different from the large pale specimens. There is also 
variation as to how rounded are the apical projections on the ventral aspect of the 
fourth tergite in the male. All intermediates exist within the same population if a large 
enough sample is examined. New to Sicily and Sardinia. 

Aphaniosoma micromacro Carles-Tolrâ, 2001 

Material examined: ITALY: Toscana: 2 c? o* , 69 9 , Parco Nazionale della Maremma, 
Collelungo, beach and dunes, 2. ix. 2003, P. Gatt. 

Described from Spain, this species is one of the very few with modified tarsi. It 
is also known from Malta. New to Italy. 

Aphaniosoma propinquans Collin, 1949 

Material examined: ITALY: Sicily: 26 6, 29 9, Siracusa, Noto, Vendicari, coastal 
marshes and dunes, M.J. Ebejer. 

Originally described from Britain, this species is now known to have a wide dis- 
tribution in Europe. It is relatively easy to identify in both sexes and so it should not 
be overlooked. New to Sicily. 



866 M - J - EBEJER 

Aphaniosoma proximum Ebejer, 1998 

Material examined: ITALY: Sardinia: 1 9 , Bosa reg., m, Bosa Marina, 20.vi.2002, 
B. Merz & M. Eggenberger; 26 6, Cuglieri Reg., Sinis Peninsula, m, Capo Mannu, 
21 .vi.2002, B. Merz & M. Eggenberger. Sicily: 66 6 ,79 2 , Siracusa, Noto, Vendicali, coastal 
marshes and dunes, M.J. Ebejer; 1 6 , 4 2 9 , same data, but B. Merz. 

This species belongs to the approximatum Becker group. It is difficult to iden- 
tify without dissection. It is probably widespread in the Mediterranean, but overlooked. 
New to Sardinia and Sicily. 

ACKNOWLEDGEMENTS 

I would like to thank my colleagues Dr John Deeming (Cardiff, UK), Dr Paul 
Gatt (Rabat, Malta) and Dr Bernhard Merz (Geneva. Switzerland) who kindly put at 
my disposal all their material of Chyromyidae. I thank the authorities of the Parco dei 
Nebrodi (Sicily) for allowing access and collecting in the National Park (permit 
number 3294). Dr Gatt collected in the Parco Nazionale della Maremma as a parti- 
cipant in the MECO Project. Finally, I am grateful to Dr Giorgio Sabella of the 
University of Catania for sparing his time to discuss with us the best sites to visit and 
for arranging permission to collect on Etna and at Vendicari. 

REFERENCES 

Andersson, H. 1971. The Swedish species of Chyromyidae (Diptera), with lectotype 

designations. Entomologisk Tidskrift 92: 95-99. 
Andersson, H. 1976. Chyromya miladae n.sp. from Czechoslovakia (Diptera: Chyromyidae). 

Entomologica scandinavica 7: 187-189. 
Canzoneri, S., Gorodkov, K., Krtvosheina, N.P., Munari, L., Nartshuk, E., Papp, L. & Süss, 

A. 1995. Diptera Opomyzoidea, Carnoidea, Sphaeroceroidea. In: Mtnelli, A., Ruffo, S. 

& La Posta, S. (eds). Checklist delle specie della fauna Italiana 75, Bologna, Calderini, 

27 pp. 
Carles-Tolrâ, M. 1992. New and interesting records of Diptera Acalyptrata from Spain. Part 

II: Heleomyzidae, Trixoscelididae, Chyromyidae. Curtonotidae, Camillidae, Diastatidae 

and Campichoetidae. Graellsia 48: 19-24. 
Carles-Tolrâ, M. 2001. Eight new chyromyid species from Spain (Diptera, Chyromyidae). 

Boletin de la Asociación espaiiola de Entomologia 25(3-4): 45-62. 
Chandler, P. 1998. Chyromyidae (p. 151). In: Chandler, P. J. (ed.). Checklists of Insects of the 

British Isles (New Series), Part 1: Diptera. Handbooks for the Identification of British 

Insects 12: iii-xx + 1-234. Royal Entomological Society, London. 
Collin, LE. 1933. Five new species of Diptera. Entomologists monthly Magazine 69: 272-275. 
Collin, J.E. 1949. On Palaearctic Species of the Genus Aphaniosoma . Annals and Magazine of 

Natural Histoiy (12) 2: 127-147. 
Czerny, L. 1927. 53c. Chiromyidae (pp. 51-54). In: Lindner, E. (ed.). Die Fliegen der 

Paläarktischen Region, Vol. 5. E. Schweizerhart' sehe Verlagsbuchhandlung, Stuttgart, 

56 pp. 
Deeming, J.C. 1998. A description of the puparium of Gymnochiromyia inermis (Collin, 1933) 

from England (Diptera: Chyromyidae). Studia Dipterologica 5 (1): 30-32. 
Ebejer. M.J. 1993. The Chyromyidae of Malta with descriptions of new species of Aphaniosoma 

(Dipt.. Schizophora). Entomologists Monthly Magazine 129: 125-133. 
Ebejer. M.J. 1998a. A new species of Gymnochiromyia Hendel (Diptera: Chyromyidae) from 

the Mediterranean, with notes, lectotype designations and a key to the species from the 

West Palaearctic. Studia Dipterologica 5 (1): 19-29. 



CHYROMYIDAE OF ITALY 867 



Ebejer, M.J. 1998b. A Review of the Palaearctic Species of Aphaniosoma Becker (Diptera, 

Chyromyidae), with Descriptions of New Species and a Key for the Identification of 

Adults. Mitteilungen Museum für Naturkunde Berlin, Deutsche Entomologische 

Zeitschrift 45 (2): 191-230. 
Merz, B. 1997. Die Megamerinidae, Strongylophthalmyiidae, Pseudopomyzidae, Chyromyidae 

und Camillidae der Schweiz (Diptera, Acalyptrata). Mitteilungen der entomologischen 

Gesellschaft Basel 47(4): 1 30- 1 38 . 
Sinclair, B. 2000. Morphology and terminology of Diptera male terminalia, 1 .2. (pp. 53-74). In: 

Papp, L. & Darvas, B. (eds). Contributions to a Manual of Palaearctic Diptera. Vol. 1. 

Science Herald, Budapest, 978 pp. 
Soós, A. 1979. Einige neue Acalyptraten Museiden aus der Paläarktischen Region (Diptera). 

Acta Zoologica Academiae Scientiarum Hungaricae 35 (3-4): 409-414. 
Soós, A. 1984. Family Chyromyidae (pp. 56-60). In: Soós, A. & Papp, L. (eds). Catalogue of 

Palaearctic Diptera. Vol. 10. Akadémiai Kiado, Budapest, 402 pp. 



Revue suisse de Zoologie 112 (4): 869-941; décembre 2005 



Hemiscorpiidae (Scorpiones) from Iran, with descriptions of 
two new species and notes on biogeography and phylogenetic 
relationships 

Lionel MONOD 1 & Wilson R. LOURENÇ0 2 

1 Muséum d'histoire naturelle, route de Malagnou 1, case postale 6434, 
CH-1211 Genève 6, Switzerland. E-mail: liocheles@gmail.com 

2 Département de Systématique et Evolution, USM 0602 - CP 53, Section Arthropodes 
(Arachnologie), Muséum National d'Histoire Naturelle, 61 rue de Buffon, 
F-75005 Paris, France. E-mail: arachne@mnhn.fr 



Hemiscorpiidae (Scorpiones) from Iran, with descriptions of two new 
species and notes on biogeography and phylogenetic relationships. - The 

family Hemiscorpiidae is closely related to the Liochelidae. Within the 
Hemiscorpiidae, the Oriental species are particularly interesting. Most of 
them exhibit highly derived characters in comparison to their African rela- 
tives. Males possess a strongly elongated metasoma and a similarly elon- 
gated telson bearing a pair of tuberculiform processes at the base of the 
aculeus. Furthermore, Hemiscorpius lepturus Peters, 1861, which occurs in 
Iraq and Iran, is known to have an extremely virulent venom with cytotoxic 
and haemolytic components. It is responsible for severe dermonecrotic 
scorpionism in southern Iran. This is the only non-buthid scorpion that is 
potentially lethal. In this paper an overview of the species of Hemiscorpius 
in Iran is presented with revised diagnoses and descriptions. Two new 
species from western Iran, H. enischnochela sp. n. and H. acanthocercus 
sp. n., are described. The genus Habibiella Vachon, 1974 is synonymised 
with Hemiscorpius Peters, 1861. A thorough analysis of hemisperma- 
tophores shows close phylogenetic relationships with several genera of the 
family Liochelidae. A hypothesis on the geological events that probably 
triggered the present distribution of Hemiscorpius is finally proposed. 

Keywords: Scorpiones - Hemiscorpiidae - Hemiscorpius - new species - 
new synonymy - Iran - hemispermatophore - Liochelidae. 

INTRODUCTION 

The family Hemiscorpiidae Pocock, 1893 was formerly considered as a sub- 
family (Hemiscorpiinae) of the Scorpionidae Latreille, 1802 (Fet, 2000). Sissom 
(1990) criticised the placement of the Hemiscorpiinae in the Scorpionidae, but did not 
suggest an alternative. Stockwell (1989) first suggested the transfer of Hemiscorpiinae 
directly from the Scorpionidae to the Ischnuridae, where they retained their subfamily 
rank. Recently Lourenço (2000) and Prendini (2000) simultaneously elevated the 



Manuscript accepted 11.08.2005 



870 L- MONOD & W. R. LOURENÇO 

Hemiscorpiinae to family level. While Lourenço did not give any justification, 
Prendini provided a detailed phylogenetic analysis where he demonstrated that 
Hemiscorpiidae is the sister group of Ischnuridae Simon, 1879, now Liochelidae Fet & 
Bechly, 2001 . Solegad & Fet (2003) confirmed the phylogeny of Scorpionoidea estab- 
lished by Prendini (2000), but downgraded the Hemiscorpiidae from family to sub- 
family rank under Liochelidae. Until the discussion is settled, we decided to follow 
Lourenço 's and Prendini 's view and consider the hemiscorpiids as a family. 

Two genera were traditionally listed in the family Hemiscorpiidae (see Fet, 
2000), Habibiella Vachon, 1974 and Hemiscorpius Peters, 1861a. Habibiella is a 
monotypic genus, the type species, H. gaillardi, was described by Vachon (1974) from 
a single female collected in eastern Iran. This genus is here placed in the synonymy of 
Hemiscorpius. The genus Hemiscorpius in the traditional sense includes six species 
which occur from Somalia to Pakistan, most of them in the Middle East (Fet, 2000). 
These are: H. arabicus (Pocock, 1899a), H. Upturns Peters, 1861a, H. maindroni 
(Kraepelin, 1900), H. persicus Birula, 1903, H. socotranus Pocock, 1899b and H. 
tellinii Borelli, 1904. Species of Hemiscorpius and Habibiella gaillardi (now under 
Hemiscorpius) are morphologically very similar and were distinguished only by their 
trichobothriotaxy. Habibiella is neobothriotaxic major, with 10-12 trichobothria on the 
ventral side of the pedipalp patella instead of 3 as in the species previously comprising 
Hemiscorpius, and 15 trichobothria on the external side of the pedipalp patella instead 
of 13. 

Three species belonging to the family Hemiscorpiidae are known to occur in 
Iran: H. gaillardi, H. lepturus and H. persicus. H. lepturus is of particular medical 
interest. This species is the only dangerous and potentially lethal scorpion that does not 
belong to the family Buthidae. It is responsible for significant scorpionism problems in 
the southern provinces of Iran. The venom of H. lepturus is highly cytotoxic and 
haemolytic and can cause serious wounds and skin inflammations whereas other scor- 
pions have a neurotoxic venom. The deep dermonecrotic ulcers and blisters caused by 
such a sting are slow and difficult to heal and therefore usually result in very un- 
pleasant scars. Severe complications such as serious haemolysis, internal haemor- 
rhages, secondary renal failure and death were also reported (Radmanesh, 1990, 1998). 
Venoms with such cytotoxic and hemolytic effects are also quite uncommon in spiders 
and only known in three genera, i. e. Cheiracanthium C.L. Koch, 1839 (Miturgidae), 
Loxosceles Heineken & Lowe, 1832 (Sicariidae) and Sicarius Walckenaer, 1847 
(Sicariidae), (Filmer, 1999). While severe envenomations in humans by Cheira- 
canthium and Sicarius are not known, the virulent poison of the Brown Recluse Spider, 
Loxosceles reclusa Gertsch & Mulaik, 1940, and its enzymatic activities have been 
extensively studied (Anderson, 1998; Butz et al., 1971; Dillaha et al., 1964; Foil étal., 
1979; Patek et al., 1994). Among the 13 species of Loxosceles present in the United 
States (at least 5 of them are associated with necrotic arachnidism), L. reclusa is most 
commonly responsible for dermonecrotic arachnidism in North America. Brown spi- 
ders are also of significant clinical concern in South America, with L. laeta (Nicolet, 
1 849) being responsible for several deaths each year. In Iran cases of scorpion enven- 
omation with cutaneous and viscerocutaneous reactions are usually attributed to the 
species H. lepturus. Species of Hemiscorpius are morphologically very close to each 



HEMISCORPIIDAE FROM IRAN 871 

other and difficult to distingish for a non-specialist, therefore we assume that H. lep- 
turus is probably not the only species responsible for all envenomations in that coun- 
try. Other Hemiscorpius species probably possess venoms with similar necrotic effects. 
As stated by Fet (2000), there is no recent revision and no dichotomic key for 
Hemiscorpius. In the past 40 years several checklists of scorpions from Iran, which 
also include Hemiscorpius species, have been published (Vachon, 1966; Habibi, 1971; 
Farzanpay & Pretzmann, 1974; Pérez-Minnocci, 1974; Farzanpay, 1988; Kinzelbach, 
1985; Kovafik, 1997). However, neither precise descriptions, nor specific diagnoses 
were included in these publications. The extensive cladistic analysis of Scorpionoidea 
by Prendini (2000) gives a precise diagnosis and description for the family, but not for 
the two species used in that study, i. e. H. lepturus and H. maindroni. Early descrip- 
tions are very short and not accurate enough, and therefore do not allow species iden- 
tification. The present paper gives an overview of the hemiscorpiids of Iran, with 
revised diagnoses and redescriptions of H. gaillardi, H. lepturus and H. persicus. H. 
enischnochela sp. n. and H. acanthocercus sp. n. are described from the region of 
Bandar Abbas, Hormozgan Province, south-eastern Iran. This study also allowed to 
clarify the taxonomic position of the enigmatic genus Habibiella, which is here 
synonymised with the genus Hemiscorpius . Strong evidence of a close phylogenetic 
relationship with the liochelids Iomachus politus Pocock, 1896, Hadogenes Kraepelin, 
1894 and Opisthacanthus (Opisthacanthus) Peters, 1861b is pointed out by the 
examination of hemispermatophore morphology. Finally a hypothesis on the geolo- 
gical events that presumably triggered the evolution and present distribution of the 
family is proposed. 



MATERIAL AND METHODS 

Illustrations were produce by using a Wild M5 stereo-microscope with a draw- 
ing tube. Trichobothrial notations and terminology of metasomal carination follow 
Vachon (1974), measurements follow Stahnke (1970) and are given in mm. Additional 
morphological terminology mostly follows Hjelle (1990) and Sissom (1990), termi- 
nology of carination of pedipalpal chela follows Prendini (2000). Hemispermatophore 
terminology is modified from the terminology applied by Lamoral (1979) and was used 
by Monod & Volschenk (2004). The distribution map was generated with Arc View® 
GIS 3.1, maps and drawings were subsequently edited in Adobe Illustrator® 8.0 and 
Adobe Photoshop® 6.0. 

List of acronyms 

BPBM Bernice P. Bishop Museum, Honolulu, Hawaii. 

CAS Californian Academy of Sciences, San Fransisco, USA. 

MHNG Muséum d'histoire naturelle, Genève, Switzerland. 

MNHN Muséum National d'Histoire Naturelle, Paris, France. 

NHMW Naturhistorisches Museum Wien, Austria. 

ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. 

ZMB Zoologisches Museum, Humboldt Universität, Berlin, Germany. 



872 L - MONOD & W. R. LOURENÇO 

SYSTEMATICS 

Hemiscorpiidae Pocock, 1893 

Hemiscorpinii Pocock, 1893: 306, 308. Type genus Hemiscorpius Peters, 1861a. 

Diagnosis (modified from Prendini, 2000). The Hemiscorpiidae can be distin- 
guished from all other scorpionoid taxa by their trichobothriotaxy: trichobothria ib and 
it of the pedipalp chela are situated midway on the finger, whereas both are at the base 
of the fixed finger in all other scorpionoid taxa except Nebinae and ib is situated basally 
and it distally on the fixed finger in Nebinae, Diplocentridae. Hemiscorpiids also differ 
from other scorpionoid taxa, except for Heteroscorpionidae and Urodacidae, by their 
metasomal segments I-IV, which bear a single ventromedian carina equally developed 
on all segments. Hemiscorpiidae can be distinguished from the Heteroscorpionidae and 
Urodacidae by the following characters: (1) 3 pairs of lateral ocelli present; (2) median 
ocular tubercle shallow; (3) movable cheliceral finger with distal external and distal 
internal teeth approximately equal in size and closely opposed; (4) pedipalp chela 
fingers with 2 primary rows of denticles often fused at the base; (5) trichobothrium db 
located on internal surface of fixed finger; (6) trichobothrium Db located basally on 
dorsal surface of manus; (7) trichobothrium Dt located at proximal end of fixed finger; 
(8) no accessory trichobothria in v series of chela; (9) telson vesicle of male elongated 
and laterally compressed; (10) hemispermatophores with a double lamellar hook and a 
complex median capsular structure. 

Remarks. Telson vesicles of mature Hemiscorpius leturus males are unusually 
elongated, laterally compressed, and bear a pair of distal lobes at the base of aculeus 
(Fig. 1 A-B), whereas females possess bulky telsons, which are laterally not compressed 
and have no distal lobes. Subadult males do not have an extremely elongated metasoma 
and a modified telson like mature males have, their metasoma and telson are just slight- 
ly more elongated than in females. These secondary sexual characters appear only in the 
last developmental instar. In Prendini's (2000) cladistic interpretation of relationships 
within the superfamily Scorpionoidea this very unusual feature is considered as a valid 
character for the whole family Hemiscorpiidae and is mentionned in the diagnosis: tel- 
sons of H. lepturus and H. maindroni males are described as possessing an extreme 
elongation and paired distal lobes. Our examination of syntypes of Hemiscorpius main- 
droni from Mascate also revealed a pronounced sexual dimorphism but not as extreme 
as in H. lepturus. Indeed H. maindroni males possess telsons (Fig. 1G-H) without any 
distal lobes or elongation as in H. lepturus. Therefore the species identified as H. main- 
droni in Prendini's cladistic revision of the superfamily Scorpionoidea (2000) undoubt- 
edly belongs to another species. All the other non-Iranian hemiscorpiids examined, i. e. 
H. socotranus, Hemiscorpius spp. from Somalia, Eritrea and Djibouti possess a simple 
telson and a less elongated metasoma than in H. maindroni (Fig. 27 A-B). The Iranian 
hemiscorpiids H. acanthocercus sp. n., H. enischnochela sp. n., H. lepturus and pro- 
bably also H. gaillardi can therefore be placed together in the H. lepturus species 
complex. However, the Iranian H. persicus does not belong to this group. The mature 
male holotype studied does not have modified metasoma and telson. 

Prendini (2000) described the hemispermatophores of hemiscorpiids as 
possessing a single lamellar hook. In his data matrix (Table 3, p. 6), the character 90 



HEMISCORPIIDAE FROM IRAN 873 

(hemispermatophore, lamellar hook) is coded (single hook) for both H. lepturus and 
H. maindroni. Mature males in the material examined have been systematically dis- 
sected in order to remove and study their hemispermatophores. Several hemisper- 
matophores in excellent shape of//, lepturus, H. acanthocercus and H. maindroni have 
been obtained. All of them bear a double lamellar hook, a very unusual feature that was 
so far only recorded for the liochelid genus Hadogenes. Unfortunately this character 
cannot be assessed for H. enischnochela sp. n., H. persicus and H. socotranus. The 
only available mature male of H. enischnochela had been dissected before but the 
hemispermatophore was not found inside the vial and is probably lost. We were not 
allowed to dissect the type of H. persicus and no other specimens of this species were 
available to us. The only available mature male of//, socotranus had poorly preserved 
paraxial organs from which it was impossible to extract hemispermatophores. 
Nevertheless the character "double lamellar hook" is used in our diagnosis of the 
family. In liochelids morphometric proportions of hemispermatophores and shape of 
the capsular median structure are usually the only features that vary between genera. 
The single lamellar hook is a constant character within the family, except for 
Hadogenes, species of which possess hemispermatophores with a double lamellar 
hook. According to Prendini (2000) Hadogenes and Cheloctonus are the more basal 
liochelid genera and Hemiscorpiidae is the basal sister- group of Liochelidae. The 
double lamellar hook can therefore be considered as a plesiomorphy for Hemi- 
scorpiidae and Liochelidae, and the single lamellar hook is synapomorphic for 
Liochelidae except Hadogenes. Therefore the probability that the double lamellar hook 
is present in the whole Hemiscorpiidae is strong. Furthermore a double lamellar hook 
was assessed for hemiscorpiids from Oman by Dr. Graeme Lowe (pers. comm.) who 
is currently carrying out a revision of the scorpion fauna of Oman (Lowe, in prep.). 
This is another argument for including the double lamellar hook into the family 
diagnosis. 

Hemiscorpius Peters, 1861 

Hemiscorpius Peters, 1861a: 426. Type species by monotypy Hemiscorpius lepturus Peters, 

1861a. 
Habibiella Vachon, 1974: 952, syn. n. Type species Habibiella gaillardi Vachon, 1974. 

Remarks. Prendini (2000) suggested in his revision of the superfamily 
Scorpionoidea that Habibiella should be considered as a synonym of Hemiscorpius, 
because the monophyly of Hemiscorpius was not supported in any of his cladistic 
analyses. However, he did not formally propose this taxonomic change and listed both 
Habibiella and Hemiscorpius in the family Hemiscorpiidae. The genus Habibiella is 
monotypic and only known from a single female. It can be distinguished from 
Hemiscorpius only on the basis of its trichobotriotaxy, 15 external trichobothria on the 
pedipal patella instead of 13, and 10-12 ventral trichobothria on the pedipalp patella 
instead of 3 . The study of a mature male from the Natural History Museum of Vienna 
that belongs to this genus and that is described here under Hemiscorpius enischnochela 
sp. n. allowed to re-evaluate the taxonomic status of Habibiella. Metasoma and telson 
of this specimen possess exactly the same morphology encountered in Hemiscorpius 
lepturus, i.e. extremely elongated metasoma, vesicle strongly elongated and bearing a 



874 L. MONOD & W. R. LOURENÇO 

pair of blunt tuberculiform projections at the base of the short and strongly curved 
aculeus. The African and Arabian hemiscorpiids examined, i. e. H. maindroni and H. 
socotranus, do not have such features. Species of "Habibiella" are therefore more 
closely related to Hemiscorpius lepturus, the type species of Hemiscorpius , than H. 
lepturus is to the other Hemiscorpius species. Habibiella is therefore considered as a 
junior synonym of Hemiscorpius . 

Diagnosis. With Habibiella now in synonymy with Hemiscorpius , the family 
Hemiscorpiidae becomes monotypic and its diagnosis is also applicable to the genus. 

Hemiscorpius acanthocercus sp. n. Figs 1C-D, 2-7, 36 

Material examined. Holotype: 3 , IRAN (Hormozgan), Abad-Geno, 38 km N Bandar 
Abbas, 3.IV.1972, K. Bilek, NHMW 4718 (39). Paratypes: 1 o\ 1 9 , no locality specified, 
H. Löffler, Austrian Iran Expedition 1949-1950, MHNG. 2 S juv., 2 9 juv., no locality speci- 
fied, H. Löffler, Austrian Iran Expedition 1949-1950, NHMW. 1 6 juv., IRAN (Hormozgan), ca. 
50 km N Bandar Abbas, 4.IV.1970, E Ressi, NHMW 3393. 1 S juv., 1 9 juv., IRAN 
(Hormozgan), ca. 41 km N Bandar Abbas, 7.IV.1972, G. Pretzman & A. Konetschnig, NHMW 
3394. 1 6 subadult, 2 S juv., 2 9,19 juv.. IRAN (Hormozgan), 115 km E Bandar Abbas, 
27 .III. 1972, K. Bilek, NHMW4716. 1 S juv., 1 9 , IRAN (Hormozgan), 22 km N Bandar Abbas, 
4.IV.1972, K. Bilek, NHMW 4717. 2 9 juv., same data as for holotype, NHMW 4718. 1 o* juv., 
19,19 juv., IRAN (Hormozgan), 40 km N Bandar Abbas, 7.IV.1972, K. Bilek, NHMW 4719. 
1 6 subadult. IRAN (Hormozgan), 41 km N Bandar Abbas, 20.IV.1974, G. Pretzman, NHMW 
4720. 1 6 subadult, 2 9 , IRAN (Hormozgan), 65 km N Bandar Abbas, 30.HI. 1972, K. Bilek, 
NHMW 4721. 1 9 juv., IRAN (Hormozgan), 22 km N Bandar Abbas, 1974, G. Pretzman, 
NHMW 21142. 1 Ô imm., 19,19 juv., IRAN (Hormozgan), 38 km N Bandar Abbas, 
28.III.1972, K. Bilek, NHMW 21143. 2 9, IRAN (Hormozgan), 65 km N Bandar Abbas, 
4.IV.1972, K. Bilek, NHMW 21144. 

Distribution. Known from the surroundings of Bandar Abbas, Hormozgan 
Province, eastern Iran (Fig. 36). 

Etymology. The name acanthocercus is a construct from the Greek words akan- 
tha [thorn, prickle] and kerkos [tail]. The name is an invariable noun in apposition and 
refers to the metasomal dorsal carinae of males, which have numerous strong spiniform 
granules, and to the telson of males which also bears numerous small spiniform gran- 
ules, especially at the posterior extremity, on the blunt tuberculiform processes at the 
base of the aculeus. 

Diagnosis. (1) Carapace longer than wide, shagreened and finely granular, with 
small smooth patches; lateral margins with small spiniform granules below lateral 
ocular tubercles; superciliary carinae finely granular; (2) pedipalps stout and bulky, 
chela fingers slightly shorter than or equal in length to chela manus; (3) internal protu- 
berance of pedipalpal patella with internodorsal carina bearing 1-2 strong spiniform 
granules; (4) pedipalp patella orthobothriotaxic, external side with 13 trichobotria (1 
est and 2 esb), ventral side with 3 trichobothria; (5) metasoma of males elongated and 
slender, with dorsal carinae bearing numerous strong spiniform granules, and ventral 
and ventrolateral carinae of segments IV- V bearing spiniform granules; (7) telson of 
males strongly elongated, bearing a pair of blunt tuberculiform processes with small 
spiniform granules at base of aculeus; (8) metasoma of females with dorsal carinae of 
segments I-IV and ventral and ventrolateral carinae of segment V bearing strong spini- 
form granules; (9) hemispermatophore with strong double lamellar hook located above 
distal transverse ridge, very close to it, and pointing anteriad. 



HEMISCORPIIDAE FROM IRAN 



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876 L. MONOD & W. R. LOURENÇO 

H. acanthocercus sp. n. is closely related to H. lepturus, both of them have stout 
orthobothriotaxic pedipalps and pronounced sexual dimorphism. However, males of 
these 2 species can be readily distinguished by (1) carapace granulation, (2) metasoma 
carination, (3) morphology of telson, (4) pedipalp carination and shape, and (5) hemis- 
permatophore morphogy. The carapace of H. acanthocercus sp. n. has lateral margins 
bearing small spiniform granules below the lateral eyes, while in H. lepturus the later- 
al carapace margins are completely smooth. H. acanthocercus sp. n. has metasomal 
dorsal carinae bearing numerous strong spiniform granules, while H. lepturus has cari- 
nae with sparse, much reduced and weaker spiniform granules, especially on the ante- 
rior segments. The ventral and ventrolateral carinae of metasomal segments IV-V are 
furnished with spiniform granules in males of H. acanthocercus sp. n., but are almost 
smooth, with weak spiniform granules (carinae of segment V with more marked spini- 
form granules in the posterior half) in H. lepturus. In H. acanthicercus sp. n. the telson 
of males is less elongated and the aculeus is less curved than in H. lepturus. The later- 
al surfaces of the telson are fairly smooth, with only small granules, and distal protu- 
berances bear small spiniform granules in H. acanthocercus, while the lateral surfaces 
and distal projections are rugose, without distinct granules, in H. lepturus. Furthermore 
the distal telson projections are bigger in H. lepturus. The pedipalp chela manus in H. 
acanthocercus sp. n. is slightly thinner than in H. lepturus. H. acanthocercus sp. n. has 
intero-dorsal carina of the patellal prominence with a single strong spiniform granule 
(rarely 2), while H. lepturus has carinae with several much reduced, weaker granules. 
Females of these two species are very similar to each other, but can be distinguished 
by the superciliary carinae, which are very finely granular in H. acanthocercus sp. n., 
while smooth, or nearly so, in H. lepturus. Hemispermatophores of H. acanthocercus 
sp. n. possess a double lamellar hook that is stronger than in H. lepturus. The lamellar 
hook is located very close to the distal transverse ridge and points anteriad in H. acan- 
thocercus sp. n., while it is located more distally and points distad in H. lepturus. 

Description. MALE (Figs 1C-D, 2A-B, 3 A, 4B, 5A-C, E-F, 6A, 7) [description 
based on the holotype (NHMW 4718)]. Measurements of holotype (in mm) carapace, 
length 5.2; posterior width 4.9; distance between anterior lateral eyes 2.7, between 
posterior lateral eyes 3.2, between median eyes 0.2; diameter of median eyes 0.3. 
Pedipalp, femur length 5.0, width 1.8; patella length 5.0, width 1.7, chela length 9.7; 
manus length 5.0, width 3.8. depth 2.3; movable finger length 4.6; fixed finger length 
3.6. Metasoma. segment I length 5.2. width 1 .4; segment V length 7.4, width 1 .2, depth 
1 .4; vesicle length 4.6. width 1 .5. depth 1 .8, aculeus length 0.9. Total length 60.0. 

Carapace (Fig. 3A). Colouration mostly uniformly dark orange to brown; me- 
dian and lateral ocular tubercles black. Carapace dorsoventraly flattened, longer than 
wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle 
weakly developed, low, distinctly situated anteriorly, with weak, finely granular super- 
ciliary carinae; frontal concavity or notch well-developed; anterior lobes truncate; 
lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. 
Carapace shagreened and finely granular, with a few smooth patches; lateral margins 
with small spiniform granules below lateral ocular tubercles; antero-median furrow 
narrow, suturiform. anteriorly bifurcated; median longitudinal furrow shallow, conti- 
nuous from the anterior suture furcation, running through ocular tubercle posteriorly 



HEMISCORPIIDAE FROM IRAN 



877 




Fig. 2 
Hemiscorpius acanthocercus sp. n. Male paratype (MHNG, H. Löffler coll.): A, dorsal aspect; 
B, ventral aspect. Female paratype (NHMW4719): C, dorsal aspect; D, ventral aspect. Scale line, 
5 mm. 



878 L. MONOD & W. R. LOURENÇO 

into a deep triangular depression; postero-lateral furrow shallow; mesolateral furrow 
weakly developed, almost indiscernible. 

Mesosoma. Tergite colouration dark orange to brown anteriorly, becoming 
lighter posteriorly (dark yellow). Tergites I-III with a shallow median depression 
divided anteriorly by a weak ridge; tergites IV- VII with median carina reduced to a 
very weak ridge surrounded by a pair of shallow, submedian depressions. Lateral and 
sublateral carinae absent on tergites I- VI, but present on the posterior 2/3 of tergite VII 
as ridges with weak spiniform granules. Surface of tergites shagreened and finely gra- 
nular; pre-tergites smooth and minutely pitted. Sternite colouration dark yellow. 
Sterni tes III- VI smooth and minutely pitted, without granulation or carinae; sternite III 
with a pair of large, very finely and densely granular depressions, covered by the 
pectines; sternites IV- VI with a pair of shallow median furrows. Sternite VII with 
similar surface texture as on preceding sternites; pair of lateral carinae developed as 
ridges; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and 
sternites VII longer than wide. 

Metasoma (Fig. 4B) very elongated and slender, shagreened, with very few 
small scattered granules. Colour dark yellow to orange. Segments I-IV with longitu- 
dinal dorsomedian furrow, dorsal carinae with strong spiniform granules. Segments 
I-IV with lateral carinae developed as weakly granular ridges, lateral carinae on seg- 
ments III-IV less granular than those on segments I-II. Segments I-II with ventrolateral 
carinae as ridges with few weak granules, almost smooth; segments III-IV with 
ventrolateral carinae as ridges of reduced spiniform granules. Segment I: ventral carina 
indistinct, reduced to a weak ridge; segments II-III with ventral carina developed as a 
distinct ridge, almost smooth; segment IV with ventral carina as a ridge, anterior 1/3 
smooth and posterior 2/3 with weak spiniform granules. Segment V: longitudinal 
dorsal furrow present in anterior half, dorsal carinae with strong spiniform granules; 
lateral carina indistinct, visible as a row of small granules in anterior 2/3 only; ventro- 
lateral and ventromedian carinae with spiniform granules. 

Telson (Figs IC-D, 4B). Vesicle yellow to orange; aculeus darker, tan, due to 
stronger sclerotisation; vesicle elliptical or ovate, strongly elongated, with a blunt 
tuberculiform projection on each side at base of aculeus; ventrolateral furrows absent; 
ventromedian ridge absent; lateral surfaces smooth, with few weak granules; posterior 
projections with small spiniform granules; dorsal surface with numerous small spini- 
form granules and a median shallow depression. Few macrosetae basally, becoming 
more numerous near base of aculeus. Aculeus short and stout, strongly curved, 
becoming markedly narrower approximately midway. 

Chelicerae. Colour dark yellow; teeth of fingers darker orange. Teeth arrange- 
ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth 
bifid; movable finger with one subdistal tooth and one basal tooth in external series; 
distal external tooth smaller than distal internal tooth; cheliceral teeth without secon- 
dary serrations. 

Pedipalp stout and bulky. Pedipalp coxa and femur (Fig. 5A, E-F). Inter- 
noventral margin of coxa with strong spiniform granules. Dorsal surface of femur 
predominantly dark yellow-orange, carinae darker, orange. Femur stout (length less 
than 2.5 times its width), pentacarinate, with 4 distinct carinae; internodorsal carina 



HEMISCORPIIDAE FROM IRAN 



879 




PQ 







880 L - MONOD & W. R. LOURENÇO 

developed as a strongly and densely granular ridge, granules spiniform; externodorsal 
carina forming a ridge with spiniform granules; interno ventral carina developed as a 
granular ridge with spiniform granules; externoventral carina developed as a ridge with 
strong spiniform granules; ventromedian carina reduced to few granules and confined 
to base of femur. Dorsal surface shagreened, finely and densely granular, distal end 
smooth, without granules; internal surface shagreened, with a few strong, scattered, 
spiniform granules; external surface shagreened; ventral surface shagreened and 
weakly granular, distal end without granules. A total of 3 trichobothria present on 
femur; d located externobasally on dorsal surface; i located dorsobasally on internal 
surface; e located dorsobasally on external surface. Pedipalp patella (Fig. 5 A, E-F) 
stout and bulky (length less than 2.5 times its width). Colour predominantly dark 
yellow to orange; carinae slightly darker. 7 carinae present, 6 of them distinct; interno- 
dorsal carina developed as a granular ridge; externodorsal carina developed as a ridge 
of indistinct granules; interno ventral carina developed as a granular ridge; externo- 
ventral carina developed as a ridge of weak granules; externomedian carina developed 
as a weakly granular ridge. Dorsal surface shagreened, sparsely and weakly granular; 
internal and external surfaces shagreened; ventral surface shagreened, sparsely and 
weakly granular, distal end without granules. Internal protuberance pronounced, bifid 
(internodorsal and internoventral tubercles separated), internodorsal carina with 1 
spiniform granule. A total of 19 trichobothria present on patella, orthobothriotaxy; dj 
located basally, external to internodorsal carina; d 2 located midway on patella; d 3 
absent; i in distal half of patella, located dorsally on internal surface. External (e) 
trichobothrial groups discernible and orthobothriotaxic: eb group composed of 5 tri- 
chobothria, esb group of 2 trichobothria, em group of 2 trichobothria, et group of 3 tri- 
chobothria; est very close to et group. Ventral (v) group orthobothriotaxic, 3 tri- 
chobothria. Pedipalp chela manus (Fig. 5A-C). Colour predominantly orange to red- 
brown; carinae darker red. Chela stout (length less than 2.5 times its width) with 5 dis- 
tinct carinae; internodorsal carina continuous, with strong spiniform granules basally 
and weak granules distally; subdigital carina vestigial, visible basally as a smooth ridge 
with weak granules; externodorsal carina distinct, visible as a ridge of indistinct fused 
granules, almost smooth; digital carina a smooth ridge with few weak granules basally; 
dorsal secondary carina (dorsomedian) a smooth ridge with few strong granules basally 
and few weak granules distally; internoventral carina continuous, visible as a weak 
ridge with few weak spiniform granules distally; externoventral carina continuous, 
forming a granular ridge, becoming almost smooth distally, running parallel to longi- 
tudinal axis of chela, its distal edge disconnected from external movable finger condyle 
and directed between external and internal condyles of movable finger; ventromedian 
carina vestigial, reduced to few coarse granules basally; internal (internomedian) 
carina vestigial, reduced to few weak spiniform granules distally; external (externo- 
median) carina indistinct, visible as raised rows of small weak granules. Dorsal surface 
forming an irregular and subreticulate network of weakly granular ridges surrounding 
shagreened patches without granules; internal surface shagreened, sparsely granular, 
with a denser patch of spiniform granules dorsodistally; external surface shagreened, 
sparsely and weakly granular; ventral surface with few weak granules surrounding 
shagreened patches, distal end smooth and pitted. A total of 15 trichobothria present on 
chela; Db trichobothria located externobasally on dorsal surface; Eb group (3 tri- 



HEMISCORPIIDAE FROM IRAN 



881 




Fig. 4 
Hemiscorpius acanthocercus sp. n., metasoma, lateral aspect: A, female paratype (NHMW 
4717); B, male paratype (MHNG, H. Löffler coll.). Scale line, 2 mm. 



882 



L. MONOD & W. R. LOURENÇO 



— h- _q 




Fig. 5 
Hemiscorpius acantiiocercus sp. n. Male paratype (MHNG, H. Löffler coll.): A, pedipalp with 
trichobothrial pattern, dorsal aspect; B, chela, external aspect; C, idem, ventral aspect; E, femur 
and patella, external aspect; F, idem, ventral aspect. Female paratype {NHMW 4719): D, pedi- 
palp, dorsal aspect. Scale line, 2 mm. 

chobothria) orthobofhriotaxic, located basally on external surface; Esb very close to Eb 
group; Em absent; Est located distally, very close to Et group: Et group composed of 5 
trichobothria, Etj located ventrally; V group comprising 4 trichobothria, V3 and V4 lo- 
cated in basal half of manus, VI and V2 located very distally. Fingers of pedipalpal 
chela (Fig. 5A-C). Basally reddish tan, becoming gradually lighter distally, tips of fin- 



HEMISCORPIIDAE FROM IRAN 



883 




Fig. 6 
Hemiscorpius acanthocercus sp. n., pectines and genital opercula, ventral aspect: A, male holo- 
type; B, female paratype (NHMW 21143). Scale line, 1 mm. 



884 L. MONOD & W. R. LOURENÇO 

gers lighter orange. Dorsal and ventral surfaces predominantly smooth and minutely 
pitted, slightly shagreened basally. Fingers slightly shorter than chelal manus. Fixed 
finger with weak basal concavity, movable finger with corresponding lobe weakly de- 
veloped. Tips of fingers with pronounced terminal hook. Fingers with distal diastema; 
edges of fingers composed of 2 rows of denticles becoming fused basally, interrupted 
at regular intervals by stronger granules, each of these coupled with an accessory gran- 
ule; rows fused into a single row above the concavity on the fixed finger and on notch 
of movable finger, single basal row running to base of fingers, in some males absent on 
movable finger. A total of 11 trichobothria present on fingers; Dt located very basally 
on dorsal surface; db located dorsally on internal surface, in basal half of fixed finger; 
dsb, dst and di on dorsal surface, in distal half of finger; eb located dorsally on exter- 
nal surface, in basal 1/3 of fixed finger, opposite db; esb, est and et in distal half of fixed 
finger, opposite dsb, dst and dt respectively; it and ib located in median 1/3 of fixed 
finger. 

Coxosternal sclerites. Dark yellow to orange; smooth and minutely pitted. 
Anterior margin of coxapohysis I with few weak granules, expanded but not sub-trian- 
gular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, penta- 
gonal, slightly shagreened; median furrow deep, more pronounced in posterior half; 
posterior pit absent. 

Legs. Pale yellow. Predominantly shagreened; ventral surface smooth; dorsal 
surfaces of trochanter and femur finely and weakly granular. Tarsus with a ventro- 
median row of microspinules and with 2 rows of rigid "spinoid" ventral macrosetae, 
tarsi I-II: prolateral row with 4/5 macrosetae, retrolateral row with 5/6 macrosetae; tarsi 
III-IV: prolateral row with 5/6 macrosetae, retrolateral row with 6 macrosetae; setae 
acuminate; tarsal claws of equal length. 

Pectines and genital operculum (Fig. 6A). Colour dark yellow; genital oper- 
culum composed of 2 subtriangular plates; genital papillae short, not protruding from 
beneath operculum; pectines with 14/15 teeth. 

Hemispermatophore (Fig. 7) lamelliform, with complex capsule; distal 
lamellum slender, basally curved, with a strong double hook pointing antero-distad and 
located very basally, flagellum more than 1 .5 times longer than basal part; distal crest 
absent. Double lamellar hook (Dh) located very slightly above distal transverse ridge 
(Tr); distal transverse ridge costate, distally strongly curved towards anterior margin, 
continuous from posterior to anterior margins, almost merging with lamellar hook, 
reaching anterior margin of distal lamella slightly below its hook. Capsule lamella (La) 
broad, with a deep longitudinal furrow, forming an acute angle with longitudinal axis 
of capsule, bearing a strong hook on its external surface and an accessory lobe (Lac) 
on its posterior margin; ventral margin straight. Distal lobe (Ld) forming a strong hook 
pointing anteriad; basal margin straight. Basal lobe (Lb) strongly reduced. Posterior 
lamella (Lp) costate. 

FEMALE (Figs 2C-D, 3B, 4A, 5D, 6B). Measurements of paratype (NHMW 
4719). Carapace, length 5.9. posterior width 5.0; distance between anterior lateral eyes 
3.1, between posterior lateral eyes 3.6, between median eyes 0.2; diameter of median 
eyes 0.2. Pedipalp. femur length 5.0. width 1.9; patella length 5.2, width 1.8, chela 
length 10.1; manus length 5.2, width 4.0, depth 2.7; movable finger length 5.2; fixed 
finger length 3.9. Metasoma, segment I length 3.3, width 1.9; segment V length 5.1, 



HEMISCORPIIDAE FROM IRAN 



885 






Fig. 7 
Hemiscorpius acanthocercus sp. n., male holotype, hemispermatophore: A, whole hemisper- 
matophore, arrows indicating its post-extrusion orientation, Ant (anterior), Bas (basal), Dist 
(distal), Post (posterior), Tr (transverse ridge); B-C, detail of the capsular region, internal aspect, 
Dh (double hook), La (lamella), Lac (accessory lobe), Lb (basal lobe), Ld (distal lobe), Lp (pos- 
terior lobe). Scale lines, 0.25 mm. 



886 L. MONOD & W. R. LOURENÇO 

width 1 .4, depth 1 .7; vesicle length 3.7, width 2.0, depth 2.1 , aculeus length 1 .2. Total 
length 53.0. 

Same characters as in males, except as follows. Carapace (Fig. 3B) shagreened 
and finely granular, each anterior lobe with a smooth, shiny and minutely pitted median 
area between median and lateral ocular tubercles; median longitudinal furrow shallow, 
continuous from the anterior suture furcation, ending after the median ocular tubercle 
and not running posteriorly into a deep triangular depression. Mesosoma. Median carina 
absent from tergites I-III. Surface of tergites smooth, shiny; median area finely granu- 
lar; median carina on tergites IV-IV smooth and pitted posteriorly. Sternites III- VI 
smooth, without granulation or carinae. Sternite VII smooth, with a pair of median cari- 
nae developed as very weak ridges, almost indiscernible. Tergites and sternites VII as 
long as wide. Metasoma (Fig. 4A). Short and stout, shagreened, without granules. 
Segments I-II with ventrolateral carinae forming almost smooth ridges. Segment I 
without ventral carina; segments II with ventral carina developed as a ridge, smooth in 
anterior half, with weak granules in posterior half; segment III with ventral carina form- 
ing a ridge of weak granules; segment IV with ventral carina developed as a ridge of 
weak spiniform granules. Segment V: dorsal carinae with spiniform granules; lateral 
carina vestigial, barely visible in anterior 2/3, composed of few very weak scattered 
granules; ventromedian carina with strong spiniform granules. Telson (Fig. 4A). Vesicle 
globular, very bulky; lateral surfaces smooth, with few extremely weak granules 
(almost indistinct); dorsal surface smooth, median shallow depression indistinct, only 
visible anteriorly. Aculeus without tubercles at its base, narrowing slightly less marked 
than in male. Pedipalp patella (Fig. 5D). Internal protuberance with internodorsal carina 
carrying 1-2 spiniform granules, less developed than in male. Pedipalpal chela (Fig. 5D) 
slightly more bulky than in male, chela fingers equal in length to chela manus or slightly 
shorter. Carinae of chelal manus generally less granular and less distinct than in males. 
Interno-dorsal carina vestigial, basally visible as few strong spiniform granules; sub- 
digital carina vestigial; externodorsal carina distinct, visible as a smooth ridge; digital 
carina developed as a weak ridge with few weak granules basally; dorsal secondary 
carina (dorsomedian) forming a weak, smooth ridge with 1-2 strong granules basally; 
internoventral carina continuous, visible as a weak ridge with few weak granules dis- 
tally; externoventral carina continuous, a ridge of indistinct granules, becoming almost 
smooth distally. Ventral surface smooth and pitted with shagreened patches. Pectines 
and genital operculum (Fig. 6B). Genital operculum composed of a single posteriorly 
truncated subtriangular plate with a median longitudinal furrow and a relatively deep 
posterior notch; pectines with 10/10 teeth. 

Intraspecific variation. Females have pectines with 8 to 11 teeth, usually 9/9 or 
10/10, males have pectines with 13 to 16 teeth, usually 14/14 or 15/15. Juveniles (males 
and females) have slender pedipalps, which are morphologically similar to the pedi- 
palps of mature males. Juveniles and subadult males do not possess an extremely elon- 
gated metasoma, they apparently acquire this feature only in the last developmental 
stage. 

Hemiscorpius enischnochela sp. n. Figs 1E-F, 8-12, 26A-B, 27 C-D, 36 

Material examined. Holotype: 3 , IRAN (Hormozgan), ca. 115 km E Bandar Abbas, 
VII. 1974, G. Pretzman, NHMW (52). Paratypes: 2 $, IRAN (Khuzestan), S of Masdjed-e- 



HEMISCORPIIDAE FROM IRAN 



887 




Fig. 8 
Hemiscorpius enischnochela sp. n. Male holotype: dorsal aspect. Scale line, 5 mm. 



Soleyman, 1.1970, R. Farzanpay ?, MHNG (SF 0057-1/2). 1 ? juv., same data as for holotype, 
NHMW(53). 1 5 subadult, IRAN (Hormozgan), 38 km N Bandar Abbas, 28.III.1972, K. Bilek. 
NHMW (54). 1 $ juv., IRAN (Hormozgan), 38 km N Bandar Abbas, 28.ffl.1972, K. Bilek, 
NHMW (58). 1 9 juv., IRAN (Hormozgan), 22 km N Bandar Abbas, VII. 1974, G. Pretzman ?, 
NHMW (60). 1 9 juv., IRAN (Hormozgan), 65 km N Bandar Abbas, 4.IV.1972, K. Bilek, 



L. MONOD & W. R. LOURENÇO 



NHMW (64). 1 S juv., IRAN (Hormozgan), 65 km N Bandar Abbas, 4.IV.1972, K. Bilek, 
NHMW (67). 1 9 , IRAN (Hormozgan), Kol River bridge on a road near Lar, 32 km W Bandar 
Abbas, 1972, K. Bilek, NHMW (76). 2 S subadults, IRAN (Hormozgan), Bandar Abbas, 
VI. 1972, G. Pretzman, NHMW (80-81). 1 9 juv., IRAN (Hormozgan), ca. 41 km N Bandar 
Abbas, 7.IV.1972, G. Pretzman & A. Konetschnig, NHMW 3395. 

Distribution. Known from surroundings of Bandar Abbas, Hormozgan 
Province, eastern Iran; S of Masdjed-e-Soleyman, Khuzestan Province (doubtful 
record, see below) (Fig. 36). 

Remark. The species occurs around Bandar-Abbas in Eastern Iran. Among the 
material studied by Vachon, there was a female found far from this area, near Masdjed- 
e-Soleyman. There was no indication about the collector, who could possibly be R. 
Farzanpay. Since much information about Vachon 's material was lost and mixed up 
after his death, it could be a case of mislabelling. 

Etymology. The name enischno chela is a construct from the Greek words 
enischnos [slender, thin] and chele [claw]. The name is an invariable noun in appo- 
sition and refers to the pedipalp chela, which is slender and with long fingers. 

Diagnosis. (1) Carapace longer than wide, shagreened and sparsely granular, 
with coarse granules on anterior margin between lateral ocular tubercles; lateral mar- 
gins finely granular, especially below lateral ocular tubercles; superciliary carinae 
weak and finely granular; (2) pedipalps elongated, chela fingers slightly longer than 
chela manus; (3) internal protuberance of pedipalp patella with internodorsal carina 
bearing 3-4 strong spiniform granules; (4) pedipalp patella neobothriotaxic, external 
side with 14 trichobotria (1 est and 3 esb), ventral side with 10-12 trichobothria; (5) 
males with elongated and slender metasoma, with dorsal carinae of segments I-V and 
ventral and ventrolateral carinae of segments IV- V bearing small spiniform granules; 
(7) telson strongly elongated, bearing a pair of blunt tuberculiform processes at base of 
aculeus, surface irregular, rugose, without distinct granules; (8) metasoma of females 
with dorsal carinae of segments I-IV and ventral and ventrolateral carinae of segments 
IV- V bearing spiniform granules. 

H. enischnochela sp. n. is closely related to H. gaillardi, both of them have 
elongated neobothriotaxic pedipalps. They can be readily distinguished by (1) their tri- 
chotriotaxy (H. gaillardi has an additional est trichobothry on the external face of the 
patella) and by (2) their pedipalp carination (H. enischnochela sp. n. has patellar 
processes with intero-dorsal carinae bearing 3 strong spiniform granules, while H. gail- 
lardi has patellar processes with carinae bearing 6 spiniform granules, which are 
slightly smaller than those in H. enischnochela sp. n.). 

Description. MALE (Figs 1E-F, 8, 10A-C, E-F, IIA, 12, 26 A, 27D) [description 
based on the holotype (NHMW 52)]. Measurements of holotype (in mm). Carapace, 
length 7.6; posterior width 7.0; distance between anterior lateral eyes 4.0, between pos- 
terior lateral eyes 4.9, between median eyes 0.2; diameter of median eyes 0.2. Pedipalp, 
femur length 8.8. width 2.7; patella length 8.6, width 2.6, chela length 15,2; manus 
length 7.4, width 4.6, depth 3.2; movable finger length 8.2; fixed finger length 7.2. 
Metasoma. segment I length 9.6, width 2.5; segment V length 12.3, width 1.6, depth 
2.0; vesicle length 7.4. width 2.3, depth 1.9. aculeus length 1.0. Total length 93.0. 

Carapace (Fig. 26A). Colouration mostly uniformly dark yellow to orange; 
median and lateral ocular tubercle black. Carapace dorsoventraly flattened, longer than 



HEMISCORPIIDAE FROM IRAN 



889 





MG. 9 

Hemiscorpius enischnochela sp. n. Female paratype (NHMW 76, Kol River): A, dorsal aspect; 
B, ventral aspect. Scale line, 5 mm. 



890 



L. MONOD & W. R. LOURENÇO 




Fig. 10 
Hemiscorpius enischnochela sp. n. Male holotype: A, pedipalp with trichobothrial pattern, dor- 
sal aspect; B, chela, external aspect; C, idem, ventral aspect; E, femur and patella, external as- 
pect; F, idem, ventral aspect. Female paratype {MHNG, Masdjed-e-Soleyman): D, pedipalp, dor- 
sal aspect. Scale line, 2 mm. 



wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle 
weakly developed, low, distinctly situated anteriorly, with weak, finely granular super- 
ciliary carinae; frontal concavity or notch well-developed; anterior lobes truncate; lat- 
eral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. 
Carapace shagreened, with sparse granules; stronger granules on the anterior margin 
between lateral ocular tubercles; edges granular, with stronger spiniform granules be- 
low lateral ocular tubercles; anteromedian furrow narrow, suturiform, anteriorly bifur- 
cated; median longitudinal furrow shallow, continuous from the anterior suture furca- 
tion, running through ocular tubercle posteriorly into a deep triangular depression; pos- 
terolateral furrow shallow; mesolateral furrow weakly developed, almost indiscernible. 



HEMISCORPIIDAE FROM IRAN 



891 




Fig. 11 
Hemiscorpius enischnochela sp. n., pectines and genital opercula, ventral aspect: A, male holo- 
type; B, female paratype (MHNG, Masdjed-e-Soleyman). Scale line, 1 mm. 



892 L. MONOD & W. R. LOURENÇO 

Mesosoma. Tergite colouration dark orange to brown, becoming lighter poste- 
riorly (dark yellow), tergite VII with a larger yellow posterior zone than on other ter- 
gites. Tergites I-III with a shallow median depression divided anteriorly by a weak 
ridge, tergites IV- VII with median carina reduced to a weak non-granular ridge sur- 
rounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae ab- 
sent on tergites I- VI, but present on posterior 2/3 of tergite VII, visible as strong ridges 
with spiniform granules. Surface of tergites shagreened, with scattered granules; pre- 
tergites smooth, shiny and minutely pitted. Sternite colouration pale yellow. Sternites 
III- VI smooth, shiny and minutely pitted, without granulation or carinae; sternite III 
with a pair of large, very finely and densely granular depressions, covered by the 
pectines; sternites IV- VI with a pair of shallow median furrows. Sternite VII with sim- 
ilar surface texture as on preceding sternites; a pair of lateral, slightly granular carinae 
present; median carinae absent. Spiracles of book lungs crescent-shaped. Tergites and 
sternites VII longer than wide. 

Metasoma (Fig. 27D). Very elongated and slender, shagreened, with scattered 
granules. Colour dark yellow to orange. Segments I-IV with longitudinal dorsomedian 
furrow and with dorsal carinae composed of spiniform granules. Segments I-II with lat- 
eral carinae developed as granular ridges; segments III-IV with lateral carinae as ridges 
of indistinct fused granules, almost smooth. Segments I-II with ventrolateral carinae as 
granular ridges; segments III-IV with ventrolateral carinae as ridges of reduced spini- 
form granules. Segment I: ventral carina almost absent, reduced to a weak ridge; seg- 
ments II-III with ventral carina forming a ridge of fused granules, almost smooth; seg- 
ment IV with ventral carina forming a ridge of fused granules, almost smooth in the 
anterior half, and with more defined spiniform granules posteriorly. Segment V: longi- 
tudinal dorsal furrow present in anterior half, dorsal carinae with reduced spiniform 
granules; lateral carina indistinct, only visible in anterior half as a ridge of fused gran- 
ules; ventrolateral carinae with reduced spiniform granules; ventromedian carina with 
spiniform granules. 

Telson (Fig. 1E-F, 27D). Vesicle orange; aculeus darker, tan, due to stronger 
sclerotisation; vesicle elliptical or ovate, strongly elongated, with a blunt tuberculiform 
projection on each side at base of aculeus; ventrolateral furrows absent; ventromedian 
ridge absent; lateral surfaces rugose, irregular, with scattered weak granules; dorsal 
surface with numerous small spiniform granules and a median shallow depression. 
Macrosetae very sparse basally, becoming more numerous near base of aculeus. 
Aculeus short and stout, strongly curved, becoming markedly narrower approximately 
midway. 

Chelicerae. Colour pale yellow; teeth of fingers darker orange. Teeth arrange- 
ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth 
bifid; movable finger with one subdistal tooth and one basal tooth in external series; 
distal external tooth smaller than distal internal tooth; cheliceral teeth without second- 
ary serrations. 

Pedipalp slender and elongated. Pedipalp coxa and femur (Fig. 10A, E-F). 
Internoventral margin of coxa with strong spiniform granules. Dorsal surface of femur 
predominantly dark yellow-orange, internodorsal, internoventral and externo ventral 
carinae darker orange. Femur slender, elongate (equal to or longer than 3 times its 



HEMISCORPIIDAE FROM IRAN 



893 




Fig. 12 
Hemiscorpius enischnochela sp. n., male holotype: left tarsus IV, ventral aspect. Scale line, 
0.5 mm. 



width), pentac annate, with 4 distinct carinae; internodorsal carina developed as a 
strongly and densely granular ridge, granules spiniform; externodorsal carina a strong 
granular ridge; internoventral carina developed as a granular ridge with spiniform 
granules; externoventral carina developed as a ridge with strong spiniform granules; 
ventromedian carina reduced to few granules and confined to base of femur. Dorsal 
surface shagreened, finely and densely granular, distal end smooth, without granules; 
internal surface shagreened, with few strong scattered spiniform granules; external sur- 
face shagreened; ventral surface shagreened and sparsely granular, distal end without 



894 L. MONOD & W. R. LOURENÇO 

granules. A total of 3 trichobothria present on femur; d located externobasally on dor- 
sal surface; i located dorsobasally on internal surface; e located dorsobasally on exter- 
nal surface. Pedipalp patella (Figs 10 A, E-F) slender (length less than 2.5 times its 
width). Colour predominantly dark yellow to orange; carinae slightly darker. 7 carinae 
present, 6 of them distinct; internodorsal carina developed as a ridge of coarse gran- 
ules; externodorsal carina forming a ridge of indistinct coarse granules; interno ventral 
carina developed as a strongly granular ridge (spiniform granules); externoventral ca- 
rina as a ridge of weak granules; externomedian carina as a granular ridge. Dorsal and 
internal surfaces shagreened, sparsely and weakly granular; external surface sha- 
greened, with a few scattered granules; ventral surface shagreened, sparsely and weak- 
ly granular, distal end without granules. Internal protuberance pronounced, bifid (in- 
ternodorsal and internoventral tubercles separated), internodorsal carina with 3-4 
strong spiniform granules. A total of 28 trichobothria present on patella, neobothrio- 
taxy major; d } located basally, external to internodorsal carina; d 2 situated in distal half 
of patella; d 3 absent; i in distal half of patella, located dorsally on internal surface. 
External (e) trichobothrial groups discernible and neobothriotaxic major: eb group 
composed of 5 trichobothria, esb group of 3 trichobothria, em group of 2 trichobothria, 
et group of 3 trichobothria; est very close to et group. Ventral (v) group neobothrio- 
taxic, 11 trichobothria. Pedipalp chela manus (Figs 10A-C). Colour predominantly 
orange to red-brown; carinae darker red. Chela slender (length more than 3 times its 
width), with 5 distinct carinae; internodorsal carina continuous, with spiniform 
granules; subdigital carina vestigial, visible basally as a smooth ridge with weak 
granules; externodorsal carina distinct, visible as a ridge of indistinct fused granules, 
almost smooth; digital carina a smooth ridge with few weak granules basally; dorsal 
secondary carina (dorsomedian) a smooth ridge with few strong granules basally and 
few weak granules distally; internoventral carina continuous, visible as a weak ridge 
with few spiniform granules; externoventral carina continuous, forming a granular 
ridge, becoming almost smooth distally, running parallel to longitudinal axis of chela, 
its distal edge disconnected from external movable finger condyle and directed 
between external and internal condyles of movable finger; ventromedian carina ves- 
tigial, reduced to few coarse granules basally; internal (internomedian) carina visible 
as a weak ridge with few spiniform granules; external (externomedian) carina hardly 
discernible, visible as raised rows of small indistinct granules. Dorsal surface sha- 
greened, sparsely and weakly granular; internal surface shagreened, sparsely granular, 
with a denser patch of spiniform granules dorsodistally; external surface shagreened, 
sparsely and weakly granular; ventral surface shagreened, sparsely and weakly granu- 
lar, distal end without granules. A total of 15 trichobothria present on chelal manus; Db 
trichobothria situated externobasally on dorsal surface; Eb group (3 trichobothria) or- 
thobothriotaxic, located basally on external surface; Esb very close to Eb group; Em 
absent; Est located distally, very close to Et group; Et group composed of 5 tri- 
chobothria, Et i located ventrally; V group comprising 4 trichobothria, V3 and V4 situ- 
ated in basal third of manus, VI and V2 located in distal quarter. Pedipalp chela fingers 
(Figs 10A-C). Basally reddish tan, becoming gradually lighter distally, tips of fingers 
yellow. Dorsal and ventral surfaces predominantly smooth, slightly shagreened basally. 
Fingers longer than chela manus. Fixed finger with a weak basal concavity, movable 



HEMISCORPIIDAE FROM IRAN 895 

finger with corresponding lobe weakly developed. Tips of fingers with pronounced 
distal hook. Fingers with distal diastema; edges of fingers composed of 2 rows of den- 
ticles becoming fused basally, interrupted at regular intervals by stronger granules, 
each of them coupled with an accessory granule; rows fused into a single row above 
concavity on fixed finger and on notch of movable finger, single basal row running 
towards base of fingers. A total of 11 trichobothria present on fingers; Dt located on 
dorsal surface, in basal third of fixed finger; db located dorsally on internal surface, ap- 
proximately midway on fixed finger; dsb, dst and dt on dorsal surface, in distal half of 
finger; eb located dorsally on external surface, approximately midway on fixed finger, 
opposite db; esb, est and et in distal half of fixed finger; esb and et opposite dsb and dt, 
respectively; it and ib located in distal half of fixed finger. 

Coxosternal sclerites. Dark yellow to orange, smooth and minutely pitted. 
Anterior margin of coxapohysis I with few weak granules, expanded but not sub-trian- 
gular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, pentago- 
nal, shagreened; median furrow deep, more pronounced in posterior half; posterior pit 
absent. 

Legs. Pale yellow. Predominantly shagreened; ventral surface smooth; dorsal 
surfaces of trochanter and femur finely granular. Tarsus (Fig. 12) with a ventromedian 
row of microspinules and with 2 rows of rigid "spinoid" ventral macrosetae; tarsi I-II: 
prolateral row with 5 or 6 macrosetae, retrolateral row with 7 macrosetae; tarsi III-IV: 
prolateral row with 7 macrosetae, retrolateral row with 7/8 macrosetae; setae acumi- 
nate; tarsal claws of equal length. 

Pectines and genital operculum (Fig. IIA). Colour pale yellow; genital opercu- 
lum composed of 2 ovoid plates; genital papillae short, not protruding from beneath 
operculum; pectines with 17/17 teeth. 

Hemispermatophore. Unknown. The specimen was already dissected and the 
hemispermatophores were probably lost. 

FEMALE (Figs 9, 10D, IIB, 26B, 27C). Measurements of paratype (MNHN 
Masdjed-e Soleyman). Carapace, length 7.4, posterior width 7.0; distance between an- 
terior lateral eyes 4.0, between posterior lateral eyes 4.7, between median eyes 0.2; di- 
ameter of median eyes 0.2. Pedipalp, femur length 7.5, width 2.4; patella length 7.2, 
width 2.6, chela length 13.6; manus length 6.8, width 4.6, depth 3.3; movable finger 
length 6.9; fixed finger length 6.2. Metasoma, segment I length 4.8, width 2.2; segment 
V length 6.9, width 1.6, depth 2.0; vesicle length 5.2, width 2.8, depth 2.7, aculeus 
length 1 .3. Total length 64.0. 

Same characters as in males, except as follows. Carapace (Fig. 26B) shagreened 
and sparsely granular, each anterior lobe with a reduced central smooth area between 
median and lateral ocular tubercles. Mesosoma. Surface of tergites shagreened, with 
few smooth and pitted patches and scattered granules. Tergites and sternites VII almost 
as long as wide. Metasoma (Fig. 27C). Short and stout. Segments I-II with ventral 
carina developed as a smooth ridge; segments III with ventral carina as a ridge of weak 
spiniform granules; segment IV with ventral carina as a ridge of spiniform granules. 
Segment V: lateral carina indistinct; ventromedian carina with spiniform granules. 
Telson (Fig. 27C). Vesicle elliptical or ovate, short and bulky; lateral surfaces smooth 
and irregular; dorsal surface smooth and irregular, with a shallow median depression 



896 L - MONOD & W. R. LOURENÇO 

posteriorly. Aculeus without tubercles at its base, narrowing slightly less marked than 
in male. Pedipalp coxa and femur (Fig. 10D). Pedipalp slightly less slender and elon- 
gated than in males. Femur: externo ventral carina developed as a ridge with spiniform 
granules, these weaker, more scattered and less numerous than in male. Pedipalp patel- 
la (Fig. 10D). Less elongated than in males. Externodorsal carina developed as a ridge 
of weak granules, almost smooth. Internal and external surface shagreened, not granu- 
lar. Pedipalp chela manus (Fig. 10D). Chela more bulky than in males. Pectines and 
genital operculum (Fig. IIB). Genital operculum composed of a hexagonal ovoid plate 
divided by a moderately deep longitudinal furrow; posterior notch weakly pronounced; 
pectines with 8-10 teeth. 

Intraspecific variation. The specimens examined possess 10 to 12 trichobothria 
on the ventral side of the patella. Females have pectines with 8 to 10 teeth, usually 8- 
8, males have pectines with 14 to 17 teeth. The same remark about pedipalps and meta- 
soma of males can be given as for H. acanthocercus . 

Hemiscorpius gaillardi (Vachon, 1974) comb. n. Figs 13-15, 26C 

Habibiella gaillardi Vachon, 1974: 952, fig. 105. 

Material examined. Holotype: 1 9 , IRAN, East, no locatity specified, MNHN-RS 4328. 

Distribution. Known from eastern Iran, no locatity specified. 

Diagnosis. (1) Carapace almost as wide as long, shagreened and sparsely 
granular, with few smooth patches and with coarse granules on anterior margin be- 
tween lateral ocular tubercles; lateral margins with small spiniform granules below lat- 
eral ocular tubercles; superciliary carinae weak, finely granular; (2) pedipalps very 
elongated, chela fingers longer than chela manus; (3) internal protuberance of patella 
with internodorsal carina bearing 6 strong spiniform granules; (4) pedipalp patella 
neothobothriotaxic, external side with 15 trichobotria (2 est and 3 esb), ventral side 
with 10-12 trichobothria; (5) metasoma of females with dorsal carinae of segments 
I-IV and ventral and ventrolateral carinae of segments V bearing weak small spiniform 
granules. 

The male of H. gaillardi remains unknown. For more details see also diagnosis 
of H. enischnochela sp. n., to which H. gaillardi is closely related. 

Description. FEMALE. Measurements of holotype (MNHN-RS 4328). 
Carapace, length 6.6, posterior width 6.7; distance between anterior lateral eyes 3.6, 
between posterior lateral eyes 4.4, between median eyes 0.2; diameter of median eyes 
0.3. Pedipalp, femur length 7.7, width 2.3; patella length 7.5, width 2.4, chela length 
14.2; manus length 6.2, width 3.8, depth 2.7; movable finger length 7.7; fixed finger 
length 6.7. Metasoma, segment I length 4.5, width 2.2; segment V length 6.5, width 
1 .7, depth 1 .9: vesicle length 4.2, width 2.3, depth 2.3, aculeus length 1 .5. Total length 
58.0. 

Carapace (Fig. 26C). Colouration mostly uniformly pale yellow to orange; 
median and lateral ocular tubercle black. Carapace dorsoventraly flattened, almost as 
wide as long, with sides nearly parallel, slightly convergent; median ocular tubercle 
weakly developed, low, distinctly situated anteriorly, with superciliary carinae weak 
but finely granular. Frontal concavity or notch well-developed; anterior lobes truncate; 



HEMISCORPIIDAE FROM IRAN 



897 





Fig. 13 
Hemiscorpius gaillardi, female holotype: A, dorsal aspect; B, ventral aspect. Scale line, 5 mm. 



L. MONOD & W. R. LOURENÇO 




B 





Fig. 14 
Hemiscorpius gaillardi, female holotype, pedipalp with trichobothrial pattern: A, chela, dorsal 
aspect; B, idem, external aspect; C. idem, ventral aspect; D, femur and patella, dorsal aspect; E, 
idem, external aspect; F. idem, ventral aspect. Scale line, 2 mm. 



HEMISCORPIIDAE FROM IRAN 



899 




Fig. 15 
Hemiscorpius gaillardi, female holotype: A, metasoma, lateral aspect (scale line, 2 mm); B, 
pectines and genital opercula, ventral aspect (scale line, 1 mm). 



900 L. MONOD & W. R. LOURENÇO 

lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. 
Carapace shagreened and sparsely granular, with few smooth patches and with coarse 
granules on anterior margin between lateral ocular tubercles, each lobe with a smooth 
central area between median and lateral ocular tubercles; edges granular below lateral 
ocular tubercles; anteromedian furrow indistinct, very shallow, suturiform, anteriorly 
bifurcated; median longitudinal furrow indistinct, almost absent, very shallow, conti- 
nuous from the anterior suture furcation, running through ocular tubercle posteriorly 
into a deep triangular depression; posterolateral furrow shallow; mesolateral furrow 
weakly developed, almost indiscernible. 

Mesosoma. Tergite colouration dark yellow, with lighter (pale yellow) posteri- 
or margins. Tergites 1-111 with a shallow median depression divided anteriorly by a 
weak ridge, tergites IV- VII with median carina reduced to a weak, non-granular ridge 
surrounded by a pair of shallow, submedian depressions. Lateral and sublateral carinae 
absent on tergites I- VI, but present in posterior 2/3 of tergite VII, developed as weak 
ridges with weak, spiniform granules. Surface of tergites shagreened, with smooth 
patches; pre-tergites smooth, shiny and minutely pitted. Sternite colouration pale yel- 
low. Sternites III- VI smooth, shiny and minutely pitted, without granulation or carinae; 
sternite III with a pair of large, very finely and densely granular depressions, situated 
under the pectines; sternites IV- VI with a pair of shallow median furrows. Sternite VII 
shagreened, very finely granular and with very few smooth patches; pair of lateral cari- 
nae, weak, smooth; median carinae absent. Spiracles of book lungs crescent-shaped. 
Tergites and sternites VII almost as long as wide. 

Metasoma (Fig. 15A). Slender, predominantly shagreened. Colour dark yellow. 
Segments I-IV with longitudinal dorsomedian furrow, their dorsal carinae with spini- 
form granules. Segments I-III with lateral carinae developed as ridges with weak spini- 
form granules, almost smooth; segments IV with lateral carinae as almost smooth 
ridges. Ventrolateral carinae developed as almost smooth ridges on segments I-II; as 
smooth ridge with weak spiniform granules in posterior half of segment III; as ridge 
with weak spiniform granules on segment IV Segment I: ventral carina almost indis- 
cernible, reduced to a very weak ridge; segments II-IV with ventral carina a smooth 
ridge with weak spiniform granules on segment IV Segment V: longitudinal dorsal 
furrow present in anterior half, dorsal carinae weakly marked, with small, very weak 
granules; lateral carina absent; ventrolateral carinae with weak spiniform granules; 
ventre median carina with small spiniform granules. 

Telson (Fig. 15A). Vesicle yellow to orange; aculeus darker, tan, due to stronger 
sclerotisation; vesicle elliptical or ovate, very bulky; ventrolateral furrows absent; 
ventromedian ridge absent; lateral surfaces smooth; dorsal surface smooth, with a 
median shallow depression anteriorly. Macrosetae very sparse basally, becoming more 
numerous near base of aculeus. Aculeus short, strongly curved, becoming markedly 
narrower approximately halfway. 

Chelicerae. Colour pale yellow; teeth of fingers darker, orange. Teeth arrange- 
ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth 
bifid; movable finger with one subdistal tooth and one basal tooth in external series; 
distal external tooth smaller than distal internal tooth; cheliceral teeth without 
secondary serrations. 



HEMISCORPIIDAE FROM IRAN 901 

Pedipalp slender and elongated. Pedipalp coxa and femur (Fig. 14D-F). Inter- 
noventral margin of coxa with few strong spiniform granules. Dorsal surface of femur 
predominantly yellow, internodorsal, interno ventral and externo ventral carinae darker, 
orange. Femur slender, elongate (longer than 3 times its width), pentacarinate, with 4 
distinct carinae; internodorsal carina developed as a strongly and densely granular 
ridge, granules spiniform; externodorsal carina as a granular ridge; internoventral 
carina as a ridge with spiniform granules; externo ventral carina as a ridge with strong 
spiniform granules; ventromedian carina reduced to few scattered granules and con- 
fined to base of femur. Dorsal surface shagreened, finely and densely granular, distal 
end smooth, without granules; internal surface shagreened, with few strong scattered 
spiniform granules; external surface shagreened; ventral surface shagreened and 
sparsely granular, distal end without granules. A total of 3 trichobothria present on 
femur; d located externobasally on dorsal surface; i located dorsobasally on internal 
surface; e located dorsobasally on external surface. Pedipalp patella (Fig. 14D-F) 
slender (length more than 2.5 times its width). Colour predominantly yellow; carinae 
slightly darker. 7 carinae present, 6 of them distinct; internodorsal carina developed as 
a ridge of coarse granules; externodorsal carina as a ridge of indistinct coarse granules 
(almost smooth); internoventral carina as a strongly granular ridge (spiniform 
granules); externoventral carina as a ridge of weak granules; externomedian carina as 
a granular ridge. Dorsal surface shagreened, sparsely and weakly granular; internal, 
external and ventral surfaces shagreened, without granules. Internal protuberance pro- 
nounced, bifid (internodorsal and internoventral tubercles separated), internodorsal 
carina with 6 strong spiniform granules. A total of 29-31 trichobothria present on 
patella, neobothriotaxy major; dj located basally, external to internodorsal carina; d 2 
located in distal third of patella; d 3 absent; i in distal third of patella, located dorsally 
on internal surface. External (e) trichobothrial groups discernible and neobothriotaxic: 
eb group composed of 5 trichobothria, esb group of 3 trichobothria, em group of 2 tri- 
chobothria, est group of 2 trichobothria, et group of 3 trichobothria. Ventral (v) group 
neobothriotaxic, composed of 10 to 12 trichobothria. Pedipalp chela manus (Fig. 14A- 
C). Colour predominantly yellow; carinae, darker orange. Chela slender (length more 
than 3 times its width), with 5 distinct carinae; internodorsal carina continuous, with 
spiniform granules; subdigital carina vestigial, visible basally as a smooth ridge; ex- 
ternodorsal carina distinct, visible as a ridge of indistinct fused granules, almost 
smooth; digital carina a smooth ridge with few weak granules basally; dorsal 
secondary carina (dorsomedian) a smooth ridge with few strong granules basally and 
few weak granules distally; internoventral carina continuous, a weak ridge with spini- 
form granules; externoventral carina continuous, a granular ridge, running parallel to 
longitudinal axis of chela, its distal edge disconnected from external movable finger 
condyle and directed between external and internal movable finger condyles; ventro- 
median carina vestigial, reduced to few coarse granules basally; internal (interno- 
median) carina visible as a weak ridge with few spiniform granules; external (externo- 
median) carina weakly pronounced, visible as raised rows of small weak granules. 
Dorsal surface of chela shagreened, sparsely and weakly granular; internal surface sha- 
greened, sparsely granular, with a denser patch of spiniform granules dorsodistally; 
external surface shagreened, sparsely and weakly granular; ventral surface shagreened, 



902 L - MONOD & W. R. LOURENÇO 

sparsely and weakly granular, distal end without granules. A total of 15 trichobothria 
present on chelal manus; Db trichobothria located externo-basally on dorsal surface; 
Eb group (3 trichobothria) orthobothriotaxic, located basally on external surface; Esb 
very close to Eb group; Em absent; Est located distally, very close to Et group; Et group 
composed of 5 trichobothria, Et 1 located ventrally; V group comprising 4 trichobothria, 
V3 and V4 located in the basal half of manus, VI and V2 situated very distally. Pedipalp 
chela fingers (Fig. 14A-C). Dark reddish tan, with tips yellow-orange. Dorsal and ven- 
tral surfaces predominantly smooth, slightly shagreened and minutely pitted basally. 
Fingers longer than chela manus. Fixed finger with a very weak basal concavity, 
movable finger with corresponding lobe very weakly developed. Tips of fingers with 
pronounced distal hook. Fingers with distal diastema; edges of fingers composed of 2 
rows of denticles becoming fused basally, interrupted at regular intervals by stronger 
granules, each of them coupled with an accessory granule; rows fused into a single row 
above concavity on fixed finger and on notch of movable finger, single basal row run- 
ning towards base of fingers. A total of 11 trichobothria present on fingers; Dt located 
basally on dorsal surface of fixed finger; db located dorsally on internal surface, ap- 
proximately midway on the fixed finger; dsb, dst and dt on dorsal surface, in distal third 
of finger; eb located dorsally on external surface, midway on fixed finger, opposite db; 
esb, est and et in distal third of fixed finger; esb, est and et, opposite dsb, dst and dt, re- 
spectively; it and ib located in distal half of fixed finger. 

Coxosternal sclerites. Dark yellow to orange; smooth. Anterior margin of 
coxapohysis I smooth, with few weak granules, expanded but not sub-triangular in 
shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, pentagonal, sha- 
greened; median furrow deep, more pronounced in posterior half; posterior pit absent. 

Legs. Pale yellow. Predominantly smooth, dorsal surfaces of trochanter and 
femur finely and weakly granular. Tarsus with a ventromedian row of microspinules 
and with 2 rows of rigid "spinoid" ventral macrosetae; tarsi I-II: prolateral row with 
5-6 macrosetae, retrolateral row with 7 macrosetae; tarsi III-IV: prolateral row with 
6 macrosetae, retrolateral row with 7 macrosetae; setae acuminate; tarsal claws of 
equal length. 

Pectines and genital operculum (Fig. 15B). Colour pale yellow; genital oper- 
culum composed of a single subtriangular plate, posterior extremity truncate, longitu- 
dinal median furrow relatively deep, posterior notch pronounced; pectines with 9/9 
teeth. 

Hemiscorpius lepturus Peters, 1861 Figs 1A-B, 16-21, 27E-F, 36 

Hemiscorpius lepturus Peters, 1861a: 426-427, 8 figs. 

Syntypes (not examined). 1 S {ZMB 43a), 1 9 (ZMB 43b), IRAQ, Baghdad; 2 specimens 
(ZMB 43), IRAQ, Mendeli; YEMEN, Aden. 

Material examined. 1 9 (7408), IRAN (Kohkiluyeh), Charam road, 22.V.1974, 
A. Senglet. MHNG. 1 ? juv. (7412b), IRAN (Kohkiluyeh), Yasudj road, 25.V.1974, A. Senglet, 
MHNG. 1 S (2002/27), IRAN (Ghilan), 2 km W Cham-Khaled (Sea Star, Hotel Resort), nsl, 
37 o 14.922'N,50°15.037'E, 10.V.2002,B. Schätti. MffiVG. 2 6,2 9 ,IRAN (Khuzestan),Ahwaz 
region, 21.XI.1995, B. Masihpour, MHNG. 1 cî, IRAN, ? Lali, VI.1961, M. Vasserot, MNHN- 
RS 4332. 1 9 juv.. IRAN, 28.III.1961 , M. Vasserot, MNHN-RS 4333. 1 S , 1 2 , without locality, 
MNHN-RS 5232 (122-123). 2 6 , 2 9 , 4 6 subadult, IRAN (Khuzestan), Masched-Soleyman, R. 
Farzanpay ?, MHNG (SF 0006/13-20). 1 S juv., 1 9 , 2 9 juv., IRAN (Fare), ca. 90 km W Shiraz, 



HEMISCORPIIDAE FROM IRAN 



903 







Fig. 16 



Hemiscorpius lepturus. Male (MNHN-RS 5232): A, dorsal aspect; B, ventral aspect. Female 
paratype (MNHN, SF0006/17): C, dorsal aspect; D, ventral aspect. Scale line, 5 mm. 



904 L- MONOD & W. R. LOURENÇO 

14.IV.1970, G. Pretzman, K. Bilek and F. Ressi, NHMW3390. 1 S subadult, 3 9 , IRAN (Fare), 
ca. 5 km NE Persepolis, 20.IV.1970, G. Pretzmann, K. Bilek and F Ressi, NHMW3391. 1 3 
juv., 19,19 juv., IRAN (Fare), ca. 18 km SE Abadeh, 20JV1970, G. Pretzman, K. Bilek and 
F. Ressi, NHMW 3392. 

Distribution (completed with data from the literature). Esfahan, Fars, Hamadan, 
Kohkiluyeh, Khuzestan and Lorestan Provinces, south western Iran, eastern Iraq (Fig. 
36). The distribution area of Hemiscorpius lepturus mentioned by Fet (2000) also in- 
cludes the west of Pakistan. However, this is not confirmed by the present study. The 
species seems to be restricted to Iraq and the western and south western regions of Iran. 

Diagnosis. (1) Carapace longer than wide, very finely granular with anterior 
area smooth and minutely pitted, and area around median ocular tubercle very finely 
granular; lateral margins smooth, without granules; superciliary carinae weak, smooth; 
(2) pedipalps stout and bulky, with weaker and less distinct granular carinae than in any 
other Iranian Hemiscorpiidae, chela fingers slightly shorter than chela manus; (3) 
internodorsal carina of patellar protuberance smooth, with 1-2 weak granules; (4) pedi- 
palp patella orthobothriotaxic, external side with 13 trichobotria (1 est and 2 esb), ven- 
tral side with 3 trichobothria; (5) metasoma of males elongated and slender, with dor- 
sal carinae bearing sparse spiniform granules (weaker granules on anterior segments), 
and ventral and ventrolateral carinae of segments TV-V developed as ridges with weak 
spiniform granules (carinae of segment V with more distinct spiniform granules in pos- 
terior half); (6) telson of males strongly elongated, bearing a pair of blunt tuberculi- 
form processes at base of aculeus, lateral surfaces rugose, without distinct granules; (7) 
metasoma of females with dorsal carinae of segments I-IV and ventral and ventrolat- 
eral carinae of segment V bearing strong spiniform granules; (8) hemispermatophore 
with double lamellar hook located above distal transverse ridge and pointing distally. 

See also diagnosis of H. acanthocercus sp. n.. to which H. lepturus is closely 
related. 

Description. MALE (Figs 1A-B, 16A-B, 17A, 18A-C, E-F, 19A, 20, 21, 27F). 
Measurements of specimen MNHN-RS 5232. Carapace, length 6.4, posterior width 
5.8; distance between anterior lateral eyes 3.6, between posterior lateral eyes 4.0, 
between median eyes 0.2; diameter of median eyes 0.2. Pedipalp, femur length 6.0, 
width 2.2; patella length 5.8, width 2.2. chela length 10.8; manus length 6.0, width 4.4, 
depth 2.7; movable finger length 5.1; fixed finger length 3.8. Metasoma, segment I 
length 8.0, width 2.2; segment V length 10.4, width 1.6, depth 1.9; vesicle length 7.2, 
width 1 .7, depth 2.0. aculeus length 0.8. Total length 85.0. 

Carapace (Fig. 17A). Colouration yellow to light orange; median and lateral 
ocular tubercle black. Carapace dorsodistally flattened, longer than wide, almost 
rectangular in shape, with sides nearly parallel; median ocular tubercle weakly devel- 
oped, very low, distinctly situated anteriorly, with superciliary carinae weak and 
smooth; frontal concavity or notch well-developed; anterior lobes truncate; lateral 
ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriore. Carapace 
very finely granular, with few smooth patches; anterior area between median ocular 
tubercle and anterior margin smooth and minutely pitted, area around median ocular 
tubercle very finely granular; edges usually smooth, sometimes with very few weak 
granules below lateral ocular tubercles; anteromedian furrow narrow, suturiform, ante- 
riorly bifurcated; median longitudinal furrow shallow, continuous from anterior suture 



HEMISCORPIIDAE FROM IRAN 



905 




CO 



CO 

OS 




906 



L. MONOD & W. R. LOURENÇO 




Fig. 18 
Hemiscorpius lepturus, male (MNHN-RS 5232): A, pedipalp with trichobothrial pattern, dorsal 
aspect; B, chela, external aspect; C, idem, ventral aspect; E, femur and patella, external aspect; 
F, idem, ventral aspect. Female (MNHN-RS 5232): D, pedipalp, dorsal aspect. Scale line, 2 mm. 



furcation, running through ocular tubercle posteriorly into a deep triangular de- 
pression; postero-lateral furrow shallow; mesolateral furrow weekly developed, almost 
indiscernible. 

Mesosoma. Tergite colouration pale yellow. Tergite VII longer than wide. 
Tergite I without any depressions or carinae, tergites H- HI with a shallow median 
depression divided anteriorly by a weak ridge, tergites IV- VE with median carina 



HEMISCORPIIDAE FROM IRAN 907 

reduced to a weak, non-granular ridge surrounded by a pair of shallow, submedian 
depressions. Lateral and sublateral carinae absent on tergites I- VI, but present in 
posterior 2/3 of tergite VII, developed as ridges with indistinct granules, almost 
smooth. Surface of tergites finely granular; pre-tergites smooth, shiny and minutely 
pitted. Sternite colouration pale yellow. Sternite III smooth, shiny and minutely pitted, 
with a large, rough, very finely granular depression under each pectine; sternite IV- VI 
smooth, shiny and minutely pitted, without granulation or carinae, with a pair of 
shallow median furrows. Sternite VII smooth, shiny, minutely pitted medially and fine- 
ly granular laterally and posteriorly; a pair of weakly developed, smooth lateral carinae 
in posterior half; median carinae absent. Spiracles of book lungs crescent- shaped. 
Tergites and sternites VII longer than wide. 

Metasoma (Fig. 27F). Very elongated and slender, slightly shagreened, with fine 
scattered granules. Colour yellow to light orange. Segments I-IV with longitudinal dor- 
somedian furrow, dorsal carinae developed as ridges with scattered spiniform granules 
becoming more numerous on posterior segments, lateral carinae as smooth ridges 
without granules. Segments I-II with ventrolateral carinae developed as smooth ridges; 
segments III-IV with ventrolateral carinae as ridges of indistinct granules, almost 
smooth. Segment I with ventral carina almost absent, reduced to a weak ridge; 
segments II-HI with ventral carina a smooth ridge; segment IV with ventral carina a 
ridge of fused granules, almost smooth. Segment V: longitudinal dorsal furrow present 
in anterior half; dorsal carinae with small spiniform granules posteriorly; lateral carina 
developed as a weak ridge; ventrolateral and ventromedian carinae as ridges of indis- 
tinct granules, almost smooth anteriorly and with reduced spiniform granules in pos- 
terior half. 

Telson (Figs 1A-B, 27F). Vesicle yellow; aculeus darker, tan, due to stronger 
sclerotisation; vesicle elliptical or ovate, strongly elongated, with a blunt tuberculiform 
projection on each side at base of aculeus; ventrolateral furrows and ventromedian 
ridge absent; lateral surfaces irregular, with numerous weak granules; dorsal surface 
with numerous small spiniform granules and a median shallow depression. Macrosetae 
very sparse basally, becoming more numerous near base of aculeus. Aculeus short and 
stout, strongly curved, becoming markedly narrower approximately midway. 

Chelicerae. Colour pale yellow; teeth of fingers darker, orange. Teeth arrange- 
ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth 
bifid; movable finger with one subdistal tooth and one basal tooth in external series; 
distal external tooth smaller than distal internal tooth; cheliceral teeth without 
secondary serrations. 

Pedipalp short and stout. Pedipalp coxa and femur (Fig. 18A, E-F). Coxa with 
internoventral margin mostly smooth, with only few strong spiniform granules. Dorsal 
surface of femur yellow, internodorsal, internoventral and externo ventral carinae 
darker, orange. Femur short, stout (length less than 2.5 times its width), pentacarinate, 
with 4 distinct carinae; internodorsal carina developed as a strongly granular ridge; 
externodorsal carina reduced to a slightly raised row of granules; internoventral carina 
developed as a strongly granular ridge; externoventral carina as a weak granular ridge; 
ventromedian carina reduced to few granules and confined to base of femur. Dorsal 
surface shagreened, finely and densely granular, distal end smooth, without granules; 



908 



L. MONOD & W. R. LOURENÇO 




Fig. 19 
Hemiscorpius lepturus, pectines and genital opercula. ventral aspect: A, male (MNHN-RS5232); 
B, female (MHNG, SF 0006). Scale Une, 1 mm. 



HEMISCORPIIDAE FROM IRAN 



909 



fjÈk'jfà 



&ù 



l 



a 






\ 



fc~s 



s?- 

TO 



m 



£=£ 



Fig. 20 
Hemiscorpius lepturus, hemispermatophores of two males, external aspect: A, male from 
Masched-Soleyman (MHNG, SF 0006); B, male from Iran, without precise locality (MNHN-RS 
4332). Scale line, 0.5 mm. 



910 L. MONOD & W. R. LOURENÇO 

internal surface shagreened, with few strong scattered spiniform granules; external 
surface slightly shagreened; ventral surface finely and densely granular, with 
shagreened patches, distal end without granules. A total of 3 trichobothria present on 
femur; d located externobasally on dorsal surface; i located dorsobasally on internal 
surface; e located dorsobasally on external surface. Pedipalp patella (Figs 18 A, E-F) 
short and stout (length less than 2.5 times its width). Colour predominantly yellow; 
carinae slightly darker, orange. 7 carinae present, 6 of them distinct; internodorsal 
carina developed as a ridge of indistinct coarse granules; externodorsal carina as a 
weak ridge of indistinct granules, almost smooth; interno ventral carina as a ridge of 
indistinct granules, almost smooth; externo ventral carina as a smooth weak ridge, 
almost indiscernible; externomedian carina as a ridge of weak coarse granulaes, almost 
smooth. Dorsal surface finely and weakly granular, with small shagreened patches; 
internal surface shagreened, with few small scattered granules; external surface 
minutely pitted, with shagreened patches; ventral surface weakly and finely granular, 
with shagreened patches, distal end smooth and minutely pitted. Internal protuberance 
weak, bifid (internodorsal and internoventral tubercles separated), dorsal carina 
smooth, with 1-3 weak granules. A total of 19 trichobothria present on patella, 
orthobothriotaxy; dj located basally, external to internodorsal carina; d 2 located sub- 
medially on patella; d 3 absent; i in distal half of patella, located dorsally on internal 
surface. External (e) trichobothrial groups discernible and orthobothriotaxic: eb group 
composed of 5 trichobothria, esb group of 2 trichobothria, em group of 2 trichobothria, 
et group of 3 trichobothria; est very close to et group. Ventral (v) group orthobothrio- 
taxic, 3 trichobothria. Pedipalp chela manus (Figs 18A-C). Colour predominantly 
yellow; carinae and fingers darker, orange. Chela short and stout (length less than 2.5 
times its width), with 5 distinct carinae; internodorsal carina weakly developed, visible 
as a row of few granules, highlighted by a slightly darker line; subdigital carina 
vestigial, a smooth, weak ridge, more distinct basally; externodorsal carina vestigial, 
visible as a ridge of indistinct granules, almost smooth; digital carina vestigial, visible 
as a smooth, weak ridge more distinct basally; dorsal secondary carina (dorsomedian) 
vestigial, visible as a very weak, smooth ridge with a few weak granules basally; 
internoventral carina not continuous, developed as a weak, smooth ridge, with a few 
weak granules distally, less distinct basally; externo ventral carina continuous, devel- 
oped as a ridge of weak fused granules, smooth distally, running parallel to longitudinal 
axis of chela, its distal edge disconnected from external movable finger condyle and 
directed between external and internal movable finger condyles; ventromedian carina 
reduced to 1-2 coarse granules basally; internal (internomedian) carina vestigial, 
visible as a row of very few sparse granules; external (externomedian) carina weakly 
developed, visible as ridge of small indistinct granules. Dorsal surface smooth and 
minutely pitted, with patches of a subreticulate network of weak granular ridges and 
shallow shagreened dimples; internal surface sparsely and finely granular, with small 
shagreened patches, with a patch of stronger spiniform granules dorsodistally; external 
surface minutely pitted, forming an irregular and subreticulate network of weakly and 
finely granular ridges and shallow shagreened dimples; ventral surface smooth and 
minutely pitted, with few small shagreened patches. A total of 15 trichobothria present 
on chela manus; Db trichobothria located externobasally on dorsal surface; Eb group 



HEMISCORPIIDAE FROM IRAN 



911 





Fig. 21 
Hemiscorpius lepturus, hemispermatophores of two males, detail of capsular region, internal as- 
pect: A-B, male from Iran, without precise locality (MNHN-RS 4332), Dh (double hook), La 
(lamella), Lac (accessory lobe), Lb (basal lobe), Ld (distal lobe), Lp (posterior lobe), Tr (trans- 
verse ridge); C, male from Masched-Soleyman (MHNG, SF 0006). Scale lines, 0.5 mm. 

(3 trichobothria) orthobothriotaxic, located basally on external surface; Esb very close 
to Eb group; Em absent; Est located distally, very close to Et group; Et group composed 
of 5 trichobothria, Et 1 located ventrally; V group comprising 4 trichobothria, V3 and 
V4 located in the basal half of manus, VI and V2 located very distally. Pedipalp chela 
fingers (Figs 18A-C). Dark orange to reddish tan, smooth, minutely pitted basally. 



912 L. MONOD & W. R. LOURENÇO 

Fingers slightly shorter than chela manus. Fixed finger with a weak basal concavity, 
movable finger with corresponding lobe weakly developed. Tips of fingers with distal 
hook. Fingers with distal diastema; edges of fingers composed of 2 rows of denticles 
becoming fused basally, interrupted at regular intervals by stronger granules, each of 
these coupled with an accessory granule; rows fused into a single row above concavity 
on fixed finger and on notch of movable finger, single basal row running towards base 
of fingers. A total of 11 trichobothria present on chelal fingers; Dt located very basally 
on dorsal surface of fixed finger; db located dorsally on internal surface, in basal half 
of fixed finger; dsb, dst and dt on dorsal surface, in distal half of finger; eb located dor- 
sally on external surface in basal half of fixed finger, opposite db; esb, est and et in 
distal half of fixed finger; esb, est and et, opposite dsb, dst and dt, respectively; it and 
ib located midway on fixed finger. 

Coxosternal sclerites. Yellow; smooth and minutely pitted. Anterior margin of 
coxapohysis I smooth, with few weak granules, expanded but not sub-triangular in 
shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, pentagonal, smooth 
and minutely pitted; median furrow deep, more pronounced in posterior half; posterior 
pit absent. 

Legs. Pale yellow. Predominantly shagreened; dorsal surfaces of trochanter and 
femur finely and densely granular; ventral surface smooth and minutely pitted. Tarsus 
with a ventromedian row of microspinules and with 2 rows of rigid "spinoid" ventral 
macrosetae; tarsi I-II: prolateral row with 5-6, retrolateral row with 7 macrosetae; tarsi 
III-IV: prolateral and retrolateral rows with 7 macrosetae; tarsal claws of equal length. 

Pectines and genital operculum (Fig. 19A). Pale yellow; genital operculum 
composed of 2 ovoid plates; genital papillae short, not protruding from beneath 
operculum; pectines with 14-16 teeth, ususally 15/15. 

Hemispermatophore (Figs 20-21). Lamelliform, with complex capsule; distal 
lamella slender, basally curved, with a double hook pointing distad and located basally, 
flagellum approximately 1.5 times longer than capsule region; distal crest absent. 
Double lamellar hook (Dh) located above transverse distal ridge (Tr), the latter costate, 
distally strongly curved towards anterior margin, continuous from posterior to anterior 
margins, not merging with lamellar hook, reaching anterior margin of distal lamella 
below its hook. Capsule lamella (La) broad, with a deep longitudinal furrow and 
forming an acute angle to longitudinal axis of capsule, bearing a strong hook on its 
external surface and an accessory lobe (Lac) on its posterior margin; ventral margin 
straight. Distal lobe (Ld) forming a strong hook pointing anteriad; basal margin 
straight. Basal lobe (Lb) absent. Posterior lamella (Lp) costate. 

FEMALE (Figs 16C-D, 17B. 18D, 19B, 27E). Measurements of specimen 
MNHN-SF0006/18. Carapace, length 6.0, posterior width 5.6; distance between 
anterior lateral eyes 3.4, between posterior lateral eyes 3.8, between median eyes 0.2; 
diameter of median eyes 0.2. Pedipalp, femur length 5.7, width 2.1; patella length 5.6; 
width 2.1, chela length 10.7; manus length 6.0, width 4.5, depth 2.5; movable finger 
length 4.9; fixed finger length 3.6. Metasoma, segment I length 3.9, width 2.3; segment 
V length 5.7, width 1.6. depth 2.0; vesicle length 3.9, width 2.3, depth 2.3, aculeus 
length 1 .1 . Total length 56.0. 

Same characters as in males, except as follows. Carapace (Fig. 17B) less elon- 
gated than in male, almost as wide as long. Carapace finely granular, with larger 



HEMISCORPIIDAE FROM IRAN 913 

longitudinal median area smooth and minutely pitted; area around median ocular 
tubercle finely and densely granular. Mesosoma. Tergite VII as wide as long. Surface 
of tergites smooth, shiny and minutely pitted, finely granular laterally; pre-tergites 
smooth, shiny and minutely pitted, tergites VI- VII more granular than others. Sternite 
VII as wide as long. Metasoma (Fig. 27E). Short and relatively stout, slightly 
shagreened, with fine scattered granules. Carinae generally more granular than in 
males, spiniform granules stronger. Segments I-IV, dorsal carinae developed as ridges 
with spiniform granules (denser than in males), lateral carinae as weakly granular 
ridges. Segments I-II with ventrolateral carinae developed as weakly granular ridges, 
segment III-IV with ventrolateral carinae bearing weak spiniform granules. Segments 
I-II with ventral carina reduced to a smooth ridge; segments III-IV with ventral carina 
a ridge of weak spiniform granules. Segment V: dorsal carinae with weak spiniform 
granules; lateral carina vestigial, reduced to a row of scattered granules, ventrolateral 
carinae with spiniform granules; ventromedian carina with spiniform granules. Telson 
(Fig. 27E). Vesicle ovoid, globular and bulky; lateral surfaces smooth, slightly irre- 
gular. Aculeus without tubercles at its base, narrowing slightly less marked than in 
male. Pedipalp (Figs 18D). Slightly shorter and stouter than in males. Pedipalp chela 
fingers (Figs 18D). Fixed and movable fingers with slightly weaker basal concavity and 
lobe, respectively. Pectines and genital operculum (Fig. 19B). Genital operculum 
composed of a single subtriangular plate, posterior extremity truncate, longitudinal 
median furrow relatively deep, posterior notch pronounced; pectines with 9-11 teeth, 
usually 10/10. 

Intraspecific variation. The size of males can vary greatly from 52 mm up to 85 
mm body length, females display less variation in size. Females have pectines with 
8-11 teeth, usually 10-10, males haves pectines with 14-16 teeth. The same remark 
about pedipalps and metasoma of males can be given as for H. acanthocercus. 

Hemiscorpius persicus Binila, 1903 Figs 22-25, 36 

Hemiscorpion persicum Binila, 1903: 77-80. 

Material examined. Syntypes: 1 6 , IRAN (Slstän va Balûchestan), "Province Ssarbas", 
village Riss, 24.11.1901, N. Zarudny, ZISP 1120; 1 S subadult, IRAN (Sïstân va Balûchestan), 
"Province Ssarbas", on the road between village Riss and locality Kaptegin-Dukan, 24- 
26.11.1901, N. Zarudny, ZISP 1120. 

Distribution. Known only from Sïstân va Balûchestan Province, Eastern Iran 
(Fig. 36). 

Diagnosis. (1) Carapace longer than wide, shagreened and finely granular, with 
small smooth patches; lateral margins with small spiniform granules below lateral oc- 
ular tubercles; superciliary carinae finely granular; (2) pedipalps relatively stout and 
bulky, chela fingers slightly shorter or equal in size to chela manus; (3) internal protu- 
berance of pedipalpal patella with internodorsal carina bearing 1-2 strong spiniform 
granules, usually 2; (4) pedipalp patella orthobothriotaxic, external side with 13 tri- 
chobotria (1 est and 2 esb), ventral side with 3 trichobothria; (5) metasoma of males 
relatively elongated and slender, with dorsal carinae of segments I-V and ventral and 
ventrolateral carinae of segments IV-V bearing spiniform granules; (7) telson of males 
not strongly elongated, without blunt tuberculiform processes at base of aculeus. 



914 



L. MONOD & W. R. LOURENÇO 




Fig. 22 

Hemiscorpius persicus, male syntype (ZISP 1120): A, dorsal aspect; B, ventral aspect. Scale line, 
2.5 mm. 



HEMISCORPIIDAE FROM IRAN 



915 




Fig. 23 
Hemiscorpiuspersicus, male syntype (ZISP 1120): A, carapace, dorsal aspect; B, metasoma, lat- 
eral aspect. Scale lines, 1 mm. 



916 



L. MONOD & W. R. LOURENÇO 




Fig. 24 
Hemiscorpius persicus , male syntype (ZISP 1120), pedipalp with trichobothrial pattern: A, chela, 
dorsal aspect; B, idem, external aspect; C, idem, ventral aspect; D, femur and patella, dorsal as- 
pect; E, idem, external aspect; F. idem, ventral aspect. Scale line, 1 mm. 



HEMISCORPIIDAE FROM IRAN 



917 




Fig. 25 
Hemiscorpius persicus , male syntype (ZISP 1120): pectines and genital opercula, ventral aspect. 
Scale line, 0.5 mm. 



H. persicus is very different from all the others Iranian Hemiscorpius. Despite 
possessing a relatively slender metasoma, the male does not show the extreme sexual 
dimorphism of metasoma and telson observed in the other Iranian species and can 
therefore be easily distinguish from them. 

Description. MALE (description based on the syntype, ZISP 1120). 
Measurements of syntype (ZISP 1120). Carapace, length 4.0; posterior width 3.4; 
distance between anterior lateral eyes 2.0, between posterior lateral eyes 2.4, between 
median eyes 0.1; diameter of median eyes 0.1. Pedipalp, femur length 4.2, width 1.5; 
patella length 4.2, width 1.4, chela length 8.0; manus length 4.2, width 2.8, depth 1.7; 
movable finger length 4.1; fixed finger length 3.2. Metasoma, segment I length 3.2, 
width 1.4; segment V length 4.6, width 1.1, depth 1.3; vesicle length 2.8, width 1.1. 
depth 1.4, aculeus length 0.9. Total length 39.0. 

Carapace (Fig. 23 A). Colouration mostly uniformly dark yellow to orange; 
median and lateral ocular tubercles black. Carapace dorsodistally flattened, longer than 
wide, almost rectangular in shape, with sides nearly parallel; median ocular tubercle 
weakly developed, low, distinctly situated anteriorly, with weak, finely granular super- 



918 L. MONOD & W. R. LOURENÇO 

ciliary carinae; frontal concavity or notch well-developed; anterior lobes truncate; 
lateral ocular tubercles with 3 ocelli, the posterior one smaller than the 2 anteriors. 
Carapace shagreened, finely and densely granular, with a few smooth patches; larger 
smooth patches in median area of anterior lobes between median and lateral ocular 
tubercles; lateral margins with small spiniform granules below lateral ocular tubercles; 
anteromedian furrow narrow, suturiform, anteriorly bifurcated; median longitudinal 
furrow shallow, continuous from anterior suture furcation, running through ocular 
tubercle, becoming wider posteriorly before merging with a deep triangular depression; 
posterolateral furrow shallow; mesolateral furrow weakly developed, almost indis- 
cernible. 

Mesosoma. Tergite colouration dark yellow to orange anteriorly, becoming 
lighter (yellow) posteriorly. Tergites I-III with a very shallow median depression 
divided anteriorly by a very weak ridge, tergites IV- VII with median carina reduced to 
a very weak ridge surrounded by a pair of shallow, submedian depressions. Lateral and 
sublateral carinae absent on tergites I- VI, but present in posterior 2/3 of tergite VII, 
developed as ridges bearing strong spiniform granules. Surface of tergites shagreened, 
finely and densely granular; pre-tergites smooth. Sternite colouration dark yellow. 
Sternites III- VI smooth, slightly granular laterally, without carinae; sternite III with a 
pair of large, very finely and densely granular depressions covered by the pectines; 
sternites IV- VI with a pair of shallow median furrows. Sternite VII shagreened, finely 
and densely granular; pair of lateral carinae developed as granular ridges; median 
carinae absent. Spiracles of book lungs crescent-shaped. Tergites and sternites VII 
longer than wide. 

Metasoma (Fig. 23B). Relatively elongated and slender, shagreened, with very 
few scattered small granules. Colour dark yellow to orange, carinae darker. Segments 
I-IV with longitudinal dorsomedian furrow and with dorsal carinae composed of strong 
spiniform granules. Segments I-IV with lateral carinae developed as ridges bearing 
sparse spiniform granules, lateral carinae on segments ffl-IV less granular than those 
on segments I-II. Segments I-II with ventrolateral carinae developed as ridges with 
weak granules; segments III-IV with ventrolateral carinae as ridges of spiniform 
granules. Segment I-II with ventral carina a ridge with few very weak granules, almost 
smooth; segments III-IV with ventral carina a ridge with more distinct spiniform 
granules. Segment V: longitudinal dorsal furrow present in anterior half, dorsal carinae 
with strong spiniform granules; lateral carina vestigial, only visible in anterior 2/3 as a 
row of small scattered granules; ventrolateral and ventromedian carinae with spiniform 
granules. 

Telson (Fig. 23B). Vesicle yellow to orange; aculeus darker, tan, due to stronger 
sclerotisation; vesicle elliptical or ovate, without blunt tuberculiform projections at 
base of aculeus; ventrolateral furrows and ventromedian ridge absent; lateral surfaces 
smooth, with sparse weak granules; dorsal surface with very small spiniform granules 
and a median shallow depression anteriorly. Macrosetae very sparse basally, becoming 
more numerous near base of aculeus. Aculeus short and stout, strongly curved, 
becoming markedly narrower approximately midway. 

Chelicerae. Colour dark yellow; teeth of fingers darker, orange. Teeth arrange- 
ment as in Scorpionidae (see Vachon, 1963); fixed finger with median and basal teeth 



HEMISCORPIIDAE FROM IRAN 919 

bifid; movable finger with one subdistal tooth and one basal tooth in external series; 
distal external tooth smaller than distal internal tooth; cheliceral teeth without 
secondary serrations. 

Pedipalp relatively elongated and slender. Pedipalp coxa and femur (Fig. 24D- 
F). Internoventral margin of coxa with few strong spiniform granules. Dorsal surface 
of femur predominantly dark yellow-orange, carinae darker, orange. Femur relatively 
short and stout (length less than 2.5 times its width), pentacarinate, with 4 distinct 
carinae; internodorsal carina developed as a strongly and densely granular ridge, spini- 
form granules strong; externodorsal carina developed as a ridge with spiniform 
granules; internoventral carina as a granular ridge with strong spiniform granules; 
externoventral carina as a ridge with spiniform granules; ventromedian carina reduced 
to few spiniform granules and confined to base of femur. Dorsal surface shagreened, 
finely and densely granular, distal end smooth, without granules; internal surface 
shagreened, with a few strong scattered spiniform granules; external surface 
shagreened; ventral surface shagreened and weakly granular, distal end without 
granules. A total of 3 trichobothria present on femur; d located externobasally on dorsal 
surface; i located dorsobasally on internal surface; e located dorsobasally on external 
surface. Pedipalp patella (Fig. 24D-F) relatively stout (length less than 2.5 times its 
width). Colour predominantly dark yellow to orange; carinae slightly darker. 7 carinae 
present, 6 of them distinct; internodorsal carina developed as a granular ridge; externo- 
dorsal carina as a ridge of indistinct granules; internoventral carina as a strongly gran- 
ular ridge with spiniform granules; externoventral carina as a ridge of weak granules; 
externomedian carina as a weakly granular ridge. Dorsal surface shagreened, sparsely 
and weakly granular; internal and external surfaces shagreened; ventral surface 
shagreened, sparsely and weakly granular, distal end without granules. Internal protu- 
berance pronounced, bifid (internodorsal and internoventral tubercles separated), 
internodorsal carina with 2 strong spiniform granules. A total of 19 trichobothria 
present on patella, orthobothriotaxy; d 1 located basally, external to internodorsal 
carina; d 2 located midway on patella; d 3 absent; i in distal half of patella, located 
dorsally on internal surface. External (e) trichobothrial groups discernible and 
orthobothriotaxic: eb group composed of 5 trichobothria, esb group of 2 trichobothria, 
em group of 2 trichobothria, et group of 3 trichobothria; est midway between em to et 
groups. Ventral (v) group of 3 trichobothria (orthobothriotaxic). Pedipalp chela manus 
(Fig. 24A-C). Colour predominantly orange to red-brown; carinae reddish brown to 
almost black. Chela relatively stout (length less than 2.5 times its width) with 5 distinct 
carinae; internodorsal carina continuous, with spiniform granules, stronger basally; 
subdigital carina vestigial, visible basally as a smooth ridge with weak granules; 
externodorsal carina distinct, visible as a ridge of indistinct fused granules, almost 
smooth; digital carina as a smooth ridge with few weak granules basally; dorsal 
secondary carina (dorsomedian) as a smooth ridge with very few strong granules 
basally and few weak granules distally; internoventral carina continuous, visible as a 
weak ridge with very weak granules; externoventral carina continuous, developed as a 
granular ridge, becoming almost smooth distally, running parallel to longitudinal axis 
of chela, its distal edge disconnected from external movable finger condyle and direc- 
ted between external and internal movable finger condyles; ventromedian carina 



920 L. MONOD & W. R. LOURENÇO 

vestigial, reduced to few coarse granules basally; internal (internomedian) carina 
vestigial, reduced to few weak spiniform granules in distal half; external (externome- 
dian) carina indistinct, visible as raised rows of granules. Dorsal surface shagreened 
and sparsely granular; internal surface shagreened, sparsely granular, with a denser 
patch of bigger spiniform granules dorsodistally; external surface shagreened, sparse- 
ly and weakly granular; ventral surface shagreened and sparsely granular, distal end 
smooth and pitted. A total of 15 trichobothria present on chela manus; Db trichobothria 
located externobasally on dorsal surface; Eb group (3 trichobothria) orthobothriotaxic, 
located basally on external surface; Esb very close to Eb group; Em absent; Est located 
distally, very close to Et group; Et group composed of 5 trichobothria, Etj located ven- 
trally; V group comprising 4 trichobothria, V3 and V4 located in basal third of manus, 
VI and V2 located very far distally. Pedipalp chela fingers (Fig. 24A-C). Basally red- 
dish tan, becoming gradually lighter distally, tips of fingers yellow, edges black. Dorsal 
and ventral surfaces predominantly smooth and pitted, slightly shagreened basally. 
Fingers slightly shorter than chela manus. Fixed finger with weak basal concavity, 
movable finger with corresponding lobe weakly developed. Tips of fingers with pro- 
nounced terminal hook. Fingers with distal diastema; edges of fingers composed of 2 
rows of denticles becoming fused basally, interrupted at regular intervals by stronger 
granules, each of them coupled with an accessory granule; rows fused into a single row 
above concavity on fixed finger and on notch of movable finger, single basal row run- 
ning towards base of fingers. A total of 11 trichobothria present on chelal fingers; Dt 
located very basally on dorsal surface; db located dorsally on internal surface, in the 
basal half of fixed finger; dsb, dst and dt on dorsal surface, in distal half of finger; eb 
located dorsally on external surface, in basal 1/3 of fixed finger, opposite db; esb, est 
and et in distal half of fixed finger, opposite dsb, dst and dt, respectively; it and ib 
located in median 1/3 of fixed finger. 

Coxosternal sclerites. Dark yellow to orange; smooth, with shagreened patches. 
Anterior margin of coxapohysis I with few weak granules, expanded but not sub-trian- 
gular in shape. Sternum of type 2 (Soleglad & Fet, 2003), longer than wide, penta- 
gonal, slightly shagreened; median furrow deep, more pronounced in posterior half; 
posterior pit absent. 

Legs. Pale yellow. Predominantly shagreened; ventral surface smooth; dorsal 
surfaces of trochanter and femur finely and weakly granular. Tarsus with a ventro- 
median row of microspinules and with 2 rows of ventral rigid "spinoid" macrosetae; 
tarsi I-II: prolateral row with 4/5 macrosetae, retrolateral row with 5/6 macrosetae; tarsi 
III-IV: prolateral row with 5/6 macrosetae, retrolateral row with 6 macrosetae; setae 
acuminate; tarsal claws of equal length. 

Pectines and genital operculum (Fig. 25). Dark yellow; genital operculum 
composed of 2 subtriangular plates; genital papillae short, not protruding from beneath 
operculum; pectines with 13/12 teeth. 

Hemispermatophore. The presence of fully developed paraxial organs was 
assessed by cutting a small slit into the lateral pleural membrane between tergites and 
sternites. We did not have the permission to dissect the specimen. 

Intraspecific variation. The other specimen examined, a subadult male, has 
pectines with 15-15 teeth. 



HEMISCORPIIDAE FROM IRAN 921 

Remarks. Apparently more male and female specimens of H. persicus from the 
type locality are lodged in the collections of the Zoological Institute of Saint Petersburg 
(Fet, pers. comm.) but unfortunately we could not get this material on loan. 

DISCUSSION 

Taxonomic characters 

Sexual dimorphism. Apart from H. persicus, all Iranian species of the genus 
Hemiscorpius show a pronounced sexual dimorphism: mature males have a distinctly 
more elongated metasoma and telson than females (subadult males do not exhibit this 
sexual dimorphism) (Figs 4, 27). In H. acanthocercus sp. n., H. enischnochela sp. n., 
H. lepturus and probably also in H. gaillardi this dimorphism reaches extreme levels. 
Among scorpions extreme elongation of the metasoma in males is fairly common. It is 
encounter in the buthid genera Centruroides, Compsobuthus and Isometrus, as well as 
in genera belonging to families that are closely related to Hemiscorpiidae, i. e. 
Hadogenes Kraepelin, 1894 (Liochelidae), Heteroscorpion Binila, 1903 (Hetero- 
scorpionidae) and Urodacus Peters, 1862 (Urodacidae). 

Hadogenes spp. are ultralithophilous scorpions occurring in the Afrotropical 
region. They inhabit weathered rock outcrops where their extremely flattened mor- 
phology allows them to creep into narrow crevices and fissures. The very long tails of 
the males of most Hadogenes species are an essential adaptation to this specialized en- 
vironment. During courtship the male usually stings the female in the pleural mem- 
brane between the sternites and tergites. This behaviour is known as sexual sting. In 
very narrow cracks where courtship usually takes place, the male must reach around 
the side of the female's body to sting her (Leeming, 2003). 

The endemic Malagasy genus Heteroscorpion is closely related to the 
Urodacidae. As in Hadogenes, the four known Heteroscorpion species are well adapted 
to live in rocky habitats and occur in narrow fissures between layers of splintered but 
still standing rocks. The metasoma of males is also very elongated compared to 
females (Lourenço & Goodman, 2002, 2004; Lourenço et al., 2004). However, these 
scorpions do not show an extreme dorsoventral depression of the body as in 
Hadogenes. 

On the other hand, scorpions of the genus Urodacus are mostly burro wers, 
therefore they are usually very stocky. However, males of four species, i. e. U. elon- 
gatus L.E. Koch, 1977, U. megamastigus L.E. Koch, 1977, U. mckenziei Volschenk, 
Smith & Harvey, 2000 and U. varians Glauert, 1963, show an extreme elongation of 
their metasoma (L. E. Koch, 1977; Volschenk et al., 2000) but they are not strictly 
lithophilous. U. varians is a burrowing species and the others excavate shallow scrapes 
under stones and are sometimes found in rock crevices (Volschenk, pers. com.). 
Ecomorphological adaptations to rocky habitats also occur in the scorpionid genus 
Opisthophthalmus C.L. Koch, 1837, which is mostly composed of burrowing species 
(Prendini, 2001). Because of soil hardness in their habitats, O. austerus Karsch, 1879. 
O. karooensis Purcell, 1898 and O. pattisoni Purcell, 1899 have abandoned burrow 
construction in favour of a semi-litophilous existence under slabs of rock. All these 
species display morphological adaptations similar to those in Hadogenes, i. e. 
dorsoventral depression of the body and greatly elongated pedipalp chelae. 



922 



L. MONOD & W. R. LOURENÇO 




g 



ft 



pq 



E 









CO 

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E 







— 


„ 




a; 


a 


e 


o 
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— 


p 


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- 


Si 
o 


o 


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HEMISCORPIIDAE FROM IRAN 



923 




Fig. 27 
Hemiscorpius spp., metasoma, lateral aspect. Hemiscorpius maindroni: A, female syntype: B, 
male syntype. Hemiscorpius enischnochela sp. n.: C, female paratype (MNHG, Masdyed- 
Soleyman); D, male holotype. Hemiscorpius lepturus: E, female (MNHN, Ahwaz); F. male 
(MNHN-RS 5232). Scale lines, 2 mm. 



924 



L. MONOD & W. R. LOURENÇO 




Fig. 28 
Rhopalurus agamemnon (C. L. Koch, 1839), male, ventral view of central part of body with left 
pecten removed showing the granular depressed region of the third sternite III (from Lourenço 
& Cloudsley-Thompson, 1995). 



There is no information on the labels of the specimens studied referring to col- 
lecting methods, ecology, or habitat. However, the rugged, mountainous topography of 
their environment and their morphology, i. e. strongly elongated metasoma and 
dorsoventral depression of the body, indicate that Iranian Hemiscorpiidae are 
lithophilous species. Lithophilous scorpions adapted to live in narrow cracks and 
crevices of rocks are characterized by the following features: (1) dorsoventral depres- 
sion of the body; (2) elongation of the metasoma and/or pedipalps; (3) spiniform 
macrosetae on the telotarsi, operating in conjunction with highly curved telotarsal 
ungues to provide grip on rock surfaces (Prendini, 2001). All these adaptations are 
present in hemiscorpiids. 



HEMISCORPIIDAE FROM IRAN 



925 





Fig. 29 
Hemiscorpius maindroni, male syntype (MNHN-RS 4328), hemispermatophore: A, hemisper- 
matophore in toto; B, detail of the capsular region, internal aspect. Scale lines, 0.25 mm. 



926 



L. MONOD & W. R. LOURENÇO 



è 



f 




fffcte 



R) 



fA 



m 



B 



Fig. 30 
Iomachus politus . hemispermatophores of two males, external aspect: A, male from German East 
Africa, now TANZANIA (NHMW 1733): B. male from Handeni, Tanganyika, TANZANIA 
(CAS). Scale line, 1 mm. 



HEMISCORPIIDAE FROM IRAN 



927 




Fig. 31 
Iomachus politus, hemispermatophores of two male, detail of the capsular region, internal 
aspect, h (hook), La (lamella), Lac (accessory lobe), Lb (basal lobe), Ld (distal lobe), Lp (pos- 
terior lobe), Tr (transverse ridge): A, male from Handeni, Tanganyika, TANZANIA (CAS); 
B-D, male from German East Africa, now TANZANIA (NHMW 1733). Scale line, 0.5 mm. 



928 



L. MONOD & W. R. LOURENÇO 



: i 



.#, 



■if, 



m 




Fig. 32 
Opisthacanthus lecomtei, male (MNHN-RS 6264, Belinga, GABON), hemispermatophore: A, 
hemispermatophore in toto (scale line, 1 mm); B, detail of the capsular region, internal aspect 
(scale line, 0.5 mm). 



HEMISCORPIIDAE FROM IRAN 



929 




Fig. 33 
Opisthacanthus cayaporum Vellard, 1932, male (MHNG, Campos de Cayapos, BRASIL), 
hemispermatophore: A, hemispermatophore in toto; B, detail of the capsular region, internal 
aspect. Scale lines, 1 mm. 



930 



L. MONOD & W. R. LOURENÇO 




Fig. 34 
Hadogenes troglodytes (Peters, 1861b), male (NHMW 1703, South Rhodesia, now 
ZIMBABWE), hemispermatophore: A, hemispermatophore with in toto; B, detail of the capsular 
region, internal aspect, Dh (double hook), La (lamella), Lb (basal lobe), Ld (distal lobe), Lp 
(posterior lobe), Tr (transverse ridge). Scale line, 1 mm. 



HEMISCORPIIDAE FROM IRAN 



931 




Fig. 35 
Liocheles australasiae (Fabricius, 1775), male (BPBM, Tanna Island, VANUATU), hemisper- 
matophore: A, hemispermatophore in toto; B, detail of the capsular region, internal aspect, h 
(hook), La (lamella), Lb (basal lobe), Ld (distal lobe), Lp (posterior lobe), Tr (transverse ridge). 
Scale lines, 0.5 mm. 

In male hemiscorpiids the genital operculum is composed of two overlapping 
plates, whereas in females it is a single undivided plate. Like in other scorpions, males 
have bigger pectines with a higher pectinal teeth count than in females. In the H. lep- 
turus species group the carapace is usually more elongated in males (longer than wide) 



932 L. MONOD & W. R. LOURENÇO 

than in females (almost as wide as long) (Fig. 26). Likewise, tergites and sternites VII 
are longer than wide in males, while as wide as long in females. Like the dimorphism 
in metasoma length, this difference is present only in the final instar. Immature males 
and females look much the same and can usually be distinguished only through exam- 
ination of their genital operculum and pectines. 

Granular depressions on sternites III. In males and females of Hemiscorpius 
acanthocercus sp. n., H. enischnochela sp. n., H. gaillardi, H. lepturus and H. persi- 
cus, the foremost sternite exhibits a pair of large, rough, very finely granular depres- 
sions. This feature is morphologically very similar to the stridulatory apparatus of 
species belonging to the neotropical buthid genus Rhopalurus (Fig. 28). The 
Rhopalurus stridulatory apparatus consists of large granular areas on the third sternite, 
which are located under the pectines and are used as a rasp. Pectinal teeth with stria- 
tions on their dorsal surface are rubbed against these areas in order to produce sound. 
Stridulation have been reported for all Rhopalurus species (in both males and females), 
but only the stridulatory sounds of larger species are audible to humans (Lourenço & 
Cloudsley-Thompson, 1995; Lourenço, Huber & Cloudsley-Thompson, 2000). 
Production of stridulatory sound has never been reported for any Hemiscorpius 
species, but this can be attributed to the small or medium size of these scorpions 
(regardless of the length of the metasoma), which probably produce barely audible 
sounds. The dorsal surfaces of their pectinal teeth seem to be finely granular. However, 
this character could not be assessed accurately because of the very reduced size of the 
structures. More thorough investigations by means of a stereo-electron microscope are 
needed. 

Hemispermatophores . The hemispermatophores of Hemiscorpiidae (Figs 7, 20, 
21, 29) share morphological similarities with hemispermatophores of Hadogenes 
species (Fig. 34) and of a liochelid group composed of Iomachus politus (Fig. 30-31) 
and Opisthacanthus (Opisthacanthus) species (Figs 32, 33). The general shape of 
hemispermatophores is the same in these taxa, i. e. distal lamella longer than the basal 
part, lamellar hook situated above the distal transverse ridge and not merging with it. 
However, Hemiscorpiidae have hemispermatophores with the distal lamella bearing a 
double hook, whereas all the other liochelid genera except Hadogenes have distal 
lamellae with a single hook. 

The capsular median structure of hemiscopiid hemispermatophores is complex 
(Figs 7B-C, 21A-C, 29B), composed of 4 distinct lobes/lamellae: (1) capsule lamella 
(La) complex, bearing a strong hook on his external surface and an accessory lobe 
(Lac) pointing to the posterior side. (2) distal lobe (Ld) forming a strong hook pointing 
anteriad, (3) basal lobe (Lb) very reduced, (4) posterior lobe (Lp) costate. Iomachus 
politus and Opisthacanthus (Opisthacanthus) species (both African and neotropical 
representatives) possess hemispermatophores (Figs 30-33) very similar to those 
encounter in hemiscorpiids. i. e. with distal lobe with a strong hook pointing anteriorly 
and with a complicated capsular lamella bearing an internal hook and an additional 
basal lobe. In Hadogenes (Fig. 34) and other liochelid genera the capsule lamella is 
much more simple, without any hook or accessory lobe. In Liocheles Sundevall, 1833 



HEMISCORPIIDAE FROM IRAN 933 

the distal lobe is not modified into a large hook and the basal lobe is much more de- 
veloped (Fig. 35). 

Based on this analysis Hadogenes and the group composed of Iomachus politus 
and Opisthacanthus {Opisthacanthus) spp. are considered to be liochelid taxa closely 
related to hemiscorpiids. The double lamellar hook can therefore be considered as a 
plesiomorphy for Hemiscorpiidae and Liochelidae, and the single lamellar hook is apo- 
morphic for all Liochelidae except Hadogenes. On the other hand, the complex capsule 
lamella can also be considered as a plesiomorphy for Hemiscorpiidae and Liochelidae. 
However, the derived, more simple structure examined in Hadogenes and other 
liochelid genera (e. g., Liocheles) is assumed to have evolved independently in these 
two groups. 

BlOGEOGRAPHICAL AND GEOLOGICAL IMPLICATIONS 

Iomachus politus occurs in East Africa (Ethiopia, Kenya Tanzania, Uganda) and 
the only African representative of the subgenus Opisthacanthus, O. (O.) lecomtei 
(Lucas, 1858), was recorded from approximately the same latitude in West Africa 
(Cameroon, Gabon). The neotropical species of Opisthacanthus (O.) are found in 
Central and South America and Hadogenes is a South African genus. Hemiscorpius is 
present all over the Middle East (Iran, Iraq, Oman, Pakistan, Saudi Arabia, United Arab 
Emirates and Yemen), but has also some representatives in north-eastern Africa 
(Djibouti, Eritrea, Somalia), not far from the distribution area of Iomachus politus . The 
presence of Hemiscorpiidae in Djibouti, Eritrea, Somalia, and on the island of Socotra 
indicates that the ancestor of Hemiscorpius was probably present on Gondwanaland. 

Since Hemiscorpiidae is the sister group of Liochelidae and Hemiscorpius 
shares characters with certain African liochelids, i. e. Iomachus politus and 
Opisthacanthus (O.) lecomtei, the occurrence of Hemiscorpius in Africa is probably 
due to a Gondwanian origin and not a recent invasion from the Middle East (Laurasia). 
However, the Iranian Hemiscorpius, i. e. the H. lepturus species group, are very 
divergent from the Hemiscorpius species from Africa, Oman, Saudi Arabia, United 
Arab Emirates and Yemen. The African species of Hemiscorpius do not show the 
highly derived features that their Iranian relatives possess, i.e. extreme sexual dimor- 
phism and cytotoxic venom. Iranian Hemiscorpius have probably been separated from 
the other hemiscorpiids for quite a long time. H. persicus represents an exception 
among Iranian Hemiscorpius; this species is more similar morphologically to African 
and Arabian representatives of the genus than to the Iranian ones. This could be the 
result of a more recent invasion from the Arabian peninsula. 

An old separation of the Iranian hemiscorpiids can be explained by the geolo- 
gical history of the region. In the Precambrian and Palaeozoic the Cimmerian terrane, 
which included Iran, was part of Gondwana (Fig. 37, 38A). The common ancestor of 
Liochelidae and Hemiscorpiidae was probably widespread on this super-continent. 
Approximately 250 Ma BP (Fig. 38B) the Neotethys ocean started to open (Golonka, 
2000; Scotese, 2000; Stampfli & Borei, 2002), inducing the rift of the Cimmerian super 
terrane off the northern margin of Gondwana and its northward drift toward Laurasia. 
Iran was separated from Gondwana in the Middle Triassic. The ancestor population of 
the Middle East hemiscorpiids was probably isolated on this rafting continent and 
diverged from its relatives. 



934 



L. MONOD & W. R. LOURENÇO 




Fig. 36 
Distribution map of Hemiscorpius spp. In Iran and Iraq. 



HEMISCORPIIDAE FROM IRAN 



935 




936 



L. MONOD & W. R. LOURENÇO 



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Palaeogeographic reconstructions of the Tethys Oceans and surrounding landmasses 280, 250 
and 220 Ma BP. The opening of the Neotethys separated the Cimmerian Superterrane from 
Gondwana, inducing the Cimmerian Terrane to drift toward Eurasia. Its accretion to Laurasia 
occurred 220 Ma BP when the oceanic crust of the Palaeotethys was completely subducted to the 
north. Landmasses are indicated by grey shading and current distribution areas of 
Hemiscorpiidae and related Liochelidae are indicated by pictograms. Positions of equator and 
tropics are also shown. Modified from Stampili & Borei, 2002. 



HEMISCORPIIDAE FROM IRAN 937 

The Hemiscorpiidae lineage was supposedly separated from the Liochelidae 
lineage before the rifting of the Cimmerian terrane, probably by climatic changes on 
Gondwana that occurred during the massive Permian global warming (290-250 Ma 
BP; Rees, 2002). Four main terrestrial climate zones (biomes) have been determined 
from fossil floras and climate-sensitive sedimentary rocks for the Late Carboniferous- 
Early Permian (300-280 Ma BP) (Gibbs et al., 2002; Rees et al, 1999; Rees et al, 
2002; Scotese, 2000; Willis & McElwain, 2002). In both hemispheres, climates 
gradually changed from the equator to the poles as follow: (1) tropical everwet (equa- 
torial region), (2) subtropical desert, (3) cool temperate and (4) cold (polar region). At 
that time Gondwana was situated south of the equator and was therefore under mostly 
lush and green vegetation (cool temperate), except for its north-eastern part where arid 
conditions prevailed (subtropical desert). Throughout the Permian, the climate change 
from icehouse to hothouse conditions (Rees, 2002). With global warming and 
increasing aridity, tropical forests disappeared and cool temperate biotopes were 
segregated in the southern part of Gondwana as deserts spread across central Pangea; 
tropical everwet biomes were only well-developed on the Chinese microcontinent 
(Rees, 2002; Scotese, 2000). Therefore the hemiiscorpiid ancestor adapted to an envi- 
ronment (in Africa and Arabia) that became more and more arid during the Permian, 
while liochelids remained in the more humid and cool ecosystems prevailing in the 
southern part of Gondwana (Scotese, 2000; Willis & McElwain, 2002). By the Late 
Triassic-Early Jurassic (210-200 Ma BP), large subtropical deserts formed a climatic 
barrier (Rees, 2002; Rees et al., 1999) between the liochelid ancestor in warm tem- 
perate forests of Southern Gondwana and the African hemiscorpiid ancestor in the 
Equatorial summerwet biotopes of the Arabian region, and interrupted gene flow 
between these two lineages. Most of the modern scorpion families probably emerged 
at the end of the Palaeozoic Era, when changing biomes, high rates of extinction and 
the resulting isolation of surviving populations in scattered "biotopes islands" were 
favourable for diversification. 

After being detached from North and South America, Africa began to rotate 
counterclockwise toward Eurasia and closed the western part of the formerly extensive 
Neotethys sea (Brown & Lomolino, 1998; Golonka, 2000). A bridge between Asia and 
Africa was formed by Arabia, following their collision in the middle Tertiary (35 Ma 
BP) that created the Zagros mountains of Iran. Then hemiscorpiids could have easily 
established new colonies through dispersal from the Arabian peninsula to Eurasia. 
Another hypothesis is the drift of microcontinents from the northern margin of 
Gondwana towards the southern margin of Laurasia during the last 200 million years, 
scorpions could have been carried on them to Eurasia. 

ACKNOWLEDGEMENTS 

We express our sincere gratitude to all individuals and institutions that assisted 
in the realization of the present study: the Natural History Museum of Geneva for 
providing working space and access to the collection; Dr Peter Schwendinger for 
kindly reviewing the manuscript and for his extensive help, constructive comments and 
suggestions; Dr Jürgen Gruber from the Natural History Museum of Vienna for 



938 L- MONOD & W. R. LOURENÇO 

arranging loans, providing information on collectors and help in deciphering old labels; 
Dr Victor Fet and Dr Graeme Lowe for kindly allowing the examination of the type 
specimens of Hemiscorpius persicus and for constructive criticism on the manuscript; 
Dr Erich Volschenk for further discussion; to M. Claude Ratton for taking the habitus 
photographs. 

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Vachon, M. 1963. De l'utilité, en systématique, d'une nomenclature des dents des chélicères 

chez les scorpions. Bulletin du Muséum National d'Histoire Naturelle, Paris (2), 35(2): 

161-166. 
Vachon, M. 1966. Liste des scorpions connus en Egypte, Arabie, Israel, Liban, Syrie, Jordanie, 

Turquie, Iraq, Iran. Toxicon 4: 209-218. 
Vachon, M. 1974. Etude des caractères utilisés pour classer les familles et les genres de scor- 
pions (Arachnides). Bulletin du Muséum National d'Histoire Naturelle, 3 e série 

140(104): 857-958. 
Vellard, J. 1932. Mission scientifique au Goyaz et au Rio Araguaya. Scorpions. Mémoires de 

la Société Zoologique de France 29(6): 539-556. 
Volschenk, E. S., Smith, G. T. & Harvey, M. S. 2000. A new species of Urodacus from 

Western Australia, with additional descriptive notes for Urodacus megamastigus 

(Scorpiones). Records of the Western Australian Museum 20: 51-61 . 
Walckenaer, C. A. 1847. Araneae (pp. 365-564). In: Walckenaer, C. A. & Gervais, R (eds). 

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Oxford. 



Revue suisse de Zoologie 112 (4): 943-961; décembre 2005 



New genera and species of nematode parasites (Drilonematoidea: 
Ungellidae) from coelomic cavity of Neotropic acanthodrilids 
deposited in the Natural History Museum of Geneva, Switzerland 

Elena S. IVANOVA 1 & Claude VAUCHER 2 

institute of Parasitology, Russian Academy of Science, 33 Leninskii pr., Moscow, 

119071, Russia. 
2 Muséum d'histoire naturelle, case postale 6434, CH-1211 Genève 6, Switzerland. 



New genera and species of nematode parasites (Drilonematoidea: 
Ungellidae) from coelomic cavity of Neotropic acanthodrilids deposited 
in the Natural History Museum of Geneva, Switzerland. - Two new 

genera and two new species of nematodes (Ungellidae, Drilonematoidea) 
parasitic in Neotropic acanthodrilids from the collection of the Natural 
History Museum of Geneva, Switzerland, are described. Yagansiella longi- 
collis gen. n., sp. n. and Ungella chileana sp. n. are parasitic in Yagansia 
spatulifera, whereas Ungella micronychium sp. n. in Y. diversicolor and 
Patagoniella capitoporus gen. n., sp. n. in Y. papillosa. Yagansiella longi- 
collis gen. n., sp. n. resembles Ungella in general morphology but differs by 
anterior position of ovary top cell and posterior vulva position, charac- 
teristic body and caudal fimbriate organs shape. U. chileana sp. n. is distin- 
guished by large circular fimbriate organs in tail, female body shape and 
arrangement of female genital tube and spicule shape. Ungella micro- 
nychium sp. n. differs by minute cephalic hooks and shape of fimbriate 
organs and spicules and gubernaculum. Patagoniella capitoporus gen. n., 
sp. n. is distinguished by anteriormost position of excretory pore, anterior 
vulva position and shape of fimbriate organs and spicules and guberna- 
culum. 

Keywords: Earthworms, nematode parasites - Ungellidae - Drilonema- 
toidea - Yagansiella longicollis gen. n., sp. n. - Ungella chileana sp. n. - 
Ungella micronychium sp. n - Patagoniella capitoporus gen. n., sp. n. - 
Yagansia spatulifera - Yagansia diversicolor - Yagansia papillosa. 

INTRODUCTION 

Earthworm collection deposited in the Natural History Museum of Geneva 
contains European lumbricids as well as annelids from around the globe. From 1 to 6 
specimens of 142 species of earthworms from museum collection were studied on the 
presence of nematodes (Drilonematoidea, Rhabditida) parasitic in coelomic cavity of 
earthworms. The bulk of museum collection consists of Neotropic glossoscolecids and 
acanthodrilids. So far, the only drilonematid parasite had been recorded from these 



Manuscript accepted 07.04.2005 



944 S. IVANOVA & C. VAUCHER 

earthworms. A new genus and the new Ungella species from coelomic cavity of 
Yagansia spatulifera, the new Ungella species from Y. diver sicolor and the new genus 
from Y. papillosa all belonged to Ungellidae family are described below. 

MATERIAL AND METHODS 

Earthworms were stored in ethanol after formaldehyde fixation which found did 
not affect the nematode morphology. Earthworms were dissected at anterior end and 
body cavity content was rinsed out and nematode parasites removed. Four specimens 
of Yagansia spatulifera, six ones of Y. diver sicolor and three of Y. papillosa were 
dissected. Three females and three males of Yagansiella longicollis gen. n., sp. n. and 
a male and a female of Ungella chile ana sp. n. were obtained from coelomic cavity of 
single specimen of Yagansia sparulifera. Two males of U. micronychium sp. n. were 
recovered from a specimen of Y. diversicolor and nine males, five females, three juve- 
niles and fragments of Patagoniella capitoporum gen. n., sp. n. from a specimen of Y. 
papillosa. Nematodes were processed into glycerol using slow evaporation technique 
(Seinhorst, 1959) and mounted on slides. Drawings and measurements were made 
using "Jenaval" microscope. De Manian indices and absolute measurements are given, 
where D is maximal diameter (if not indicated), Oes is oesophagus length, Cd is tail 
length and Ex is distance from base of cephalic hooks to excretory pore. A name 
"fimbriate organs" was used when describing caudal structures of sensory nature 
(phasmids?) instead of "suckers" (Ivanova & Hope, 2004). 

DESCRIPTIONS 

Yagansiella gen. n. 

Diagnosis. Rhabditida, Drilonematoidea, Ungellidae. Very long and thin neck 
region and swollen body. Spacious pseudocoel. Paired blade-like cephalic hooks 
present. Cuticular ridges situated posteriorly to mouth. Amphids elliptical. Stoma 
lacking; oesophagus with corpus, isthmus and bulb. Nerve ring situated around 
isthmus. Excretory pore at isthmus level. Large paired deep transversely oriented 
fimbriate organs in caudal region with prominent lip-like basal rim. Females mono- 
delphic, prodelphic, spermatheca set-off, distal tip of ovary near oesophageal base, 
vulva posterior with oblique vagina. Males monorchic; spicules paired and curved; 
thick gubernaculum with dorso-caudal apophysis and crurae, bursa absent. 

Type and only species. Yagansiella longicollis sp. n. 

Etymology. Generic name refers to annelid host of the species. 

Yagansiella longicollis sp. n. Figs 1-2 

Measurements. Holotype female . L = 1216 pm; D = 153 pm; Oes = 213 pm; 
Ex = 129 pm; NR = 105 pm; Cd = 123 pm; Ovum = 54 x 27 pm; V% = 89.44%; a = 
7.95; b = 5.71; c = 9.89. 

Paratype females (n = 2).L= 1 185-1458 pm; D = 108-140 pm; Oes = 215 pm; 
Ex = 131-135 pm; NR = 81-90 pm; Ova = 52-54 x 25-27 pm; Cd = 112-127 pm; V% 
= 90.55-90.81; a = 10.41-10.97; b = 5.51-6.78; c = 10.58-11.48. 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 945 




Fig. 1 
Yagansiella longicollis gen. n., sp. n. Female. A - paratype, entire body; B - paratype, head 
end; C - holotype, anterior end; D - holotype, posterior end; E - holotype, spermatheca 
region. All in lateral position. Scale bars in firn. 



Paratype males in = 3). L = 1093.67±142.1 (930-1 185) /im; D = 83±25.2 (60- 
110) /im; Oes = 188.33±9.5 (179-198) /im; Ex = 118.33+6.7 (114-126) /im; NR = 
99.33±7.8 (93-108) /im; Cd = 200.67±8.1 (195-210) /im; Sp (arc)= 104.33±5.5 (99- 
110) /im; Sp (chord)= 67±5.6 (62-73) /im; Gub = 30.33±2.1 (28-32) /im; a = 14.32±5.7 
(8.45-19.75); b = 5.8+0.6 (5.2-6.3); c = 5.47±0.9 (4.43-6.02). 



946 S. IVANOVA & C. VAUCHER 

Description. Adults. Very long and thin neck region and swollen body. Body 
starts to widen gradually behind the base of esophagus and turns uniformly wide from 
the point of where genital tube forms flexure. Terminal portion of tail filamentous. 
Epidermis thick. Spacious pseudocoel. Cuticle 1 thick, annulated at both ends, with 
annules 1.5 piva thick. Head bluntly rounded. No cephalic sensilla present. Head with 
paired, independently-moveable, submedian, claw-like cephalic hooks situated on 
anterior surface of head; base of each hook consists from pair of thick equal pointed 
diverged processes 2-3 piva long; longer pointed blade directed dorsad. Minute oral 
aperture shifted slightly dorsad near distal tips of blades. Stoma absent. Cuticular 
ridges on dorsal side just beneath mouth. Amphid faint, situated close to hooks base; 
its aperture transversely elongated. Anteriormost portion of esophagus slightly 
expanded. Slightest constriction of corpus at 10-12 /<m from head end. Oesophagus 
slender, muscular, from very long corpus 6-8 /<m wide, thinner long isthmus and bulb 
with granular secretion of dorsal oesophageal gland. Bulb elongated with nearly 
rectangular base, 30-40 pi m long and 10-11 pi m wide. One large nucleus of dorsal gland 
at posterior of bulb and two smaller ones of subventral glands situated more anteriorly. 
Nerve ring wide, encircling anterior part of isthmus. Excretory pore in 20-40 piva 
behind nerve ring, 2 pirn wide. Excretory duct strongly cuticularized, 2 piva wide and 
105-153 piva long. Excretory cell large, with granular content, observed untili mid- 
body. Cardia from two elongated cells. Intestine thick- walled, contains debris. Large 
paired symmetrically placed deep transversely oriented fimbriate organs in caudal 
region with prominent lip-like basal rim. 

Females. Blades of head hooks 7.67±0.6 (7-8) piva long, hook base 5.33±1.2 (4- 
6) piva wide. Amphidial aperture 3 x 2 piva. Oesophagus 9.33±0.6 (9-10) piva wide at 
anterior, corpus 8 pirn and isthmus 6 pirn wide. Bulb 37±2.6 (35-40) piva long and 
10.67±0.6 (lO-ll)^m wide. Genital tube starts in 150-201 pi m behind the oesophageal 
base, sometimes extending prior to spermatheca, which situated in 388.33±30.4 (363- 
422) pi m from anterior. Tube thick, runs posteriorly to fimbriate organs, then turns back 
by convoluted course and forms rounded or elongated off-set spermatheca 74.67±17.5 
(60-94) x 50±26.5 (30-80) piva with thick walls filled with spherical or bean-like sper- 
matozoa 1.5-2 piva, in diameter. Descending and ascending branches of genital tube 
spirally twisted. Spermatheca joined with thick-walled oviduct by thin tube. Thin- 
walled muscular uterus obscured by twists of ovary. Muscular vagina 36.33±4 (32-40) 
^m long. Vulva at posterior. No post-uterine sack present. Anus in 43.33±7.6 (35-50) 
piva posteriorly to vulva. Up to 6 eggs with thin finely punctated shells. Fimbriate 
organs with deep chamber, 39±5.3 (35-45) piva long and 66±5.3 (60-70) pi m wide with 
prominent lip-like muscular rim 5-7 piva thick. A channel inside tail end runs through 
fimbriate organs. Terminal portion of tail 50.67±4 (47-55) long and 5.33±.6 (5-6) piva 
wide. 

Males. Resembles females in body shape but a little shorter and less pro- 
minently expanded behind neck region. Amphidial aperture 4.67±1.2 (4-6) x 2.33±0.6 
(2-3) piva. Testis reflexes at 333.67±94.2 (255-430) piva from oesophageal base, flexure 
curved, 180.67±19.9 (159-198) piva long. Developing spermatocytes in 7-8 rows, 
small, spherical (2 piva in diameter). Testis very wide, ejaculatory duct muscular, sepa- 
rated from vas deferens by constriction. Spermia in vas deferens spherical or bean-like. 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 947 




Fig. 2 
Yagansiella longicollis gen. n., sp. n. Male. A - paratype, entire body; B 
C - paratype, anterior end; D - paratype, spicules and gubernaculum; E - 
lateral position. Scale bars rnjim. 



- paratype, head end; 
paratype, tail. All in 



resemble those from female spermatheca. Very long strongly cuticularized 
sickle-shaped paired spicules, with elongated manubria 12.33±1.2 (11-13) ^m long 
and 10.33±1.5 (9-12) pirn wide. Spicule shafts 5.66+0.6 (5-6) pirn wide at mid-length 
tapering gradually to rounded tips and bearing two front ridges. Gubernaculum 
massive, dark-brown in colour, with dorso-caudal apophysis 12.33±3.1 (9-15) //m long 
and crurae as long as gubernaculum body. A slit in gubernaculum body parallel to 
spicules. Nine pairs of sub ventral precloacal and four (five) pairs of postcloacal papilli- 
form sensilla. First pair of postcloacal sensilla subventral, situated between anus and 



948 s - IVANOVA & C. VAUCHER 

fimbriate organ; second one dorsal in front of fimbriate organ; third subventral between 
fimbriate organ and terminal portion of tail; fourth subdorsal close to the latter and fifth 
lateral symmetrically to the fourth in a paratype specimen. Copulatory muscles pro- 
minent but bursa lacking. Fimbriate organs smaller than in females: 24.67±3.1 (22-28) 
pm long and 22.33±2.3 (21-25) pm wide with basal rim 4-5 //m thick. Thick channel 
ends blindly posterior to fimbriate organ. Filamentous portion of tail 71±3.6 (68-75) 
pm long and 4 pm thick. 

Type host and locality. Yagansia spatulifera (Mich.) AF 4262, Chile, 
Villarica Vulcan, 1200 m, 14.02.01, coll. and identified by C. Czudi. 

Type habitat. Coelomic cavity at mid-body region. 

Type specimens. Holotype female MHNG 35445 INVE and paratype female 
and two paratype males MHNG 35446 INVE deposited in Natural History Museum 
(Geneva, Switzerland). 

Differential diagnosis. The present species is closely related to Ungella 
Cobb, 1928. From the latter genus, it can be distinguished by position of extremity of 
ovary which is anterior in Yagansiella and posterior in Ungella and vulva position 
which places in front of anus whereas it is typically post-median. It differs from 
Ungella as well by having swollen body posterior and large and broad fimbriate organs 
of different shape. From Plesioungella Yeates et al., 1998, which it resembles by 
swollen posterior, it differs by having paired spicules. From Onychonema Baylis, 1943 
it differs by presence of prominent fimbriate organs which lacked in the former. From 
Acanthungella (Ivanova & Hope, 2004) it differs in lack of somatic sensilla. 

Etymology. The species name derived from Latin words longus (long) and 
collum (neck) and reflects appearance of the nematodes. 

Ungella chileana sp. n. Figs 3-4 

Measurements. Holotype male. L = 1205 pm; D = 48 pm; Oes = 182 pm; Ex 
= 149 //m; NR = 105 pm; Cd = 183 //m; Sp (arc) = 70 pm; Sp (chord) = 51 pm; Gub 
= 13 pm; a = 25.1; b = 6.63; c = 6.58. 

Paratype female. L = 1218 pm; D = 90 pm; Oes = 215 pm; Ex = 170 pm; NR 
= 130 pm; Cd = 163 pm; V% = 64.5; a = 13.53; b = 5.67; c = 7.42. 

Description. Adults. Anterior of body thin. Terminal portion of tail filamentous. 
Cuticle 1 pm thick, annulated. Epidermis well-developed. Head truncate, with paired, 
independently-moveable, submedian cephalic hooks situated on anterior surface of 
head. Each hook from thick bone head-like base and pointed outwardly-curved blade 
and directed dorsad. Mouth shifted slightly dorsad near distal tips of blades. Stoma 
absent. Amphid pocket-like, situated close to hooks base; amphidial aperture trans- 
versely elongated. Anteriormost portion of esophagus slightly expanded. No head 
sensilla visible. Oesophagus straight, slender, from long finely muscular corpus, no 
distinct isthmus and narrow bulb with 3 nuclei of oesophageal glands. Base of 
oesophagus not rounded. Nerve ring situated a little farther back from mid-oesophagus. 
Excretory pore opposite anterior of bulb, 1 .5 pm thick. Excretory duct 1 .5-2 pm wide 
90- 100;/ m long. Excretory cell with fine granular content visible till mid-body. Cardia 
prominent from two large oval cells. Intestine thin. 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 949 




Fig. 3 
Ungella chileana sp. n. Male holotype. A - entire body, lateral view; B - head end, apical view; 
C - tail, lateral view; D - spicules and gubernaculum, lateral view. Scale bars in/<m. 



950 S. IVANOVA & C.VAUCHER 

Male. Body slender, tapering to both ends and slightly expanded in testis region. 
Cephalic hooks with blades 8 //m long and base 5 /im wide. Amphidial aperture 6x2 
piva. Oeasophageal corpus 9 piva wide at anterior, isthmus 8 piva wide, bulb 13 piva wide, 
32 piva long. Intestine collapsed. Testis reflexed in 298 pirn from anterior, flexure 95 piva 
long. Developing spermatocytes numerous, in 4, then 8 rows, spherical, 1.5-2 piva in 
diameter. Vas deferens separated from ejaculatory duct by constriction. Spermia 
spherical, 1-1.5 pirn in diameter. Spicular muscles and levator ani muscle prominent. 
Copulatory muscles present preanally. Bursa absent. Spicules paired, light brown in 
colour, cephalated, highly sclero tized, bent at one third length from distal part. Manu- 
bria elongated, 1 piva wide and 14 piva long, shafts distally 6 piva wide, then gradually 
tapering to tips which curved anteriad and bluntly rounded, 2 piva thick. Gubernaculum 
darker than spicules, massive, parallel to spicules, with crurae 13 piva wide embracing 
spicules and thick dorso-caudal apophysis 16/* m long. Four pairs of preanal papilliform 
ventral sensilla. Two (?) pairs of postanal lateral papilla around left-side fimbriate organ. 
Fimbriate organs asymmetrically disposed, large, circular, rised, with fibrous basal rim 
2 pi m thick, internal chamber covered by transparent membrane 11 piva and aperture 4 
piva va diameter. Filamentous portion of tail 46 pirn long and 3 piva thick. 

Female. Body stout, very slightly tapering posterior to vulva. Tail abruptly 
changes into bluntly rounded terminus 8 piva thick 98 pi m long. Cephalic hooks with 
blades \0 piva long and base 7 piva wide. Minute oral aperture 1 piva wide. Amphidial 
aperture oval, 4x2 piva, pouch 4 piva wide 5 piva long. Corpus 10 piva wide, isthmus 8 
pi m wide and bulb 14 ^m wide 37 piva long very slightly displaced dorsally. Cuticu- 
larized meandering channel 5 //m thick observed subventrally in vulva region running 
through gland with numerous nuclei (excretory gland?). Distal tip of ovary in 110 pi m 
posterior to vulva. Ovary runs back nearly to end of expanded part of tail, then returns 
to the same distance, then runs back to start and turns to anterior where reflexes in 408 
}im from oesophageal base and forms spermatheca. Spermatheca 95 piva long 35 piva 
wide packed with spherical spermatozoa resembled those from male testis. Thick- 
walled uterus with two smooth-shelled eggs 60 x 25 piva. Vagina straight, muscular, 38 
pirn long, with slightly protruded lips. Vulva post-median. Anus in 15 piva before right- 
side fimbriate organ. Fimbriate organs situated slightly asymmetrically near extremity 
of expanded part of tail (distance between centres of these 20 piva), circular, raised, with 
basal rim 7 piva thick 34 piva in diameter and aperture 23 piva in diameter covered with 
transparent membrane. Channel 4 pi m thick runs through fimbriate organs. 

Type host and locality. Yagansia spatulifera (Mich.) AF 4262, Chile, 
Villarica Vulcan, 1200 m, 14.02.01, coll. and identified by C. Czudi. 

Type habitat. Coelomic cavity at mid-body region. 

Type specimens. Holotype male MHNG 35447 and paratype female MHNG 
35448 deposited in Natural History Museum (Geneva, Switzerland). 

Differential diagnosis. The genus Ungella comprises 7 species: U. secta 
Cobb, 1928, U. astrida Spiridonov, 2001, U. burmensis Ivanova & Hope, 2000, 
U. ituriensis Spiridonov. 2001, U. kivuensis Spiridonov, 2001, U. mexicana Ivanova & 
Hope, 2000 and U. sucofera Timm, 1962. 

Ungella chileana sp. n. is characterised by having swollen posterior of females 
and filamentous portion of tail; not-embedded cephalic hooks with characteristic head 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 95 1 




Fig. 4 
Ungella chileana sp. n. Female paratype. A - entire body; B - head end; C - fimbriate organ; 
D - tail. All in lateral position. Scale bars in firn. 



952 S. IVANOVA & C. VAUCHER 

bone-like base and pointed blades; pocket-like amphids; oesophagus from slender 
corpus, no isthmus and narrow basal bulb not rounded at base; nerve ring around mid- 
oesophagus; excretory pore opposite anterior of bulb; long excretory duct; large cir- 
cular asymmetrically placed caudal fimbriate organs; paired highly sclerotised arcuate 
cephalated spicules and massive gubernaculum; 4 pairs of precloacal and 2 pairs post- 
cloacal male genital sensilla; extremity of ovary between vulva and anus; several loops 
of ovary; postmedian vulva; and by absence of cephalic sensilla. 

The present species differs from the rest of Ungella by largest spicules of 
different shape and circular fimbriate organs, shorter female tail, more posterior vulva 
position, ovary arrangement and female body shape which is swollen not slender in 
posterior. 

The present species resembles U. kivuensis, U. astrida and U. ituriensis in 
location of excretory pore opposite basal bulb and U. ituriensis as well in location of 
ovary tip cell but differs by having differently shaped surfactantly attached vs em- 
bedded cephalic hooks, differently shaped much longer spicules, expanded vs slender 
posterior of females, filamentous terminal portion of tail vs gradually pointing tail, 
post-median vs median vulva position, shorter and thinner oesophagus and fimbriate 
organs of different shape and size. From U. seda, U. burmensis, U. mexicana, U. suco- 
fera, U. micronychium n. sp. the present species differs in location of excretory pore 
which in former species is well behind the oesophageal base, by having differently 
shaped much longer spicules, expanded vs slender posterior of females. From U. sec- 
ta, U. burmensis, U. mexicana the present species differs also by having asym- 
metrically placed larger fimbriate organs vs small elliptical ones. From U. secta, U. 
burmensis, U. sucofera, U. kivuensis and U. astrida the present species is distinguished 
by location of ovary tip cell between vulva and anus vs in tail and from U. secta by 
absence vs presence of bursa. 

Etymology. The species name refers to locality of annelid host. 

Ungella micronychium sp. n. Fig. 5 

Measurements. Holotype male. L = 862 pim; D = 34 //m; Oes = 99 //m; Ex = 
124 //m; NR = 69 pm; Cd = 107 Jim; Sp (arc) = 51 pirn; Sp (chord) = 43 pirn; Gub = 
17; a = 25.35; b = 8.71; c = 8.06. 

Paratype male. L = 964 ptm; D = 35 ;^m; Oes = 92 }im\ Ex = 124 pirn; NR = 
62 }im; Cd = 106 //m; Sp (left, arc) = 47 pira; Sp (right, arc) = 40 ptm; Sp (left, chord) 
= 29 pirn; Sp (right, chord) = 32 //m; Gub = 17 pirn; a = 27.54; b = 10.48; c = 9.09. 

Description. Males. Body slender, slightly tapering to anterior end, tail curved, 
conoid, spicate. Cuticle thin, smooth. Epidermis well-developed, up to 10 pirn, thick. 
Head truncate, hooks triangle-shaped, 5 /im long 2 ^m thick, slightly embedded. 
Mouth shifted dorsad, oral aperture minute, stoma absent. No head sensilla. Amphids 
with oval apertures 3-5 x 1 .5 ;/m and pouch 5 x 4//m. Oesophagus from corpus 43-48 
ptm long and 6-7 pira wide at mid-length, with basal widening as wide as bulb, isthmus 
5 //m wide 23-30 ;<m long and bulb 9-12 pira and 17-21 ptm long. Bulb piriform with 
granular secretion of dorsal oesophageal gland. Nerve ring encircles anterior of 
isthmus. Excretory pore in 25 posterior to oesophageal bulb, 1 pira wide, disposed on 
hillock 1.5 piva high. Excretory duct thin, 20-22 pira long, leads to excretory cell 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 953 




Fig. 5 
Ungella micronychium sp. n. Male. A- paratype, entire body, lateral view; B - holotype, anterior 
end, lateral view; C - paratype, head end, subdorsal view; D - holotype, tail, lateral view; E - fim- 
briate organ, lateral view; F - paratype, spicules and gubernaculum, lateral view; G - holotype, 
spicules and gubernaculum, lateral view; H - holotype, gubernaculum, lateral view. Scale bars in 
pim. 



954 S. IVANOVA & C. VAUCHER 

extending till mid-body. Three large nuclei observable at anterior region of cell: one 
near testis flexure and two smaller near distal part of duct. Content of excretory cell in 
anterior region from well-defined granules and homogenous in posterior one. Testis 
reflexed at 260-320 jdm from oesophageal base. Flexure thin and 120-135 firn long. 
Developing spermatocytes in 2, then 3 rows, 3-4 x 4 ^m, then 6-7 x 7 pira in size. 
Spermatids amoeboid, 5-7 x 6-8 //m. Vas deferens posteriorly set off from ejaculatory 
duct. Ejaculatory duct from large cells. Spermia numerous, spherical 1 jim. in diameter. 
Spicules light brown in colour, strongly cuticularized, curved, not distinctively cepha- 
late. Holotype male spicules equal, thin, with nearly straight proximal and curved distal 
parts, bent at one third of its length from tips on nearly right angle. Manubria elon- 
gated, a little wider then shafts, which are 5 /im wide. Distal spicule tips pointed, 
curved upward and separated by cleft 5-6 /<m deep. Paratype spicules unequal; left 
spicule the same shape as holotype but manubrium more prominent and the right one 
shorter with wider shaft and no distinct manubrium. Dark massive gubernaculum with 
small crurae, lateral ridge and short lateral apophysis. Tail curved, terminal portion 
24-28 pirn long and 2 //m wide. Fimbriate organs prominent, asymmetrically placed, 
right one in 20-25 pim posterior to anus, left in 10-12 pim farther back. Fimbriate 
organs with bowl-shaped chamber 30 pim. long and 20 pim wide and circular aperture 
6-7 pim in diameter partially covered with transparent membrane surrounded by raised 
fibrous basal rim 7-8 pim thick. Single sensillum protruded from fimbriate organ aper- 
ture. A pair of sub ventral preanal papilliform sensilla, two pairs of subventral post- 
cloacal sensilla around posterior fimbriate organ and two pairs of lateral and subdorsal 
sensilla situated close to the end of conical portion of tail. 

Females. Not found. 

Type host and locality. Yagansia diversicolor Beddard, 965.171 , Chile, Prov. 
Valdivia, 25.10.65, coll. and identified by A. Zicsi. 

Type habitat. Coelomic cavity at anterior. 

Type specimens. Holotype male MHNG 35449 ENVE deposited in Natural 
History Museum (Geneva, Switzerland). 

Differential diagnosis. Ungella micronychium sp.n. is characterised by 
having truncate head bearing minute cephalic hooks, small pocket-like amphids, short 
oesophagus from corpus, isthmus and bulb, nerve ring on isthmus, excretory pore 
posterior to oesophageal bulb, arcuate not distinctively cephalate spicules with bifur- 
cate distal tips, dark massive gubernaculum, prominent asymmetricaly placed caudal 
fimbriate organs with bowl-shaped chambers, two pairs precloacal and four pairs post- 
cloacal genital sensilla. 

Present species is closest to U. mexicana Ivanova & Hope, 2000 by size of 
cephalic hooks, shape of oesophagus, tail, spicules, gubernaculum and excretory pore 
position. It can be distinguished from the former species by longer spicules, oeso- 
phagus and tail, less prominent amphids, absence of cephalic sensilla and much larger 
asymmetrically placed fimbriate organs. It differs from the rest of Ungella by having 
minute cephalic hooks, different shape of fimbriate organs in tail and spicules and 
gubernaculum. 

Etymology. The species name derived from Greek words «mikros» (small) and 
«onychium» (claw or talon) and reflects the size of cephalic hooks. It is a noun in 
apposition. 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 955 

Patagoniella gen. n. 

Diagnosis. Rhabditida, Drilonematoidea, Ungellidae. Blade-like cephalic 
hooks with amalgamated base present. Amphids with thickened rim and no pouch. 
Stoma short; displaced dorsally, esophagus nearly cylindrical. Nerve ring situated 
around anterior of bulb. Excretory pore situated closely to head end. Paired fimbriate 
organs in caudal region asymmetrically placed, with large subcuticular chamber, small 
external aperture and thick, basal rim. Males monorchic; spicules paired and curved; 
small boat-like gubernaculum with dorso-caudal apophysis. Bursa present. Female 
prodelphic, monodelphic, spermatheca set off; vulva at anterior third of body length 
with oblique vagina. No uterine sack present. 

Type and only species. Patagoniella capitoporus sp. n. 

Etymology. Generic name refers to geographic region where the annelid host 
was found. 

Patagoniella capitoporus sp. n. Figs 6-7 

Measurements. Holotype female . L = 1494 ^m; D (max) = 40 pm; D (behind 
V) = 26 pm; NR = 95 pm; Oes = 122 pm; Ex = 6 pm; Cd = 325 pm; V% = 36.81; 
Ovum = 60 x 26 pm, a = 37.35; b = 12.25; c = 4.6. 

Paratype females (n = 4). L= 1592±170.9 (1451-1831) /mi D (max) = 42.5±2.9 
(40-45) pm; D (behind V) = 25±3.6 (22-30) pm; NR = 89.25±8.7 (79-100) pm; Oes = 
117.8+7.9 (107-126) pm; Ex = 6.5+1 (5-7) pm; Cd = 325±26.3 (302-360) pm; V% = 
35.1 (34.29-36.27); Ova = 53.25±5.4 (49-61) pm x 22.5±3.3 (18-25) pm; a = 
37.69±5.8 (32.24-45.78); b = 13.56+1.5 (11.52-15.26); c = 4.91±0.5 (4.44-5.55). 

Paratype males (n = 7). L = 1424.43±214.1 (1234-1813) pm; D = 29±3.6 (22- 
34)#m; NR = 86.57±6.5 {15-95) pm; Oes = 112+6.1 (102-119) pm; Ex = 5.57±0.8 (5- 
l)pm; Cd= 158.14+10.4 (144-172) pm; Sp (arc) = 33.57±4.7 (29-42) pm; Sp (chord) 
= 27±3.8 (22-34) pm; Gub = 8.86±2 (6-12) pm; a = 49.98±10.6 (64.75-36.29); b = 
10.94±4.6 (1.5-16.94); c = 9.03±1.4 (7.69-11.55). 

Juveniles (n = 3). L = 550±94.3 (479-657) pm; D = 19+5.3 (15-25) pm; NR = 
63.33±2.9 (60-65) ^m; Oes = 104.3±5.5 (98-108) pm; Ex = A pm; Cd = 98±43.7 (57- 
144) pm; a = 29.57±4.2 (26.28-34.3); b = 5.31+1.2 (4.44-6.7); c = 6.16±2.0 (4.56-8.4) 

Description. Adults. Body cylindrical, long, slender, tapering to tail. Head 
bluntly rounded or truncate, inclined dorsally. Cuticle 1-2 p m thick, annulations 1- 
1.5 pm visible in tail. Epidermal layer from 1 pm in anterior end to 5 pm in tail. 
Cephalic hooks slightly embedded into head tissue. Base of hooks amalgamated with 
processes inclined. Dorsal hook blades diverged, longer than base. No head sensilla 
visible. Mouth and stoma slightly displaced dorsally. Amphidial apertures elliptical, 
with thin cuticularized rim, situated just behind hooks base and slightly displaced dor- 
sally. Amphidial pouch not observed. Excretory pore in 5.57-6.4 pm from anterior, flat- 
tened in dorso- ventral direction. Excretory duct 1.14-1.2 pm wide, proceeds up to 
35.8-41 .43 pm, then narrowing to less than 1 pm wide and runs until after oesophagus. 
Excretory cell large, with granular content and huge nucleus. Oesophagus muscular, 
slender, nearly cylindrical, with slightest expansion at posterior bulb. Isthmus less than 
1 pm thinner than corpus and about 1 .5-2 pm than bulb. Three nuclei of oesophageal 



956 



S. IVANOVA & C. VAUCHER 




Fig. 6 
PatagonieUa capitoporus gen. n., sp. n. Female. A - holotype, entire body, lateral view; B - 
paratype, head end, apical view; C - paratype, head end, lateral view; D - paratype, fimbriate or- 
gan, lateral view; E - paratype, part of oviduct, lateral view; F - paratype, spermatheca, oviduct 
and uterus, lateral view: G - paratype, tail, lateral view. Scale bars in//m. 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 957 

glands in posterior of bulb. Nerve ring wide, encircling anterior of bulb. Cardia large. 
Intestine collapsed, with walls composed from large cells and cuticularized lining. Pair 
of asymmetrical fimbriate organs in tail. 

Females. Body tapered dramatically just behind vulva. Lateral chord 6-9 
(7.4±1.1) pim wide at mid-body. Hook blades 4-6 (5.4±0.9) pim long, hook base 3-4 
(3.6±0.5) pim long. Stoma 5-6 (5.4±0.5) pim long and 1-2 (1 .8+0.4) pi m wide. Amphids 
5-4 (4.4±0.5) x 2 (2±0)pim, situated close to hook base. Excretory pore in 5-7 (6.4±0.9) 
pim from head end. Nerve ring about 10 pirn, thick. Anterior of corpus 8-10 (9.2±0.8) 
jim wide, corpus 8-9 (8.6±0.5) pim wide, isthmus 6-8 (7±1) pira wide, bulb 27-35 
(29.6±3.2) piva long and 10-11 (10.4±0.5) pirn wide. Genital tube starts in tail, runs on 
dorsal side and reflexed in 174-272 (229±40.4) pim from anterior, where forms long 
narrow set-off spermatheca 63-134 (91.4±26.2) pim long and 12-17 (14.6+1.9) pim 
wide filled with oval or spherical spermatozoa 1.5-4 x 2-4 pim in size. Oviduct from 
large cells, about 80-152 pim long. Single mature smooth-shelled egg 49-61 (54.6±5.6) 
x 18-26 (23.2±3.3) pim in uterus. Spermatozoa in uterus as in spermatheca. Egg shell 
1 pim thick. Vulva at the level of one-third length of the body. Lips protruded. Vagina 
muscular, obliquely inclined, 15-18 (17±1.2) pim long. No post-uterine suck present. 
Tail long, conical, initially as wide as before anus, then gradually pointing and ends in 
conoid portion 23-34 (28.8±4.1)//m long and 2-3 (2.6±0.5)pim wide. Fimbriate organs 
wider than tail diameter, cause slight swelling of tail. First fimbriate organ situated in 
80-95 (87.2±7) pim behind anus, the next one in 35-52 (43.8±7.1) pim farther back. 
Fimbriate organs slightly raised, longitudinally oval, with fibrous basal rim 29-36 
(31 ±2.8) x 17-24 (19. 6±2. 1 ) pt m , fringed internally with thin rim and deep chamber 17- 
24 (20.8±2.8) x 12-16 (14.2+1.6) pim, half-covered from surface by transparent mem- 
brane. In several specimens circular membrane aperture 13-14 pim in diameter 
observed. Bottom of organ chamber covered by fibers. Posterior fimbriate organ 
slightly smaller in several specimens. 

Males. Body length, anterior end, oesophagus and excretory system as in 
females with cephalic hooks little smaller in size than in females (blades 4.71 ±0.5 
(3-40) pim; base 3.43±0.8 (2-4) pirn). Amphids 4-7 (4.86+1.1) x 1-2 (1.71 ±0.8) pi m . 
Lateral chord 4.71±0.5 (4-5) pim. Cuticularized spot 1 pim in size situated opposite 
excretory pore on dorsal side observed in several specimens. Testis reflexed at 
271.71+31.2 (220-310) pim from anterior, testis flexure 177.57±46.2 (124-250) pim 
long. Developing spermatocytes hexagonal, arranged in 2, then 3 rows, 4x4, then 7 x 
7; immature sperm in testis with irregular outlines arranged in no rows, 4-7 in diameter, 
spermatids in vas deferens spherical, numerous, about 1-1.5 in diameter. Constriction 
between vas deferens and ejaculatory duct, spermatids in ejaculatory duct 2.5-3.5 in 
diameter. Tail conical with short filamentous terminal portion 15-22 (18.29±2.6) pim 
long. Spicules not distinctively cephalate, sickle-shaped, with manubria 2.86±0.4 (2-3) 
x 2.71±0.5 (2-3) pim blades 2.14±0.4 (2-3) pim wide at mid-length and pointed distal 
tips. Gubernaculum minute, boat-shaped, with proximal hook 3.14±2.6 (15-22) pim 
long bent posteriad. Long thin ribbed bursa present. Four-five pairs of postanal sub- 
ventral sensilla. First one in 12-25 posteriorly to anus, next one in 11-13 farther back 
and situated more laterally, third in front of anterior fimbriate organ, fourth between 
fimbriate organs, last one just behind posterior fimbriate organ. Each sensillum with 



958 



S. IVANOVA & C. VAUCHER 



20 




Fig. 7 
Patagoniella capitoporus gen. n., sp. n. Male and juvenile. A- paratype male, entire body, lateral 
view; B, C - anterior end, paratype male, lateral view; D - head, paratype male, ventral view; 
E - male tail, lateral view; F - spicule s and gubernaculum, lateral view; G -juvenile anterior 
end, lateral view; H - entire juvenile, lateral view. Scale bars in//m. 



prominent base 1 .5 high and fine bristle 4-10 high. Fimbriate organs the same in struc- 
ture as in females but smaller in size, first one situated in 46.43±7.2 (36-60) //m 
posterior to anus and second one in 18±7.9 (7-31) ptm farther back. First fimbriate 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 959 

organ with basal rim 17±2.8 (15-23) x 12.71+2.5 (10-17) //m, nearly spherical internal 
chamber 8. 57±1.1 (8-11) x 8.1 4±1. 5 (6-11)/* m; second one with rim 14±2.2 (11-18) x 
11.20+1.4 (10-14) pm and chamber 7.86±0.4 (7-8) x 7.43+1.1 (5-8) pm. Circular 
aperture in membrane covered the organ 3-5 pm in diameter. 

Juveniles. Slender body, anterior end rounded, tail conical with filamentous 
terminal portion 15+6.6 (9-22) pm long. Cephalic hooks thin with blades 3.33±0.6 
(3-4) pm and base 1.33+0.6 (1-2) pm long. Amphids not observable. Excretory pore 
and duct Iunior less wide, duct visible until after oesophagus base. Oesophagus with 
corpus 5 pm and isthmus 4 pm and basal bulb 8±1 (7-9) pm wide. Cardia and intes- 
tine well-developed. Genital primordium in 231+18.2 (210-242) pm from anterior, 
reflexed, 98.3±53.9 (60-160) pm long. Fimbriate organs circular, first situated in 
14.67±6.7 (9-22) pm from anus and second one in 13.33+5 (8-18) jim.. Both fimbriate 
organs with basal rim 8.67+1 .5 (7-10) pm. in diameter and chamber 5±1 (4-6) pm in 
diameter. Aperture in covering membrane not observable. 

Type host and locality. Yagansia papillosa AF 653, Chile, prov. Bahia Pond, 
Patagonic forest, 6.10.89, collected and identified by Mr Covarubias. 

Type habitat. Coelomic cavity. 

Type specimens. Holotype female MHNG 35450 INVE, three paratype females 
and seven paratype males MHNG 35451 INVE and two juveniles MHNG 35452 INVE 
deposited in Natural History Museum (Geneva, Switzerland). 

Differential diagnosis. The species resembles most members of Ungella 
Cobb, 1928 in general morphology but differs from all of them in having of excretory 
pore closely to anterior end and more anterior position of vulva (at one third body 
length y s median or post-median position). From Plesioungella Yeates et al., 1998, it 
differs by presence of paired spicules of different shape and slender posterior of 
females. From Onychonema Baylis, 1943, it can be distinguished by presence of 
fimbriate organs. From Acanthungella (Ivanova & Hope, 2004), it differs by absence 
of somatic sensillae along the body. 

Etymology. Species name derived from Latin words caput (head) and porus 
(pore) and reflects the anteriormost position of excretory pore. It is a noun in apposition. 

KEY TO GENERA OF UNGELLIDAE WITH SPICULAR APPARATUS 

1 Somatic sensilla present Acanthungella Ivanova & Hope, 2004 

Somatic sensilla absent 2 

2 Single spicule Plesioungella Yeates, Spiridonov & Blakemore, 1998 

Paired spicules 3 

3 Caudal fimbriate organs («suckers») absent Onychonema Baylis, 1943 

Caudal fimbriate organs present 4 

4 Excretory pore near head end Patagoniella gen. n. 

Excretory pore at the level of mid-oesophagus or well behind the oeso- 
phageal base 5 

5 Extremity of ovary behind vulva which is median or post-equatorial 

Ungella Cobb, 1928 

Extremity of ovary behind oesophageal base, vulva in front of anus 
Yagansiella gen. n. 



960 S. IVANOVA & C. VAUCHER 



TAXONOMICAL REMARKS 

Drilonematid fauna of Neotropic acanthodrilids is the less (if at all) studied 
amongst all Drilonematoidea. Till now only Ungella mexicana was described from 
Howascolex sp. from Mexico. Two new Ungella species and two new genera were 
described above. Both Ungella species demonstrate similarity in shape of oesophagus, 
spicules and gubernaculum and presence of prominent fimbriate organs. From type 
species U. secta Cobb, 1928 they differ by structure of cephalic hooks attached to head 
surface whereas in U. secta they have deeply embedded shafts. Excretory pore position 
seems to be variable in the genus and takes place or opposite nerve ring (U. kivuensis 
Spiridonov, 2001, U. astrida Spiridonov, 2001, U. ituriensis Spiridonov, 2001, U. 
chileana sp. n.) either behind oesophageal bulb (U. secta, U. burmensis Ivanova et 
Hope, 2000, U. mexicana, U. sucofera Timm, 1962, U. micronychium sp. n.) whereas 
nerve ring position on isthmus is constant. Oesophagus shape, presence of few (or 
more often single) thin-shelled eggs and median/post-median vulva position and struc- 
ture of excretory system is as well constant. Typically, ovary is leading straight to 
anterior end though in U. chileana it makes some loops at posterior. Short post-uterine 
sack present (U. kivuensis, U. astrida, U. ituriensis, U. sucofera) or absent. For male 
reproductive system, morphologically distinct vas deferens and ejaculatory duct, 
spherical small-sized immature sperm, curved paired spicules, sometimes bifurcated 
(U. mexicana, U. micronychium) are characteristic. Bursa present (U. secta) or absent 
(the rest of species). Fimbriate organs vary greatly in shape, size and position. 

Yagansiella longicollis gen. n., sp. n. possess some unusual features. Cuticular 
ridges on its anterior end reminds of cephalic armature of Homungellidae though these 
not joined with hooks which is single in the latter. Anterior placement of female 
gonoduct extremity in Yagansiella is unusual among drilonematids, as well as an- 
teriormost position of excretory pore in Patagoniella. Appearance of Yagansiella 
longicollis resembles in some way of oxyurids inhabiting earthworm gut - for instance, 
Posterovulva danieli Spiridonov, Ivanova, 1998 - in body shape, prominent annu- 
lation, spacious pseudocoel, muscular vagina though the present species undoubtedly 
belongs to Ungellidae. 

ACKNOWLEDGEMENTS 

The first author gratefully acknowledges the financial support related with re- 
search project provided by the Natural History Museum of Genève, Switzerland and 
thanks for the loan of nematode specimens. The study was also partially supported by 
RFBR grant 05-04-48140 and "Bioresources" programme. 

REFERENCES 

Cobb, N. A. 1928. Ungella secta n. gen., n. sp., a nemic parasite of the Burmese Oligochaete 

(earthworm) Eutyphoeus rarus. Journal of the Washington Academy of Sciences. 

Contribution to the Science of Nematology 18: 394-397. 
Baylis, H. A. 1943. Some nematode parasite of earthworms from the Indo-Malay region. 

Parasitology 35 (3):112-127. 
Ivanova, E. S. & Hope, W. D. 2000. Two new species of Ungella Cobb, 1928 (Rhabditida: 

Drilonematoidea) from earthworms deposited in the Smithsonian Institution Natural 

History Museum. Russian Journal of Nematology 8 (1): 57-64. 



NEMATODE PARASITES FROM COELOMIC CAVITY OF NEOTROPIC ACANTHRODILIDS 961 



Ivanova, E. S. & Hope, W. D. 2004. Two new nematode genera (Rhabditida: Drilonematoidea) 
with somatic sensory organs from earthworms deposited in the Smithsonian Institution's 
Natural History Museum. Nematology 6(3): 423-436. 

Seinhorst, J. W. 1959. A rapid method for the transfer of nematodes from fixative to anhydrous 
glycerin. Nematologica 4: 54-60. 

Spirtdonov, S. E. 2001. Parasitic organisms from tropical earthworms deposited in the Royal 
Museum of Central Africa. Annales des Sciences Zoologiques, Musée Royal de l'Afrique 
Centrale, Tervuren, Belgique 285: 19-27. 

Spirtdonov, S. E. & Ivanova, E. S. 1998. Parasitic nematodes of tropical megascolecid earth- 
worm Pheretima leucocirca from Ba Vi National Park in Viet Nam. Vestnik zoologii 32 
(1-2): 40-50. 

Timm, R. W. 1962. Nematode parasites of the coelomic cavity of earthworms. I. The genera 
Synoecnema and Ungella. Biologia (Dacca) 8: 1-7. 

Yeates, G. W., Spirtdonov, S. E. & Blakemore, R. 1998. Plesioungella kathleenae gen. et sp. 
n. (Nematoda: Drilonematoidea) from the Australian endemic megascolecid earthworm 
Fletcherodrilus unicus (Fletcher, 1889). New Zealand Journal of Zoology 25: 205-212. 



Revue suisse de Zoologie 112 (4): 963-982; décembre 2005 



New species of Leiodidae (Coleoptera) and new records from the 
Neotropical Region 

José Maria SALGADO COSTAS 

Dpto. de Biologia Animal, Facultad de Biologia, Universidad de Leon, 

E-24071 Leon, Espana. E-mail: dbajsc@unileon.es 

New species of Leiodidae (Coleoptera) and new records from the 
Neotropical region. - Four new species of Neotropical Leiodidae are de- 
scribed: Adelopsis Catarina sp. n., Adelopsis curvipes sp. n., Eucatops 
(Eucatops) troglodytes sp. n. and Parapaulipalpina giachinoi sp. n. New in- 
formation is given on certain basic diagnostic characters of several species, 
as well as further records on their distribution in the Neotropical Region. 

Keywords: Coleoptera - Leiodidae - Cholevinae - Camiarinae - taxonomy - 
Adelopsis Catarina sp. n. - Adelopsis curvipes sp. n. - Eucatops (Eucatops) 
troglodytes sp. n. - Parapaulipalpina giachinoi sp. n. 

INTRODUCTION 

This paper on the family Leiodidae, subfamilies Cholevinae and Camiarinae, is 
the continuation of research started by the author in 1991 for the fauna of Neotropical 
Region. The 11 publications which are of listed in the bibliography. 

Four new species belonging to the genera Adelopsis, Eucatops and Para- 
paulipalpina are described in this study. New data on the distribution of other species 
are given, as well as descriptions and illustrations of structures considered essential for 
differentiating the species but which had not been indicated in original descriptions or 
later papers, such as the male genital segment and spermathecal complex. 

MATERIAL AND METHODS 

A total of 477 specimens of the collection of the Muséum d'histoire naturelle, 
Geneva (Switzerland), from now on referred to as MHNG and of the private collection 
of Dr P. M. Giachino, Turin (Italy), from now on referred to as CPMG were studied. 

The types of Adelopsis insolitus Szymczakowski, 1961 and Adelopsis trian- 
gulifer Szymczakowski, 1961 deposited in the Swedish Museum of the Natural 
History, Stockholm (Sweden) were also examined. 

As all the studied material was dry, for the examination of many of the struc- 
tures the specimens had to be softened in boiling water for 10-15 minutes. Some of the 
examined pieces were placed in hot 10% KOH to separate adhérences. All the 
extracted structures were mounted on a small rectangular piece of plastic in a drop of 
"Hoyer liquid". Illustrations were by camara lucida. 



Manuscript accepted 15.03.2005 



964 J- M. SALGADO COSTAS 

SYSTEMATICS 
Cholevinae Kirby, 1837 

Ptomaphagini Jeannel, 1911 

Adelopsis Catarina sp. n. 

Type material. Holotype, S . Nova Teutônia, Santa Catarina State (Brazil), IV.1976, Fr. 
Plaumann leg. (MHNG). Paratypes: Nova Teutônia, Santa Catarina State (Brazil). 29 9, III- 
1976; 29 9, 1-1977; IS S, VI- 1977; lo\ IX-1978, Fr. Plaumann leg. (MHNG); IS (CPMG) 
and 1 S (CJSC), Zoology Department, University of Leon. 

Diagnosis. Length, 2.40-2.80 mm. Segments of antennal club of almost equal 
thickness; protarsi male dilated; genital segment with median region of spiculum 
gastrale wide and lobulate; aedeagus with right lobe arcuate and pointed in apical 
region, bearing numerous microsetae on right dorsal face of median lobe; spermathecal 
complex with numerous spirals forming arches in posterior region. 

Description. Male. Holotype: length, 2.80 mm; width, 1.30 mm. (Paratypes: 
length, 2.40-2.80 mm; width, 1.20-1.34 mm). Overall shape oval, fairly robust (Plate 
1: Fig. 1). Metathoracic wings fully developed. Reddish brown colour, with legs and 
base of antennae lighter. Pubescence very short, fine, golden and laid back. Transverse 
striolae weakly visible on head and clearly visible on pronotum and elytra. 

Head 1 .60 times narrower than pronotum; eyes developed. Antennae short, 3.20 
times shorter than body, not surpassing basal region of pronotum; last five segments 
almost of equal width; 2 nd to 6 th segments decreasing progressively in length; 6 th seg- 
ment transverse, the longest being the 1 st (Fig. 2). Using the length of the 9 th segment 
as a basis, relative length of each segment from 1 st to 11 th : 1.80-1.30-1.08-0.85-0.77- 
0.72-1 .05-0.57-1 .00-1 .00-1 .25. Proportions between the length and width of each club 
segment, from the 7 th to the 11 th : 1.28-0 .69-0 .96-0 .96-1. 26. 

Pronotum transverse, 1 .65 times wider than long, convex, with sides subparallel 
in basal half; posterior angles weakly protruding; as wide as anterior region of elytra. 
Elytra oval, weakly convex and narrowing slightly posteriorly, with scarcely rounded 
tip; transverse striae quite noticeable, as close together as those of pronotum and 
slightly oblique. Anterior tibiae robust in anterior region; protarsi dilated, first tar- 
somere 1.10 times narrower than the apical region of the tibiae (Fig. 3). 

Genital segment as long as wide, spiculum gastrale with median branch of 
anterior region very wide, bearing three lobes, one central and two lateral ones; the two 
lateral branches of the posterior region strongly arcuate (Fig. 4). 

Aedeagus wide, 1.65 times longer than wide and belonging to the ultraevolved 
group (Szymczakowski, 1961). In lateral view (Fig. 5), with very large, oval apical 
orifice, closed by the two strongly curved asymetrical lobes. Left lobe narrow and 
highly developed, with two apical setae; right lobe arcuate, much more developed than 
left one, narrowing to a pointed tip. Parameres fused with median lobe, left one 
differentiated in apical region bearing two inserted short setae very close together. 
Stylet or flagellum of internal sac very long and spiral-shaped, much more expanded 
in basal region. Right anterior median region of median lobe with numerous microsetae 
(Fig. 5). In ventral view, right lobe peak-shaped and flagellum long and wide in median 
region, forming folds, and end region filiform (Fig. 6). 



NEW NEOTROPICAL LEIODIDAE 



965 




Plate 1 
Fig. 1. Habitus of Adelopsis Catarina sp. n. (body length, 2.80 mm); Fig. 8. Habitus of Adelopsis 
curvipes sp. n. (body length, 2.75 mm); Fig. 29. Habitus of Parapaulipalpina giachinoi sp. n. 
(body length, 1.45 mm); Fig. 36. Habitus of Eue atop <s (Eucatops) troglodytes sp. n. (body length, 
3.05 mm). 



966 



J. M. SALGADO COSTAS 







Figs 2-7 
Adelopsis Catarina sp. n. 2. Antenna; 3. Protarsus male; 4. Genital segment, ventral view; 5. 
Aedeagus, lateral-left view; 6. Aedeagus, ventral view; 7. Spermathecal complex. (Scale bars: 
0.20 mm). 



Female. Generally smaller, though with the same external morphological 
characters as the male, except for the slender protarsi. 

Spermathecal complex wih the spermatheca around 3.5 times longer than wide, 
weakly expanded in anterior region and wider than apical lobe which is blunt at tip, 
narrowing in median region; highly complex posterior region, formed by numerous 
spires (9-10), posterior ones slightly wider and set out in archs (Fig. 7). 

Discussion. The most similar species to Adelopsis Catarina sp. n. is A. trian- 
gulifera Szymczakowski, 1961, captured in the same locality, and although the new 
taxon is larger, the body shape, striation, antennae and protarsi are the same. However, 



NEW NEOTROPICAL LEIODIDAE 967 

examination of the holotype reveals enormous differences between the two species in 
the spiculum gastrale of the genital segment and in the shape of the aedeagus, with the 
two parameres clearly differentiated and the large apical orifice in the new species. 

The structure of the aedeagus belongs to the ultrarevolved group species, 
including Adelopsis asper Jeannel, 1936, A. asperoides Szymczakowski, 1963, 
A. bernardi Portevin, 1923, A. grouvellei Jeannel, 1936, A. insolita Szymczakowski, 
1961, A. leo Gnaspini, 1993 and A. piruapuera Gnaspini, 1993, all with a large apical 
orifice in the aedeagus. However, A. Catarina shows some similarity in the shape of the 
lobules with A. insolita, though is noticeably different when compared with the holo- 
type of the latter species. Also, the general shape of the aedeagus is similar to 
A. bernardi and A. grouvellei, however the genitalia of these two species was examined 
in this study and the differences between A. Catarina are in the shape and direction of 
the parameres and of the left and right lobes, and also the lack of microsetae on the 
median lobe. In conclusion, besides the differentiating characters observed in the 
aedeagus with regard to the shape of the lobes and the number of setae, those observed 
in the spiculum gastrale of the genital segment and in the spermathecal complex should 
also be added as they make this new taxon unmistakeable. 

Etymology. The name is a noun in apposition referring to the Brazilian state 
where a great diversity of species belonging to the genus Adelopsis is found. 

Adelopsis curvipes sp. n. 

Type material. Holotype, 6 . Nova Teutônia, Santa Catarina State (Brazil), V.1977, 
Fr. Plaumann leg. (MHNG). 

Diagnosis. Length: 2.75 mm. Wide body. Apical club of antenna flattened, 
segments clearly transverse. Right lobe of aedeagus developed and pentagonal, with 
membranous protuberance in mid apical region. Left paramere with apical region 
curved outwards and bearing three setae, right paramere with five setae. 

Description. Male. Length: 2.75 mm; width: 1.45 mm Metathoracic wings 
fully developed. Slightly ovoid; wide and robust appearance; maximum width in 
anterior sixth of elytra (Plate 1: Fig. 8). Shiny dark brown body. Pubescence golden, 
short, fine and laid back. Transverse striolae visible on head, prothorax and elytra, with 
no other punctuation. 

Head 1.65 times narrower than pronotum; eyes developed. Antennae quite 
robust, surpassing posterior margin of pronotum and 3.10 times shorter than body; first 
five segments yellowish, the following ones darker in colour; 6 th segment slightly 
wider than long, 3 rd shorter than 2 nd ; antennal club with segments flattened and trans- 
verse, even 11 th , which ends in a whitish membranous structure (Fig. 9). Using the 
length of the 9 th segment as a basis, relative length of each segment from 1 st to 11 th : 
1.40-1.13-1.00-0.75-0.70-0.65-1.04-0.50-1.00-0.95-1.50. Similarly, the proportions 
between the length and width of each club segment, from 7 th to 11 th : 1.03-0.50-0.84- 
0.76-0.98. 

Pronotum transverse, 2.00 times wider than long, base scarcely narrower in 
anterior region of elytra; sides weakly blunt in anterior half, almost rectilinear in pos- 
terior half; posterior angles acute, but weakly pointed; transverse striolae quite fine and 
close together, slightly undulate, more marked than those on the head. Elytra elongate, 



968 J- M. SALGADO COSTAS 

weakly convex, with apical region widely arcuate, slightly truncate; 2.50 times pre- 
notai length, 1.25 times longer than wide; transverse striolae spaced further apart than 
on pronotum, weakly oblique, areas between them smooth. Anterior tibiae short and 
expanded in apical region; protarsi dilated, first tarsomere 1 .35 times narrower than 
maximum width of pro tibia (Fig. 10). Median tibiae slightly arcuate and posterior ones 
straight, both robust at end. 

Aedeagus, in lateral view (Fig. 11), slightly curved in ventral face, with very 
sy metrical tip, bearing three differentiated lobes set out at different levels, two bearing 
a short seta and the third with microsetae; right paramere slightly dilated before tip, 
pointed at tip, with 5 setae of unequal length inserted in apical region. In dorsal view 
(Fig. 12), apical orifice of median lobe displaced towards the left, and feebly 
developed; completely lacking left lobe; right lobe highly developed, forming trans- 
verse pentagonal plate, without visible setae, and with protruding, more membranous 
anterior median region. Parameres laterally joined along entire length of median lobe, 
except for the apical area of the left paramere which is curved outwards and surpasses 
the apical orifice of the median lobe, where three equidistant setae are inserted. Internal 
sac with robust curved stylet which appears to be resting on a rectangular sclerotized 
plate. 

Female not known. 

Discussion. In many species of the genus Adelopsis the apical area of the right 
lobe is triangular, rectangular or pentagonal, for example: Adelopsis bellatrix 
Szymczakowski, 1968; A. galea Gnaspini & Peck, 1996; A. heterocera Poitevin, 1907; 
A. onorei Salgado, 2002; A. palata Gnaspini & Peck, 2001; A. peruviensis Bias, 1980 
and A. santamarta Gnaspini & Peck, 2001 . Differences in the shape of the mid apical 
region of the membranous right lobe, the absence of a left lobe and the number and 
layout of the setae on the parameres make A. curvipes sp. n. easily distinguishable by 
the general structure of the aedeagus in all the previously-mentioned species. 

Greater similarity to A. brunnea Jeannel, 1936, from Colombia, and its different 
subspecies from Venezuela, all collected in caves (Szymczakowski, 1975) and to A. 
chapadaensis Salgado, 1999, from Mato Grosso, Brasil, is observed in the structure of 
the right lobe. Differences in external morphology between these two species and A. 
curvipes sp. n. are observed in the size, A. curvipes is larger and wider. The aedeagus 
of the new tax on is differentiated from A. brunnea and A. chapadaensis by the 
membranous protuberance in the mid apical area of the right lobe, and from A. cha- 
padaensis by the lack of setae on the right lobe in dorsal view; it is also differentiated 
from these two species by its length, shape, apical position and number of setae on the 
parameres. 

Etymology. The specific name refers to one of the diagnostic characters of the 
species, the apical curvature of the left paramere. 

Adelopsis ascutellaris (Murray, 1856) 

Material examined. 12 km N Luepa, Gran Sabana, Bolivar Province (Venezuela). 2$ ? , 
l/ll-VII-1987, S. & J. Peck leg. (MHNG). 

Description. Female. 2.00 mm long, with very weakly arcuate prenotai sides; 
antennal segments with 6 th transverse, 2 nd as long as the sum of 3 rd and 4 th segments 



NEW NEOTROPICAL LEIODIDAE 969 

together. The spermathecal complex is formed by an elongate oblong apical lobe, as 
wide as anterior region of spermatheca which is elongate and of similar width; the 
posterior region of the spermathecal complex is formed by 4 turns of equal thickness 
(Fig. 13). See Jeannel (1936) for the description of other structures. 

Distribution. The only record known to date is from Caracas D.F. (Venezuela) 
(Jeannel, 1936), the record from Bolivar Province is now made for the first time. 

Adelopsis asperoides Szymczakowski, 1963 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 1<?-19, IV- 1976; 
2<?<?-3 9 9,V-1976; lcM 9 , XII-1976; la, 1-1977; 1 9 , VI-1977, Fr. Plaumann leg. (MHNG 
andCPMG). 

Description. Length 2.15-2.40 mm. Male genital segment as long as wide, 
spiculum gastrale with median branch of anterior region around 2.5 times longer than 
wide and with parallel sides; lateral branches of posterior region long, narrow and 
slightly arcuate (Fig. 14). 

Spermathecal complex with spermatheca about four times longer than wide, of 
uniform thickness and weakly arcuate shaped; apical lobe ovoid and protruding; 
posterior region coiled with two-three turns (Fig. 15). 

For the description of other structures and also figures see Szymczakowski 
(1963) and Gnaspini (1993). 

Distribution. This species is known from Sao Paulo and Minas Gerais States 
(Brazil), (Peck et al., 1998). The record from Santa Catarina State is new for Brazil. 

Adelopsis bernardi Portevin, 1923 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 1<?-19, III- 1976; 
666, IY-1976; 26 6, V-1976; 46 6-22 9, XII-1976; 1Ó*-19, 1-1977; 366-19, V-1977; 
26 6-1 9 , VI-1977, Fr. Plaumann leg. (MHNG and CPMG). 

Arroyo Itabo, Guazu, Alto Parana Province (Paraguay). \6, 4-XI-1979, Expedition 
Muséum de Genève leg. (MHNG). 

Description. Size 2.20-2.50 mm. Male genital segment as long as wide; spi- 
culum gastrale with median brach of anterior region as wide as long, slightly angulose 
at tip; lateral branches of posterior region narrow, long and almost straight (Fig. 16). 

Spermathecal complex with spermatheca of uniform thickness, bearing an arch 
dorsally and four times longer than wide, with apical lobe and wider than anterior 
region of spermatheca; posterior region coiled with four turns (Fig. 17). 

See Jeannel (1936) and Szymczakowski (1963) for the description of other 
structures and illustrations. 

Distribution. Teresópolis, Rio de Janeiro State (Brazil). The record from Santa 
Catarina State is new for Brazil. 

Adelopsis grouvellei Jeannel, 1936 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). \6 , III-1976; 16- 
49 9, IV-1976; 26 6-1 9 , V-1976; 29 9, XII-1976; 36 6, 1-1977; 16-1 9 , V-1977; 36 6, VI- 
1977; 1 9 , IX-1978, Fr. Plaumann leg. (MHNG and CPMG). 

Description. Size 2.10-3.00 mm. Male genital segment somewhat wider than 
long, spiculum gastrale with median branch of the anterior region arcuate and wider 



970 



J. M. SALGADO COSTAS 





Figs 9-13 
Adelopsis curvipes sp. n. 9. Antenna; 10. Protarsus male; 11. Aedeagus, lateral-right view; 
12. Aedeagus, dorsal view. Adelopsis ascutellaris . 13. Spermathecal complex. (Scale bars: 
0.20 mm). 



NEW NEOTROPICAL LEIODIDAE 971 

than long; lateral branches of posterior region long and slightly curved upwards 
(Fig. 18). 

Spermathecal complex with arcuate spermatheca, wider in anterior region, 
apical lobe with anterior region blunt and wider than the anterior region of the 
spermatheca; posterior region formed by numerous superposed turns of uniform 
thickness (Fig. 19). 

See Jeannel (1936) and Szymczakowski (1963) for the description of other 
structures and also graphs. 

Distribution. Recorded from Bahia, Parana, Rio de Janeiro and Sao Paulo States 
(Brazil) by Szymczakowski (1963) and Peck et al. (1998). The record from Santa 
Catarina State is new for Brazil. 

Adelopsis insolita Szymczakowski, 1961 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 1 8 , IV- 1976; 1 5 , VI- 
1977, Fr. Plaumann leg. (MHNG). 

Description. Size 2.00-2.50 mm. The aedeagus observed in the holotype has a 
very large apical orifice; very long and strongly undulate stylet, and curved asym- 
metrical parameres. Spiculum gastrale with median region wider than long and no 
anterior branch is observed, unlike in the other species, the lateral branches of the 
posterior region are long, straight and narrow, though feebly dilated in apical region 
(Fig. 20). 

The spermathecal complex is not described as it is absent in the studied female 
specimen. 

See Szymczakowski (1961) for extensive description and for illustrations. 

Distribution. This species has only been recorded in Nova Teutônia, Santa 
Catarina State (Brazil). 

Adelopsis luculenta Szymczakowski, 1963 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 2c? S -22 9 , IV-1976; 
1 cM 9 , V-1976; 1 6 , XII-1976, Fr. Plaumann leg. (MHNG and CPMG). 

Description. Size 2.60-3.70 mm. Male genital segment wider than long, spicu- 
lum gastrale with median branch of anterior region wider than long, weakly angulose 
at the tip, the two lateral branches of the posterior region slightly curved (Fig. 21). This 
structure is similar in shape to A. bernardi, but the spiculum gastrale in this species is 
less robust than in A. luculenta (Fig. 16). 

Spermathecal complex with anterior region of spermatheca strongly expanded; 
apical lobe oval, weakly blunt at the tip; posterior region formed by two robust turns 
(Fig. 22). 

See Szymczakowski (1963) for extensive description and for illustrations. 

Distribution. Only the record for Sao Paulo State is known, its distribution has 
now been extended to Santa Catarina State. 

Paulipalpina claudicans (Szymczakowski, 1980) 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 112c? cî-129 ? 5 , 
captured in III- 1976, IV-1976, V-1976, XII-1976, 1-1977, V-1977, VI- 1977, Fr. Plaumann leg. 
(MHNG and CPMG). 



972 



J. M. SALGADO COSTAS 







Figs 14-17 
Adelopsis asperoides. 14. Genital segment, ventral view; 15. Spermathecal complex. Adelopsis 
bernardi. 16. Genital segment, ventral view; 17. Spermathecal complex. (Scale bars: 0.20 mm). 



Description. Size 2.00-2.30 mm. Male genital segment as long as wide, spi- 
culum gastrale narrow, long and straight (Fig. 23). 

Spermathecal complex with small lobe in mid anterior region of spermatheca in 
a widely arcuate shape which narrows progressively; posterior region formed by 
numerous superposed turns and ending in a structure resembling a flattened lobe 
(Fig. 24). 

See Szymczakowski (1980) for extensive description and for illustrations. 

Distribution. Only data from Nova Teutônia, Santa Catarina State 
(Szymczakowski, 1980) and from Rio de Janeiro State (Salgado, 1999) are known. 

Parapaulipalpina giachinoi sp. n. 

Type material. Holotype, la. Samiria, Loreto Province (Peru). 28-X/8-XI-1980, CI. 

Vaucherleg. (MHNG). 



NEW NEOTROPICAL LEIODIDAE 973 

Diagnosis. Length, 1.45 mm. Antennal club segments transverse, except for 
11 th . Aedeagus very long and narrow, with lateral margins slightly undulate. Lobes of 
apical region of aedeagus touching, both robust, especially the right one. Flagellum as 
long as median lobe. 

Description. Male. Length, 1.45 mm; width, 0.75 mm. Metathoracic wings 
fully developed. Body ovoid, quite elongate (Plate 1: Fig. 29). Shiny reddish colour, 
legs lighter in colour with whitish protarsi and pale yellow antennae, except for slightly 
darker 7 th to 10 th segments. Pubescence very short, fine golden and laid back. Eyes 
developed. Maxilliary palps with final segment very fine, pointed and around twice as 
short as penultimate. 

Antenna 1.30 times as long as pronotum, almost reaching base of pronotum 
(Fig. 25); 8 th , 9 th and 10 th club segments weakly transverse, 4 th also transverse; in 
lateral view, last segment ventrally concave, forming a small pointed lamina. Using the 
length of the 9 th segment as a basis, relative length of each segment from 1 st to 11 th : 
1.48-1.50-0.80-0.60-0.70-0.70-1.10-0.50-1.00-1.00-1.60. Proportions between length 
and width of each club segment, from 7 th to 11 th : 1.36-0.60-0.98-0.95-1.85. 

Pronotum strongly transverse, 2.25 times as wide as long, maximum width in 
posterior angles which are acute and protruding posteriorly; sides uniformly rounded, 
basal edge weakly sinuous; transverse striae clearly visible and quite close together. 
Elytra together 1.35 times as long as wide, the same width as the posterior region of 
the pronotum; sides weakly arcuate, apical edges wide, subtruncate; transverse striolae 
clearly defined and further apart than those on the pronotum. Anterior tibiae simple, 
about 3.5 times longer than wide, anterior tarsi dilated, 0.60 times as wide as maximum 
widh of tibia (Fig. 26). Metafemur bearing a small tubercle near the middle at the 
posterior margin (Fig. 27). 

Genital segment with lobes of sternite together as long as wide; spiculum 
gastrale very straight and long, as long as sternite (Fig. 28). 

Aedeagus very long and narrow, 5 times longer than wide. In dorsal view 
(Fig. 30), with sides slightly undulate; apical orifice oval, small completely closed by 
two lobes of median lobe which are unequal and short, the right one being more robust 
and blunt, with 4-5 small setae inserted; left lobe with truncate tip and one small seta. 
In lateral view (Fig. 31), on ventral side median lobe uniformly curved, dorsally with 
two curvatures, one anterior and concave and the other posterior, longer and convex; 
apex forming very characteristic point bearing four small setae and dorsal hump; 
internal sac enclosing heliocoidal membranous structure, and a long flagellum. Para- 
meres scarcely visible, joined to whole length of median lobe. Internal sac with stylet 
or flagellum very long and ondulate, as long as aedeagus (note: when extracting the 
holotype genitalia the end section of the less sclerotized stylet may have broken off); 
no other structures visible. 

Female unknown. 

Discussion. Parapaulipalpina giachinoi sp. n. is included in the genus 
Parapaulipalpina due to the short antennae, slightly elongate antennal segments, 
aedeagus somewhat S-shaped in dorsal view and the stylet very elongate. The only two 
species in the genus, Parapaulipalpina dentata Gnaspini, 1996, from Venezuela, and 
P.filicornis (Jeannel, 1936), from Colombia, P. giachinoi sp. n. are easily differentiated 



974 



J. M. SALGADO COSTAS 




Figs 18-22 
Adelopsis grouvellei. 18. Genital segment, ventral view; 19. Spermathecal complex. Adelopsis 
insolita. 20. Genital segment, ventral view. Adelopsis luculenta. 21. Genital segment, ventral 
view; 22. Spermathecal complex. (Scale bars: 0.20 mm). 

by: smaller body size, antennal club segments more transverse, aedeagus less curved 
in dorsal view and, in particular, the two lobes closing the apical orifice more dilated 
and rounded. 

This is the first record of the genus Parapaulipalpina in Peru. 

Etymology. This species is named after the entomologist Dr Pier Mauro 
Giachino, in recognition of his extensive research work in the study of the families 
Leiodidae and Carabidae. 



Ptomaphagus (Adelops) spelaeus (Bilimek, 1867) 

Material examined. Cacahuamilca Cave, Guerrero State (Mexico). 1 o*-l 9 , 16-IV-1981, 
Strinati leg. (MHNG). 



NEW NEOTROPICAL LEIODIDAE 



975 






Figs 23-28 
Paulipalpina claudicans. 23. Genital segment, ventral view; 24. Spermathecal complex. 
Parapaulipalpina giachinoi sp. n. 25. Antenna; 26. Protarsus male; 27. Femur, tibia and tarsi 
posterior; 28. Genital segment, ventral view. (Scale bars: 0.20 mm). 



Description. Size 3.00 mm. See Peck (1973) for extensive description and for 
illustrations. 

Distribution. Troglophil species recorded in Guerrero, Jalisco and Oaxaca 
States (Mexico), (Peck et al, 1998). 



EUCATOPINI Jeannel, 1921 

Eucatops (Eucatops) glabricollis (Reitter, 1884) 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 19, IV- 1976; lo*. 
V-1976; 36 6-3 9 9 , XII-1976; 26 6-1 9 , 11-1977; 1 6-1 9 , V-1977, Fr. Plaumann leg. (MHNG 
and CPMG). 



976 



J. M. SALGADO COSTAS 





34 





35 



Figs 30-35 
Parapaulipalpina giachinoi sp. n. 30. Aedeagus, ventral view; 31. Aedeagus, lateral-right view. 
Eucatops (E.) glabricollis. 32. Genital segment, ventral view; 33. Spermathecal complex. 
Eucatops (E.) troglodytes sp. n. 34. Antenna; 35. Protarsus male. (Scale bars: 0.20 mm). 

Description. Size 1.80-2.60 mm. Male genital segment subcylindrical, 1.90 
times as long as wide, with 8-10 setae in the inner ventral margin (Fig. 32). 

Spermathecal complex exhibiting a globose spermatheca, a very long and very 
fine spermathecal duct forming a jumble of turns and a weakly-defined membranous 
elongate final region (Fig. 33). 



NEW NEOTROPICAL LEIODIDAE 977 

See Jeannel (1936) for extensive description and for illustrations. 

Distribution. This species was mentioned by Blumenau, Santa Catarina State 
(Szymczakowski, 1963), Sao Paulo State (Jeannel, 1936) and Alagoas and Para States 
(Salgado, 1999). 

Eucatops (Eucatops) troglodytes sp. n. 

Type material. Holotype S , Grotte de Caiman, Route de Kaw, French Guiana, 2- VIII- 
1987, C. Juberthie leg. (CPMG). Paratypes: 5 â â , same data as holotype. (Collections: 1 6 , 
MHNG; 3 3S, CPMG and 1 Ö\ CJSC). 

Diagnosis. Length, 2.75-3.15 mm. Antennae with the 2 nd segment somewhat 
longer than the 3 rd ; 8 th to 10 th segments weakly transverse. Apical area of elytra not 
truncate. Protarsi very thin, none of the tarsomeres dilated. Aedeagus with triangular 
dorsal valve, longer than wide, ventral valve with strongly pointed tip, very narrow and 
protruding towards ventral face; parameres short, not surpassing tip of dorsal valve, 
setae quite long and inserted in apical area; stylet not distinguishable amongst sclero- 
tized pieces; basal lamina long, with ventral median carina and convex posterior 
margin. 

Description. Male. Holotype. Length, 3.05 mm; width, 1.80 mm (Paratypes: 
length 2.95-3.15 mm; width, 1.72-1.90 mm). Metathoracic wings fully developed. 
Strongly convex body (Plate 1: Fig. 36). Eyes developed. Very shiny reddish-brown 
colour. Pubescence very fine, laid back and yellowish. Punctuation on head somewhat 
stronger than on pronotum, fine in both cases, very abundant, with points clearly dis- 
tant and surface smooth. 

Antennae clearly longer than pronotum, 0.39 times length of body (Fig. 34); 
club segments progressively expanding towards the apical area, 8 th segment slightly 
transverse and 9 th and 10 th segments as long as wide; also, 4 th to 6 th segments of equal 
length, 2 nd being slightly longer than 3 rd and as long as 1 st using the length of the 9 th 
segment as a basis, relative length of each segment from 1 st to 11 th : 1.02-1.04-0.84- 
0.64-0.64-0.64-1.04-0.60-1.00-1.02-1.80. Proportions between length and width of 
each club segment from 7 th to 11 th : 1.30-0.76-1.00-0.98-1.54. 

Pronotum campaniform, strongly transverse, 2.25 times wider than long; later- 
al margins slightly sinuate in posterior region; basal region as wide an anterior region 
of elytra; posterior angles pointed. Maximum width of elytra in anterior fourth, 1.15 
times longer than wide; apical area of elytra not truncate, but slightly pointed and arch 
shaped; transverse striation clearly visible, defined by well defined points, striae sepa- 
rate and not in continuous curve. Legs long and slender. Protarsi very fine and long, 
none of the tarsomeres dilated, and much narrower than protibiae which are also 
slightly expanded (Fig. 35). Posterior femurs lacking spine or tubercle in posterior 
margin. Posterior tibiae not arcuate, with first metatarsomere shorter than following 
two together; internal metatibial spur slightly longer than first metatarsomere. 

Genital segment twice as long as wide, with 9-10 setae in internal margins of 
lobes of sternum anterior apical setae (Fig. 37), and 6 setae longer than sternal ones on 
margins of lobes of tergum (Fig. 38). 

Aedeagus with both valves of median lobe shaped very differently; dorsal valve 
longer than wide, triangular and larger than ventral one; ventral valve with undulate 
lateral margins, tip elongate and strongly pointed and sharp, also, in lateral view tip 



978 



J. M. SALGADO COSTAS 









Figs 37-43 
Eucatops (E.) troglodytes sp. n. 37. Genital segment, ventral view; 38. Genital segment, dorsal 
view; 39. Ventral valva of aedeagus; 40. Aedeagus, lateral view; 41. Left paramere, latero-dor- 
sal view; 42. Aedeagus and inner sac, dorsal view. Eunemadus chilensis. 43. Spermathecal com- 
plex. (Scale bars: 0.20 mm). 



separated from median lobe, slightly curved and protruding towards ventral (Figs. 39 
and 40). Parameres short and weakly robust, not surpassing tip of dorsal valve, bearing 
3 fairly long setae inserted in apical region (Fig. 41). Internal sac as long as dorsal 
valve, with numerous sclerotized pieces; stylet is not differentiated. Basal lamina 
longer than dorsal valve and clearly longer than wide, with strongly convex posterior 
margin; also exhibiting a weakly defined carina mid ventrally (Fig. 42). 
Female unknown. 



NEW NEOTROPICAL LEIODIDAE 979 

Variability. Differences have only been observed in the number of setae in the 
apical area of the parameres, as some paratypes have 4 or 5 setae, which are always 
long and all inserted in the apical area. The varying number of setae on the parameres 
appears to be frequent in species belonging to Eucatops (Salgado, 2003). 

Discussion. Eucatops (E.) troglodytes sp. n. resembles Eucatops (E.) oblongus 
Poitevin, 1903 , from Peru, in the number of setae inserted in the apical area of the para- 
meres, and Eucatops (E.)formicetorum (Bruch, 1918) from Argentina, in the elongate 
pointed shape of the valves and the short parameres which do not surpass the tip of the 
valves. However, the shape of the protarsi and many of the characters of the aedeagus, 
in particular the shape and position of the ventral valve, clearly separate this new taxon 
from any other species in the genus Eucatops. 

Ecology. To date, this is the first record of a Eucatops being captured inside a 
cave. Due to their developed eyes, large wings and pigmentation, the specimens of this 
species should be considered troglophil. They were found in the anterior part of the 
cave, feeding on organic material. 

Etymology. The specific name, an invariable epithet, refers to the tendency of 
the specimens to colonize caves. 

Anemadini Hatch, 1928 

Eunemadus chilensis Poitevin, 1914 

Material examined. Argentina: El Bolsón, 150 m altitude, Rio Negro Province; 1 9 , 7-1- 
1996, M. Etonti leg. (CPMG). Golondrinas, Chubut Province; 19, 15-IX-1981, Kovacs leg. 
(MHNG). 

Chile: Tolhuaca, Malleco Province; 29 9, 1-1987 (CPMG). 

Description. Female. Size: 2.90-3.60 mm. Very characteristic spermafhecal 
complex formed by sacciform spermatheca, lacking differentiated lobes, around five 
times longer than wide with quite clear superficial striation. Long, fine spermathecal 
duct inserted in basal region, approximately three times longer than spermatheca, 
which expands slightly at the end (Fig. 43). 

See Jeannel (1957, 1962) for extensive description and for illustrations. 

Distribution. The species has been recorded in Argentina, Rio Negro Province, 
and now in Chubut Province; and various provinces in Chile, such as Chiloé, 
Concepción, Llanquihue and Santiago. New for the Malleco Province. 

Dissochaetus hetschkoi Reitter, 1884 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 85c?c?-469 9 cap- 
tured in 1-1977, 11-1977 and V-1977, Fr. Plaumann leg. (MHNG and CPMG). 

Description. See Gnaspini (1991) and Salgado (1991a) for extensive descrip- 
tion and for illustrations. 

Distribution. This is a widely distributed species, known from Belize, Brazil, 
Mexico and Venezuela (Peck et al., 1998). 

Dissochaetus murrayi Reitter, 1884 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 49c?(?-63 9 9 cap- 
tured in 1-1977, 11-1977 y V-1977, Fr. Plaumann leg. (MHNG and CPMG). 

Description. See Gnaspini (1991) and Salgado (1991a) for extensive 
description and for illustrations. 



980 J- M. SALGADO COSTAS 

Distribution. Known from the south eastern Neotropical region, with records 
from Paso de los Libres, Corrientes Province (Argentina) (Salgado, 1991a) and several 
states in Brazil (Peck et al, 1998). 

Dissochaetus parallelus Poitevin, 1921 

Material examined. 400 m, 35 km N Altagracia, Miranda Province (Venezuela), 29 9, 
7-14-VI-1987, S. & J. Peck leg. (MHNG). 

Description. See Jeannel (1936) for extensive description and for illustrations. 
Distribution. To date, only known from the type locality, El Simon, Caracas 
D.F. (Venezuela). 

Dissochaetus villosus Szymczakowski, 1961 

Material examined. Nova Teutônia, Santa Catarina State (Brazil). 11-1977, 9c? o*-3 9 9 , 
Fr. Plaumann leg. (MHNG and CPMG). 

Description. See Szymczakowski (1961) and Gnaspini (1991) for extensive 
description and for illustrations. 

Distribution. This species is endemic in Brazil and known from Minas Gerais, 
Parana and Sao Paulo. New for the Santa Catharina State. 

Nemadiolus (Subnemadiolus) kuscheli Jeannel, 1962 

Material examined. Ushuaia, Monte Susana, Tierra de Fuego Province (Argentina). 1 ó* , 
26-111-1975, E. Horak leg. (MHNG). 

Description. See Jeannel (1962) and Salgado (1991b, 2000) for extensive de- 
scription and for illustrations. 

Distribution. Jeannel (1962) describes this species from specimens from Chepu, 
Chiloé Province. Salgado (1991b, 2000) recorded it from several provinces including 
Aisén, Cautfn, Chiloé, Concepción, Llanquihue, Magallanes, Malleco, Nuble, Osorno, 
Palena, Talea and Valdivia, all in Chile. The record from Tierra de Fuego Province 
extends its distribution area to Arsentina, from where it is recorded for the first time. 



Camiarinae Jeannel, 1911 

Agyrtodini Jeannel, 1936 

Chiliopelates kuscheli (Jeannel, 1957) 

Material examined. Tolhueca, Malleco Province (Chile). 3 cT cT-1 9 , 1-1987 (CPMG). 

Description. See Jeannel (1962) for extensive description and for illustrations. 

Distribution. This species is endemic in Chile. It had been recorded from Aisén 
and Llanquihue provinces (Jeannel, 1957) and is now recorded for the first time from 
Malleco province. 

Dasypelates nebulosus (Jeannel, 1957) 

Material examined. Tolhueca, Malleco Province (Chile), lo*, 1-1987 (CPMG). 

Description. See Jeannel (1957, 1962) for extensive description and for illus- 
trations. 



NEW NEOTROPICAL LEIODIDAE 



Distribution. This species is endemic in Chile and is only known from Bio Bio 
Province. The record from Malleco province is now also included. 

Eupelates transversestrigosus (Fairmaire & Germain, 1859) 

Material examined. Golondrinas, Chubut province, 2??, 15-IX-1981, Kovacs leg. 
(MHNG). 

Description. See Jeannel (1957 and 1962) for extensive description and for 
illustrations. 

Distribution. The species is known from various localities in the southern 
Neotropical region, Chubut and Rio Negro Provinces (Argentina) and Aconcagua, 
Arauco, Cautin, Chiloé, Concepción, Llanquihue, Nubie, Santiago and Valdivia 
Provinces (Chile) (Jeannel, 1962; Salgado, 1991b; Peck et al, 1998). 

ACKNOWLEDGEMENTS 

I would like to express my gratitude to the Natural History Museum, Geneva 
(Switzerland) for allowing me to study his entomological material and in particular to 
my friend Dr Pier Mauro Giachino, for making this material available to me. Also, the 
Swedish Museum of Natural History, Stockholm and its curator, Dr B. Viklund, for 
sending the types of Adelopsis triangulifer and A. insolita. 

REFERENCES 

Gnaspini, P. 1991. Brazilian Cholevidae (Coleoptera), with emphasis on cavernicolous species. 

I. Genus Dissochaetus. Giornale italiano di Entomologia 5: 325-340. 

Gnaspini, P. 1993. Brazilian Cholevidae (Coleoptera), with emphasis on cavernicolous species. 

II. Subgenus Iutururuca nov., subgenus of Adelopsis. Annales de la Société Entomo- 
logique de France (N.S.) 29 (1): 77-87. 

Gnaspini, P. 1994. The genus Eucatops (Coleoptera, Cholevidae, Eucatopinae). Description of 
new species and systematic remarks. IHERINGIA, Série Zoologie, Porto Alegre, 76: 
33-42. 

Jeannel, R. 1936. Monographie des Catopidae. Mémoires du Muséum National d'Histoire 
Naturelle (n. s.) 1: 1-433. 

Jeannel, R. 1957. Sur quelques Catopides, Liodides et Camiarides du Chili (Coleoptera). 
Revista Chilena de Entomologia 5: 41-65. 

Jeannel, R. 1962. Les Silphidae, Liodidae, Camiaridae et Catopidae de la Paléantarctide occi- 
dentale (pp. 481-525). In: Delamare, C. & Rapoport, E. (eds). Biologie de l'Amérique 
Australe. Vol. 1 , Etudes sur la Faune du Sol - Centre National de la Recherche Scien- 
tifique, Paris: 1-657. 

Peck, S.B. 1973. A review of the cavernicolous Catopinae (Coleoptera: Leiodidae) of Mexico, 
Belize and Guatemala. Bulletin of the Association for Mexican Cave Studies 5: 97-106. 

Peck, S.B., Gnaspini, P. & Newton, A.F. 1998. Catalogue and generic keys the Leiodidae of 
Mexico, West Indies and Central and South America (Insecta: Coleoptera). Giornale 
italiano di Entomologia 9: 37-72. 

Salgado, J.M. 1991a. Nota sobre algunos Dissochaetus (Coleoptera, Catopidae) de Brasil y 
Argentina. Bulletin et Annales de la Société royale belge d'Entomologie 127: 211-215. 

Salgado, J.M. 1991b. Nuevos datos sobre Cholevidae y Camiaridae (Coleoptera) de Chile. 
Elytron 5: 169-179. 

Salgado, J.M. 1999. The Leiodidae (Coleoptera) of the Carnegie Museum of Natural History. 
New data and description of two new species. Pan-Pacific Entomologist 75 (1): 35-47. 



982 J- M. SALGADO COSTAS 



Salgado, J.M. 2000. Revision of the genus Nemadiolus Jeannel, 1936 (Coleoptera: Leiodidae: 

Cholevinae). Elytron 14: 159-174. 
Salgado, J.M. 2001 . Nuevos datos sobre algunos Dissochaetus Reitter, 1885 de Ecuador, con la 

descripción de una nue va especie (Coleoptera, Leiodidae, Cholevinae). Nouvelle revue 

d'Entomologie (N.S.) 18 (3): 249-258. 
Salgado, J.M. 2002. Data on the genus Adelopsis from Ecuador. Description of five new species 

(Coleoptera Leiodidae Cholevinae Ptomaphagini). Belgian Journal of Entomology 4: 

113-128. 
Salgado, J.M. 2002. Revision of the genera Nemadiopsis and Falkonemadus (Coleoptera: 

Leiodidae: Cholevinae). European Journal of Entomology 99: 505-521. 
Salgado, J.M. 2003. El gènero Eucatops en Ecuador. Descripción de dos especies nuevas 

(Coleoptera, Leiodidae, Cholevinae, Eucatopini). Nouvelle revue d'Entomologie (N.S.) 

20(1): 51-60. 
Salgado, J.M. 2004. El gènero Eucatops en Ecuador, nuevas especies y nuevos datos 

(Coleoptera: Leiodidae: Cholevinae: Eucatopini). Boletin de la Sociedad Entomològica 

Aragonesa, Zaragoza 35: 1-17. 
Salgado, J.M. (in press). Napocatops subgen. nov. Eucatops (Napocatops) giganteus sp. nov. 

of Ecuador (Coleoptera: Leiodidae: Cholevinae: Eucatopini). Nouvelle revue d'Entomo- 
logie (N.S.). 
Salgado, J.M. (in press). Description of new genera and species of the subtribe Eunemadina 

from Chile. New data on other genera (Coleoptera: Leiodidae: Cholevinae). Deutsche 

Entomologische Zeitschrift. 
SzYMCZAKOWSKi, W. 1961. Espèces néotropicales nouvelles ou peu connues de la famille 

Catopidae (Coleoptera). Polskie Pismo Entomologiczne 31 (14): 139-163. 
Szymczakowski, W. 1963. Catopidae (Coleoptera) récoltés au Brésil par J. Mrâz. Acta Entomo- 
logica Musei nationalis Pragae 35: 667-680. 
Szymczakowski, W. 1975. Formes cavernicoles d' 'Adelopsis brunneus Jeann. du Venezuela et 

d'Ile de Trinidad (Coleoptera, Catopidae). Boletin de la Sociedad Venezolana de 

Espeleologia 6 (11): 13-24. 
Szymczakowski, W. 1980. Deux espèces nouvelles de Ptomaphaginae (Coleoptera, Catopidae). 

Polskie Pismo Entomologiczne 50 (4): 515-521. 



Revue suisse de Zoologie 112 (4): 983-1028; décembre 2005 



Fruit flies (Diptera, Tephritidae) from Saudi Arabia, with 
descriptions of a new genus and six new species 

Bernhard MERZ 1 & Hassan A. DAWAH 2 

1 Muséum d'histoire naturelle, C. P. 6434, CH-1211 Genève 6, Switzerland. 
E-mail : bernhard .merz @ mhn .ville-ge .ch 

2 King Khalid University, College of Science, Department of Biology, 
P. O. Box 9004, Abha, Saudi Arabia. 

E-mail: dawahhassan@hotmail.com 

Fruit flies (Diptera, Tephritidae) from Saudi Arabia, with descriptions 
of a new genus and six new species. - During a survey of the insect fauna 
of the Abha area (Aseer province, southwestern part of Saudi Arabia) with 
Malaise traps 51 species of Tephritidae were recorded, 40 of them being 
new records for this country. Nine species are not named. The first compre- 
hensive checklist for Saudi Arabia is presented, including 62 species of 
which 10 species are considered to be pests. Ar abode sis gen. n., with its 
type A. reductiseta sp. n., is described from Saudi Arabia and Yemen. 
Further new species are Neoceratitis flavoscutellata sp. n. (Saudi Arabia), 
Dicheniotes multipunctatus sp. n. (Kenya, Saudi Arabia, Tanzania, Yemen), 
Sphenella setosa sp. n. (Saudi Arabia), Tanaica maculata sp. n. (Ethiopia, 
Kenya, Saudi Arabia), and Tanaica pollinosa sp. n. (Saudi Arabia, South 
Africa). The status of Tephritomyia despoliata Hering and Trupanea aucta 
var. repleta Bezzi is discussed. Sphenella marginata austrina Munro is 
synonymized with S. marginata s. str. (syn. n.). Keys for the genera of the 
Campiglossa genus group and for the species of Tanaica Munro are pro- 
vided. 

Keywords: Diptera - Tephritidae - Saudi Arabia - new genus - new species - 
new synonymy. 

INTRODUCTION 

Saudi Arabia is a large country of about 2.15 Million km 2 . Its biogeographical 
position is quite peculiar as a crossroad of various realms, included in the Palaearctic 
Region which lies primarily in the North (at the border with Jordan), adjacent to the 
Afrotropical Region in the South West (separated from Eritrea by the narrow Red Sea), 
and to a lesser extent to the Oriental Region in the South East. It may be expected 
therefore that its fauna is reflecting this special biogeographical position, and elements 
of the various realms may be found. 

Very little information is available on the Tephritidae fauna of this country 
although species of this family are usually attractive with patterned wings and bright 
eyes, easy to collect, often abundant on their host plants, and economically and ecolo- 



Manuscript accepted 01.06.2005 



984 B. MERZ & H.A. DAWAH 

gically important. About 100 species of Tephritidae out of about 4500 described 
species worldwide are known as pests (White & Elson-Harris, 1992). The only 
comprehensive list of Saudi Arabian Tephritidae containing 12 species was given by 
Al-Ahmadi & Salem (1999), most of them species which cause damage in fruit plan- 
tations. Freidberg & Kugler (1989) added 6 species. Four further species are mentio- 
ned in isolated publications. Thus, 22 species are known up to date from Saudi Arabia 
(Tab. 1). Compared with the diversity of other countries (Tab. 2) it may be anticipated 
that any research which is not intended to study exclusively pest species will add new 
species to this list. 

In the course of a survey of the insect fauna of the Aseer (= ' Asir ) area (south- 
western part of Saudi Arabia) an important number of Tephritidae was collected by the 
second author using Malaise traps. After identification of the specimens by the first 
author it became clear that this collection improves considerably the knowledge on 
Tephritidae of Saudi Arabia. This paper brings together our present knowledge on its 
fruit fly diversity. In addition to the results of the identifications all available literature 
about Tephritidae of Saudi Arabia is summarized. 

MATERIAL AND METHODS 

Some 440 specimens of Tephritidae were collected using Malaise traps at 4 si- 
tes in South-Western Saudi Arabia by the second author. All sites belong to Aseer 
(='Asir) and are situated between 260 and 2200 m within a radius of about 100 km 
around the town of Abha. The names of the localities and further information are given 
in Tab. 3. Abha has the highest rainfall of Saudi Arabia and it is the only region where 
open forests may grow. Consequently, the region is comparatively heavily populated 
and most of its surface is cultivated. 

In order to resolve some taxonomic questions, about 80 specimens from other 
countries are included in this study. 

The depository of the specimens is as follows: 

CHD collection H. Dawah 

FMNH Finnish Museum of Natural History, Helsinki, Finland 

KMMA Koninklijk Museum voor Midden Afrika, Tervuren, Belgium 

MCSNM Museo Civico di Storia Naturale, Milano, Italy 

NHML Natural History Museum London (= British Museum of Natural History), 

England 

MHNG Muséum d'histoire naturelle, Genève, Switzerland 

NMWC National Museum of Wales, Cardiff, United Kingdom 

SAMCT South African Museum, Cape Town, South Africa 

SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany 

TAU Tel Aviv University, Israel 

ZMUA Zoological Museum, University of Amsterdam, Netherlands 

Most specimens collected during this study are deposited in the MHNG. Dupli- 
cates are basically stored in the NMWC, TAU and the CHD except where otherwise 
mentioned. 



FRUIT FLIES FROM SAUDI ARABIA 



985 



Tab. 1 
Checklist of Tephritidae of Saudi Arabia. Classification according to Norrbom et al. (1999a, b), 
but species are listed in alphabetical order within the tribe Tephritini. Abbreviations. Column 
«Source»: 1 = Al-Ahmadi & Salem (1999); 2 = Freidberg & Kugler (1989); 3 = Norrbom et al. 
(1999b); 4 = Shalaby (1961); 5 = White & Elson-Harris (1992); 6 = Anonymous (2002); 7 = De 
Meyer & Freidberg (2005). * = Identification doubtful as species often misidentified (see also 
text). Column «PS»: Pest status according to White & Elson-Harris (1992). B = Potential spe- 
cies for Biological Control of weed; M = Minor pest species or potential pest species, attacking 
commercially grown plants; P = Pest species; B, M = Either polyphagous species whose host 
range include commercially grown species (and therefore considered potential or minor pest) as 
well as weeds (potential useful species), or a species which is attacking a plant species which is 
cultivated in some parts of the world and considered to be a pest in others. 



no Name 



Source 



PS 



Subfamily Trypetinae 
Tribe Carpomyini 
Subtribe Carpomyina 

1 Carpomya incompleta (Becker, 1903) 

2 Myiopardalis pardalina (Bigot, 1891) 

Tribe Dacini 
Subtribe Ceratitidina 

3 Capparimyia aenigma De Meyer & Freidberg, 2005 

4 Ceratitis capitata (Wiedemann, 1824) 

5 Neoceratitis efflatouni (Hendel, 1931) 

6 Neoceratitis flavoscutellata sp. n. 

Subtribe Dacina 

7 Bactrocera (Zeugodacus) Cucurbitae (Coquillett, 1899) 

8 Bactrocera (Daculus) oleae (Gmelin, 1790) 

9 Bactrocera (s. str.) zonata (Saunders, 1842) 

10 Dacus (Leptoxyda) annulatus Becker, 1903 

11 Dacus (Didacus) sp. nr. arcuatus Munro, 1939 

12 Dacus (Didacus) ciliatus Loew, 1862 

13 Dacus (Didacus) frontalis Becker, 1922 

14 Dacus (Leptoxyda) longistylus Wiedemann, 1830 

15 Dacus (Leptoxyda) semisphaereus Becker, 1903 

16 Dacus (Didacus) vertebratus Bezzi, 1908 

17 Dacus (Leptoxyda) obesus Munro, 1948 

18 Dacus sp. 1 

Subfamily Tephritinae 
Tribe Noeetini 

19 Ensina sonchi (Linnaeus, 1767) 

Tribe Schistopterini 

20 Bactropota sp. 1 

21 Rhochmopterum arcoides Munro, 1935 

22 Schistopterum moebiusi Becker, 1903 

Tribe Tephrellini 
Subtribe Platensinina 

23 Psednometopum cf. aldabrense (Lamb, 1914) 

24 Stephanotrypeta brevicosta Hendel, 1931 

25 Stephanotrypeta vittata Freidberg, 1979 

Subtribe Tephrellina 

26 Dicheniotes angulicornis (Hendel, 1931) 

27 Dicheniotes multipunctatus sp. n. 



1 , present study 

1 



7, present study 
1,4,5 

present study 
present study 

1 

1 

6, present study 

3 

present study 

1,4 

1, 5, present study 

1 , present study 

present study 

5, present study 

present study 

present study 



present study 

present study 
present study 
present study 



present study 
present study 
3 , present study 

present study 
present study 



P 
P 
P 
M 

P 
P 
B,M 

P 

M 



B,M 



986 



B. MERZ & H. A. DAWAH 



no Name 



Source 



PS 



28 Gymnaciura austeni (Munro, 1935) 

29 Metasphenisca hazelae (Munro, 1947) 

30 Metasphenisca negeviana (Freidberg, 1974) 

31 Oxyaciura tibialis (Robineau-Desvoidy, 1830) 

32 Paraspheniscoides binarius (Loew, 1861) 

33 Paraspheniscus debskii (Efflatoun, 1924) 

Tribe Tephritini 

34 Acanthiophilus helianthi (Rossi, 1794) 

35 Arabodesis reductiseta sp. n. 

36 Campiglossa ignobilis (Loew, 1861) 

37 Capitites augur (Frauenfeld, 1857) 

38 Dectodesis auguralis (Bezzi, 1908) 

39 Dectodesis sp. 1 

40 Desmella sp. nr myiopitoides (Bezzi, 1908) 

41 Dioxyna sororcula (Wiedemann, 1830) 

42 Euare stella iphionae (Efflatoun, 1924) 

43 Freidbergia mirabilis Merz, 1999 

44 Goniurellia persignata Freidberg, 1980 

45 Goniurellia spinifera Freidberg, 1980 

46 Goniurellia tridens (Hendel, 1910) 

47 Hyalotephritis complanata (Munro, 1929) 

48 Hyalotephritis planiscutellata (Becker, 1903) 

49 Spathulina acroleuca (Schiner, 1868) 

50 Sphenella marginata (Fallen, 1814) 

51 Sphenella setosa sp. n. 

52 Tanaica maculata sp. n. 

53 Tanaica poi linosa sp. n. 

54 Telaletes ochraceus (Loew, 1861) 

55 Tephritomyia despoliata (Hering, 1956) 

56 Trupanea amoena (Frauenfeld, 1857) 

57 Trupanea pseudoamoena Freidberg, 1974 

58 Trupanea pulcherrima (Efflatoun, 1924) 

59 Trupanea repleta Bezzi, 1918 

60 Trupanea stellata (Fuesslin, 1775) 

61 Trupanea sp. 1 

62 Trupanodesis sp. 1 



present study 
present study 
2, present study 
present study 
present study 
present study 

1 P 
present study 

present study B 

1*, 3*, present study 

present study 

present study 

present study 

present study BJV1 

2 (from «Arabia») 
present study 
present study 

2, present study 

2 

present study 

present study 

present study M 

present study B 

present study 

present study 

present study 

present study 

present study 

1 , present study M 

2, present study 
2 

present study 

1 M 

present study 

present study 



The species are arranged according to Norrbom et al. (1999a); the nomenclatu- 
re follows Norrbom et al. (1999b), except for the Tephritis genus group, where the sys- 
tematic arrangement of Merz (1999) is adopted. 

Morphological terminology is according to White et al. (1999), except for the 
term postpedicel, which is used instead of 1st flagellomere (Stuckenberg, 1999). 

Each species is treated in the same way. First, the material studied is recorded. 
The localities are abbreviated as numbers following the details given in Tab. 3. In the 
«Distribution» section the general distribution is given. This information is taken basi- 
cally from Norrbom et al. (1999b) and occasionally from other sources. The host plants 
are listed from the literature given in brackets. It should be noted that other plants may 
be hosts in Saudi Arabia. The indications about the pest status of each species are 
mainly taken from White & Elson-Harris (1992). Finally, comments about taxonomy, 
nomenclature and biology are given in the «Remarks» section. 



FRUIT FLIES FROM SAUDI ARABIA 



987 



Tab. 2 
Tephritidae diversity in various countries of the Near and Middle East. 



Country 



Nr species 



Source 



Afghanistan 

Egypt 

Iran 

Iraq 

Israel 

Jordan 

Lebanon 

Saudi Arabia 

Syria 

Yemen 



46 
60 
23 
31 
85 
19 
18 
62 
34 
51 



Hering (1961) 

Norrbom et al. (1999b) 

Hering (1956) 

Korneyev & Dirlbek (2000) 

Freidberg (1988) 

Korneyev & Dirlbek (2000) 

Knio et al. (2002) 

present study 

Korneyev & Dirlbek (2000) 

Merz et al. (in press) 



Tab. 3 



Collecting sites in Saudi Arabia, Province of Aseer (= 'Asir). 



Nr Locality 



Coordinates 



Altitude coll. Dates 



Remarks 



1 Abha, Farm Centre 


18.13N/42.30E 


2200 


III.-VI.2001 


25 km E of Abha 


(close to locality 2) 










2 Madenate Al-Ameer 


18.13N/42.30E 


2200 


25 .11.-25 .V.2002 


25 km E of Abha 


Sultan 










3 Wadi Hali 


18.36N/41.18E 


810 


9.1.2003 


80 km NW Abha 


4 Maraba 


17.54N/42.23E 


260 


1.-30.V.2004 


60 km S Abha 


RESULTS 











Subfamily TRYPETLNAE 
Tribe Carpomyini 
Subtribe Carpomyina 

Carpomya incompleta (Becker, 1903) 

Material. 2: 11 do*, 59 2. 

Distribution. Italy, East Africa, Near and Middle East. First recorded from 
Saudi Arabia by Al-Ahmadi & Salem (1999). 

Host plants. Zizyphus spp. (Rhamnaceae) (Freidberg & Kugler, 1989). 

Pest status. Pest species in some parts of the world (White & Elson-Harris, 
1992). 

Tribe Dacini 
Subtribe Ceratitidina 

Capparimyia aenigma De Meyer & Freidberg, 2005 

Material. 2: 2$ $ (identification by M. De Meyer). 

Distribution. Tropical Africa and Arabian Peninsula. First recorded from Saudi 
Arabia by De Meyer & Freidberg (2005). 

Host plants. Maerua spp. and Boscia spp. (Capparidaceae) (De Meyer & 
Freidberg, 2005). 

Pest status. Not a pest species. 



988 B. MERZ & H. A. DAWAH 

Remarks. This species was recently described from specimens from Kenya and 
Tanzania (De Meyer & Freidberg, 2005). The two specimens from Saudi Arabia were 
also studied but not included into the type series because they differ slightly in thora- 
cic markings from the East African population. The identification is therefore tentative 
(De Meyer & Freidberg, 2005). 

Neoceratitis ejflatouni (Hendel, 1931) 

Material. 2: 1 S . 

Distribution. Originally described from Sudan, subsequently found in Egypt 
and Israel. New for Saudi Arabia. 

Host plant. Lycium schweinfurthii (Solanaceae) (Freidberg & Kugler, 1989). 

Pest status. Not a pest species. 

Neoceratitis flavoscutellata sp. n. Figs 1-10 

Material. Holotype 6 : SAUDI ARABIA: Abha, Madenate Ameer Sultan, 22.11.- 
25.V.2002, H. A. Dawah (MHNG). Paratypes: 16c? S, 20 9 9 , same data as holotype 
(CHD, KMMA, MHNG, NMWC, TAU). The holotype is double-mounted on a minutien 
pin on a polyporus block and is in good condition with only the left midleg missing. 

Etymology. The species is named after the entirely yellow scutellum, which is a 
unique characteristic within the genus. 

Diagnosis. This is the only species of Neoceratitis with an entirely yellow 
scutellum dorsally (Fig. 3); wing with discal, anterior and posterior apical band fused 
in rl and r2+3, subapical band isolated (Fig. 2). 

Description. Weng length. Male: 3.3-4.1 mm; female: 3.65-4.35 mm. 

Head (Fig. 1). Anterior half of frons, ventral three quarters of face, and most of 
gena whitish-yellow; posterior half of frons and dorsum of occiput brownish-yellow; 
ocellar triangle, dorsal quarter of face and irregular pattern on occiput dorsal to occi- 
pital foramen blackish; gena with indistinct brown to black spot at eye margin; anten- 
na and moufhparts yellow, but postpedicel with faint black tinge. Head in profile al- 
most 1 .5 times as high as long; gena about 1/6 to 1/5 as high as compound eye, the lat- 
ter elongate oval; frons slightly longer than wide, in dorsal view slightly more than half 
as wide as entire head width; frons black setulose in middle; face not projecting at 
vibrissal corner, with indistinct antennal grooves separated by a broad, evenly convex 
carina; palpus slightly flattened, with black, stiff setulae; proboscis rather fleshy, capi- 
tate; antenna little shorter than face, postpedicel about twice as long as wide, apico-dor- 
sally indistinctly pointed; arista setulose with longest rays about as long as basal 
diameter of arista. Chaetotaxy as in other Neoceratitis, all setae black: 2 frontal, 2 
orbital (posterior about 0.75 times as long as anterior), 1 strong ocellar, 1 postocellar, 
1 medial vertical, 1 lateral vertical, 1 genal, 1 row of postocular setae. 

Thorax. Mesonotum dorsally with a pattern as in Fig. 3; postpronotum entire- 
ly whitish-yellow; a narrow yellow stripe from wing base along transverse suture; dor- 
sal surface medially grey microtrichose, laterally with a shining, black, broad border; 
grey pattern antero medially pointed and posteriorly with 1-3 black teeth penetrating in- 
to grey area; a pair of shiny black spots just posterior of transverse suture; scutellum 
swollen, whitish-yellow with a narrow brown to black spot laterally at base, white 



FRUIT FLIES FROM SAUDI ARABIA 




Figs 1-3 
Neoceratitis flavo saltellata sp. n. (paratype from Abha): 1, head, lateral view; 2, wing; 3, tho- 
rax, dorsal view. 



setulose on dorsal surface; most of anepisternum, anatergite and katatergite whitish- 
yellow, remaining parts of pleuron brown to black, subshining; setulae on grey and 
whitish-yellow surfaces white, other setulae pale brown; prosternum with pale brown 
setulae. Chaetotaxy as in other Neoceratitis , all setae black: 2 scapular, 1 postpronotal, 
2 notopleural, 1 presuturai supra- alar, 1 postsutural supra-alar, 1 intra- alar, 1 postalar, 
1 dorsocentral posterior to line of postsutural supra-alar, 1 acrostichal, 1 anepisternal, 
1 anepimeral, 1 katepisternal, 2 (pairs of) scutellar setae. 

Legs. All femora (except for knees) brown, distal parts contrastingly yellow; 
mid- and hindfemur anteroventrally with 4-7, midfemur dorsoventrally with 4-5 short, 
black setulae at apex; midtibia with a black apicoventral seta. 

Wing (Fig. 2). Rl dorsally entirely setulose, ventrally with 4-6 setulae in distal 
half of pterostigma; node of R2+3 and R4+5 ventrally with 2-3 setulae; R4+5 dorsal- 
ly setulose over level of DM-Cu, the latter oblique; distance between R-M and DM-Cu 
on M about 1.2 times the length of DM-Cu; cell bcu with conspicuous posterodistal 
lobe; anal lobe large, undulating; wing with brown to black pattern, without light 
yellow areas; arrangement of spots and bands as in Fig. 2: Base of wing largely brown, 
in particular be and surface ventral to stem vein; numerous brown streaks and spots 
from c to bcu; distally with 4 crossbands, the discal, anterior and posterior apical bands 
fused in rl and r2+3; the subapical band well isolated from remaining pattern, 
obliquely trans versing DM-Cu; rl with 2-3 darker teeth-like spots; wing along C very 



990 B. MERZ & H.A. DAWAH 

narrowly hyaline. Halter and calypteres whitish-yellow; both calypteres convex, 
subequal. 

Abdomen male. Bicoloured, with posterior margin of tergite 1 , lateral spots on 
tergite 2, entire tergite 3 and anterior third of tergite 4 dark brown, other parts of 
tergites 1-4 whitish-yellow; tergite 5 yellow medially, becoming gradually darker 
brown laterally; tergites black setulose, except for whitish-yellow parts on tergites 1-4, 
which are white setulose. Terminalia (Figs 4-6): epandrium ovoid, fused with lateral 
surstyli, the latter bifurcated and black distally, with a slightly pointed, apically 
directed, sparsely setulose branch and a broad ventral plate; medial surstylus narrow, 
with two black, subequal prensisetae; cercus large, setulose; hypandrium and asso- 
ciated structures symmetrical, glans as in Fig. 6 (drawn from compressed glans), of 
complicated 3-dimensional structure; vesica spoon-shaped, very large, bordered by a 
broad fringe of minute tubercles; its articulation with remaining glans flexible; sub- 
apical lobe well developed, with a sclerotized tip and a bar-like sclerotization basally; 
acrophallus composed of two ducts, the more straight one weakly sclerotized, sub- 
apically surrounded by a strongly sclerotized sheath. 

Abdomen female. As in male, but entire anterior third of tergite 2 brown; 
tergite 6 almost invisible in dorsal view, at most 1/6 as wide as tergite 5; oviscape 
(Fig. 7) swollen, shining orange brown, but tip dark brown, black setulose; as long as 
2-3 preceding tergites combined; aculeus (Figs 8-9) evenly pointed; sternite 8 width / 
aculeus length ratio = 0.8; aculeus length: 1.2 mm; 2 spermathecae present (Fig. 10): 
composed of a spherical apical part with a reticulate surface and a cylindrical base with 
some rather sharp projections. 

Distribution. Saudi Arabia. 

Host plants. Unknown. All specimens were collected in a Malaise trap. 

Pest status. Not a pest species. 

Remarks. The genus was diagnosed under the name Trirhithromyia Hendel, 
1931, by Hardy (1967) and Freidberg & Kugler (1989). Korneyev (1994) synonymi- 
zed Trirhithromyia with Neoceratitis Hendel, 1927, and redescribed the genus. The 
new species fits well the characteristics given in these papers. The colour of the scutel- 
lum, however, which has black spots on the dorsal surface in other species of the ge- 
nus, was considered to be of generic value by Hardy (1967) and Korneyev (1994), but 
N . flavoscutellata has only a narrow brown spot ventrobasally. Shape of head, wing 
pattern and genitalic characters show, however, that this species is congeneric with 
other Neoceratitis. 

Within Neoceratitis, the new species appears to be related to N. cyanescens 
(Bezzi, 1923) which is known from Madagascar, Mauritius and La Réunion, sharing 
the same wing pattern and the entirely yellow postpronotal lobes. The grey-microtri- 
chose pattern on the mesonotum and the colour of the abdomen are very different in 
the two species, in addition to the coloration of the scutellum (3 couples from La 
Réunion, St. Pierre (MHNG) studied, see Fig. 6 in Hancock, 1984). 

Male terminalia of the new species differ clearly from those of /V. efflatouni in 
length of vesica and sclerotization of glans (see Freidberg & Kugler, 1989, Fig. 216). 
On the other hand, they are similar to Ceratitis capitata (Wiedemann, 1824), as 
illustrated by Freidberg & Kugler (1989, Fig. 193). Unfortunately, male terminalia are 
unknown for most species of the Ceratitidina, thus, their value for phylogeny or species 
recognition is limited. 



FRUIT FLIES FROM SAUDI ARABIA 



991 




Figs 4-10 
Neoceratitis flavoscutellata sp. n. (4-6, male; 7-10, female, paratypes from Abha): 4, epandrium. 
lateral view; 5, same, caudal view; 6, glans; 7, oviscape; 8, aculeus; 9, tip of aculeus; 10, sper- 
mathecae. 



Subtribe Dacena 

Bactrocera (s. str.) zonata (Saunders, 1842) 

Material. 4: la (identification by I. M. White); 19, same, but IX.-X.2004 
(identification by J. C. Deeming) (NMWC). 

Distribution. Originally an Oriental species which was introduced to various 
countries of the Near East (Egypt, Israel, United Arab Emirates, Oman, Iran, Yemen) 
and some islands in the Indian Ocean (Réunion, Mauritius). Recorded from Saudi 
Arabia anonymously (Anonymus, 2002). 

Host plants. Reared from numerous plants belonging to different families 
(White & Elson-Harris, 1992). 



992 B. MERZ & H. A. DAWAH 

Pest status. Pest species in some parts of the world (White & Elson-Harris, 
1992). 

Dacus (Didacus) sp. nr. arcuatus Munro, 1939 

Material. 2:2â S (identifications by I. M. White). 

Distribution. Published records of D. arcuatus from Kenya and South Africa. 
New for the Arabian Peninsula. 

Host plant. D. arcuatus was reared from Pergularia sp. (Asclepiadaceae) 
(Munro, 1984). 

Remarks. According to White (pers. comm.) the two males fit rather well D. ar- 
cuatus, but differ in morphological details. At the present moment only these two mal- 
es are available from the Arabian Peninsula, and a safe identification is not possible. 

Dacus (Didacus) frontalis Becker, 1922 

Material. 2: 3 c? 6 , 1 9 ; 4: 19 (identifications by I. M. White). 

Distribution. Entire Afrotropical Region including the Arabian Peninsula. 
Recorded from Saudi Arabia by White & Elson-Harris (1992) and Al-Ahmadi & Salem 
(1999). 

Host plants. Various Cucurbitaceae (White & Elson-Harris, 1992). 

Pest status. Pest species in some parts of the world (White & Elson-Harris, 
1992). 

Dacus (Didacus) vertebratus Bezzi. 1908 

Material. 2: 1<J; 4: 36 S , 29 9 (identifications by I. M. White) (MHNG, 
NMWC). 

Distribution . Entire Afrotropical Region including Madagascar, and the Arabian 
Peninsula. Recorded from Saudi Arabia by Munro (1984) and White & Elson-Harris 
(1992). 

Host plants . Various Cucurbitaceae (White & Elson-Harris, 1992). 

Pest status. Pest species in some parts of the world (White & Elson-Harris, 
1992). 

Dacus (Leptoxyda) longistylus Wiedemann, 1830 

Material. 4: 1 9 (identification by I. M. White). 

Distribution. Afrotropical Region. Arabian Peninsula (Munro, 1984). Reported 
from Saudi Arabia by Al-Ahmadi & Salem (1999) and Carroll et al. (2002). 

Host plant. Reared from Sodom apple (Calotropis procera, Asclepiadaceae) 
(Munro, 1984). 

Pest status. A potential pest species in areas where the host is cultivated. On the 
other hand. Sodom apple is poisonous to livestock, and D. longistylus may therefore be 
useful for Biological Control in other parts of its distribution range (White & Elson- 
Harris, 1992). 

Dacus (Leptoxyda) obesus Munro, 1948 

Material. 4: 1 â (identification by I. M. White). 

Distribution. So far only known from Kenya. New for Saudi Arabia. 



FRUIT FLIES FROM SAUDI ARABIA 993 

Host plant. Reared from Sodom apple {Calotropis procera, Asclepiadaceae) 
(Munro, 1984). 

Pest status. A potential pest species in areas where the host is cultivated (White 
& Elson-Harris, 1992). 

Remarks. This is one of the most remarkable species oi Dacus, characterized by 
a number of unusual features. Males lack a pecten on tergite 3; other characters of this 
small species are the blue-grey, silvery microtrichose scutum, the compact body, the 
broad continuous black band along anterior wing margin which is covering both costal 
cells, the absence of an anal streak, the presence of 2 black spots on the face, the ab- 
sence of both supra-alar and prescutellar setae, the entirely yellow scutellum, the ab- 
sence of stripe-like medial and lateral postsutural vittae (but with a yellow transversal 
marking in posterior half of scutum), the presence of two postalar vittae across katater- 
gite and anatergite, and the entirely yellow legs. A comprehensive description and use- 
ful illustrations were provided by Munro (1984, p. 151). Because of the unusual set of 
characters Munro (1984) erected the new tribe Pionodacini and the new genus 
Pionodacus Munro, 1984, for this species. This proposal, however, was not accepted 
by subsequent workers (Norrbom et al., 1999a). 

Dacus (Leptoxyda) semisphaereus Becker, 1903 

Material. 4: 1 9 (identification by I. M. White) (NMWC). 

Distribution. Eastern Africa from Egypt to Sudan and Ethiopia. New for Saudi 
Arabia. 

Host plant. Unknown. 

Pest status. Not a pest species. 

Remarks. The data concerning this species were communicated by J. C. 
Deeming (in litt., 8.1.2005). 

Dacus (Leptoxyda) sp. 1 

Material. 4: Id, 1 2 (identifications by I. M. White). 

Remarks. This species is externally similar to D. obesus, differing chiefly in the 
colour of the scutum, which is black posterior of the suture (with a transversal yellow 
marking on posterior half of scutum in D. obesus). Currently, a revision of the genus 
is in preparation by I. M. White and the species will be formally described in this work. 

Subfamily TEPHRITINAE 
Tribe Noeetini 

Ensina sonchi (Linnaeus, 1767) 

Material. 2: 1 $ , 2 2 ? . 

Distribution. Large distribution in Palaearctic, Afrotropical, Australasian 
(Hawaii) and Oriental Regions. New for Saudi Arabia. 

Host plants. Wide range of Asteraceae of the subfamily Cichorioideae 
(Chondrilla, Crépis, Helminthia, Lactuca, Picris, Scorzonera, Sonchus and others) 
(Freidberg & Kugler, 1989; Hendel, 1927; Merz, 1994). 

Pest status. A potential or minor pest on some cultivated plants (Scorzonera in 
Ukraine). On the other hand E. sonchi may be important in the natural regulation of 
Sonchus oleraceus, which is a weed in some parts of the world (White & Elson-Harris, 
1992). 



994 B. MERZ & H.A. DAWAH 

Tribe Schistopterini 

Bactropota sp. 1 

Material. 3: 45 specimens. 

Remarks. This species differs readily from B. woodi Bezzi, the only described 
species of the genus (Freidberg, 2002), in wing pattern. In contrast to B. woodi the 
anterior margin has ill-defined, small yellow-brown areas, thus the entire wing appears 
much darker without two hyaline incisions which are present in B. woodi (see Fig. 2 A 
in Freidberg, 2002). A revision of the genus is currently prepared by Freidberg et al. 
and the species will be formally described in that study. 

Rhochmopterum arcoides Munro, 1935 

Material. 2: 1 S . 

Distribution. East Africa. New for Saudi Arabia. 

Host plant. Reared in South Africa from flower heads of Vernonia kraussii 
(Asteraceae) (Munro, 1935). 

Pest status. Not a pest species. 

Remarks. This genus is in bad need of revision. At least 10 undescribed species 
are known in addition to the 6 described species from the Afrotropical Region 
(Freidberg, 2002). The specimen at hand fits very well the description and illustration 
of Munro (1935) as well as one couple from Kenya (Magadi road, 3. III. 1993, leg. 
B. Merz, MHNG). The only difference concerns the colour of the tibiae, which are 
yellow with two black rings in basal and distal third in the specimens studied from 
Saudi Arabia and Kenya, whereas Munro (1935) did not give any indication of their 
colour. 

Schistopterum moebiusi Becker, 1903 

Material. 2: 1 c? , 4$ ? ; 3: 1 o\ 1 9 . 

Distribution. Eastern Africa north to Lebanon (Deeming, in litt.), Yemen (Merz 
et al., in press). New for Saudi Arabia. 

Host plant. Pluchea dioscoridis (Asteraceae) (Freidberg & Kugler, 1989). 

Pest status. Not a pest species. 

Tribe Tephrellini 

Subtribe Platensintna 

Psednometopum cf. aldabrense (Lamb, 1914) 

Material. 2: 28 8 . 

Distribution. This species is only known from the island of Aldabra 
(Seychelles). The record from Kenya (Merz et al., in press) is based on a misidenti- 
fication of P. nigritum Munro, 1937. New for Saudi Arabia. 

Host plants. Unknown. 

Pest status. Not a pest species. 

Remarks. P. aldabrense was recently recorded from Yemen based on one male 
(Merz et al., in press), and this specimen belongs to the same species as the two males 
from Saudi Arabia. D. L. Hancock studied recently the types and additional specimens 
of Tephritis aldabrensis Lamb from Aldabra (in NHML) and found that the specimens 



FRUIT FLIES FROM SAUDI ARABIA 995 

from the Arabian Peninsula differ from those of Aldabra by the entirely black ptero- 
stigma (with hyaline spot in specimens from Aldabra), and he concluded that they may 
belong to an undescribed species (Hancock, in litt.). However, as a very limited 
number of specimens is available and the two populations differ only slightly, the 
intraspecific variability of the species in the genus is poorly known. We prefer therefo- 
re not to describe a new species. 

Stephanotrypeta brevicosta Hendel, 1931 

Material. 2: 26 S . 

Distribution. Kenya, Sudan. New for Saudi Arabia. 

Host plants. Unknown. 

P est status. Not a pest species. 

Stephanotrypeta vittata Freidberg, 1979 

Material. 2: 1 o\ 3 9 9 . 

Distribution. East Africa, Madagascar, Arabian Peninsula. Recorded from Saudi 
Arabia by Freidberg (1979). 

Host plant. Unknown. 

Pest status. Not a pest species. 

Remarks. We follow here Freidberg (1979) for the generic placement of the 
species and do not accept the proposition of Hancock (2003), who transferred this 
species to Terpnodesma Munro. 

Subtribe Tephrellina 

Dicheniotes angulicornis (Hendel, 1931) 

Material. 2: 1 o" . 

Distribution. Described from Sudan, later found in Yemen (Merz et al., in 
press). New for Saudi Arabia. 

Host plants. Unknown. 

Pest status. Not a pest species. 

Dicheniotes multipunctatus sp. n. Figs 11-13 

Dicheniotes sp. - Merz et al. (in press). 

Material. Holotype 6: KENYA: Rt. B4 nr. Lake Baringo, 28.XI.1986, 
A. Freidberg (TAU). Paratypes: KENYA: 1 S , Voi, Rt. A109, 4,6.V.1991 , A. Freidberg 
& F. Kaplan (MHNG); 1 9 , (N. W.) Kainuk, on Morun River, 25 .XL 1989, A. Freidberg 
& F. Kaplan (TAU). SAUDI ARABIA: 1 9 , Abha, Madenate Ameer Sultan, 22.IL- 
25.V.2002, H. A. Dawah (MHNG). TANZANIA: 1 2 , Same, Rt. Bl, 8-16.LX.1992, A. 
Freidberg (TAU) ; 1 S , 1 9 , Dar es Salam , 20 km N , Rt . B 1 2 1 2 , 4 .IX . 1 996 , A . Freidberg 
(TAU). YEMEN: 1 9 , ar-Rujum, 1 6. X. 2000- 15. 1.200 1 , Malaise trap, A. van Harten & 
A. M. Hager (MHNG); 1 9 , Lahj, VIII.2000, Malaise trap, A. van Harten & A. Sallam 
(ZMUA); 1 9 , 12 km NW of Manakhah, 27 .111-5 .V.2002, Mal. trap, A. van Harten; 1 9 , 
same, but 5.V-17.VI.2002 (MHNG). The holotype is double-mounted on a minutien 
pin on a polyporus block and is in excellent condition. 

Etymology. The species is named after the wing pattern with numerous hyaline 
and subhyaline spots. 



996 



B. MERZ & H. A. DAWAH 




FlGS 11-13 
Dicheniotes multipunctatus sp. n.: 11, head, lateral view (holotype); 12, wing of male (holotype); 
13, wing of female (paratype from Yemen, slightly modified after Merz et al., in press). 



Diagnosis. This species belongs to the group of species of Dicheniotes with a 
hyaline spot in cell r2+3 at the tip of R2+3 (Figs 12-13) and normal head shape without 
oblique oral opening or large labella (Fig. 11). It differs from all other species of the 
genus by the presence of 3 hyaline spots in cell rl, which are either small, or elonga- 
ted and extending into cell r2+3; remainder of wing with numerous hyaline or subhya- 
line spots; sexual dimorphism of wing pattern present, but weak. Scutum thickly grey 
microtrichose, with distinct, coarse white setulae. Abdomen reddish-brown, sub- 
shining. 

Description. Wing length. Male: 3.15-3.35 mm; female: 2.70-3.35 mm. 

Head (Fig. 11). Predominantly yellow, but ocellar triangle and a paired spot on 
occiput dorsal to occipital foramen greyish black; posterior half of frons, frontal and 
orbital plates silvery; face indistinctly whitish yellow; palpus and antenna entirely 
yellow, only apical two third of arista brown; head in profile about 1 .2 times as high as 
long; gena narrow, about one tenth as wide as compound eye; at anterior margin with 
2-3 rows of brown, rather short subvibrissal setulae of equal length; frons slightly 
wider than long (measured from posterior margin of posterior ocelli to base of 
antenna), tapering anteriorly; covered with numerous white setulae anteriorly; face 
shallow concave, without carina, vibrissal corner produced anteriorly to about same 
level as frontofacial corner; scape and pedicel black setulose; postpedicel 1.5 times as 
long as wide, dorsally slightly concave, rounded dorsoapically; arista almost bare, rays 
at most as long as basal diameter of arista; palpus short, not projecting beyond vibrissal 



FRUIT FLIES FROM SAUDI ARABIA 997 

comer. The following paired pale brown setae are present: 1 genal, 3 frontal, 2 orbital, 

1 ocellar, 1 medial vertical seta; 2 lateral vertical setae pale brown to white depending 
on illumination; postocellar and paravertical setae white; postocular setae mixed with 
long, white and short, black setulae. 

Thorax. Scutum, dorsal surface of scutellum and pleura posterior to anterior 
spiracle black, very densely grey microtrichose, mat, black ground colour entirely 
invisible; postpronotum, posterior two third of notopleuron, pleura anterior to anterior 
spiracle and on ventral side of scutellum mat yellow to pale brown. Mesonotum 
covered with numerous short, white, coarse setulae; prosternum white setulose. The 
following pale brown, paired setae are present: 1 dorsocentral inserted closer to level 
of anterior supra-alar seta than to suture, 1 prescutellar acrostichal, 1 presuturai supra- 
alar, 1 postsutural supra-alar, 1 intra- alar, 1 postalar, 1 postpronotal, 2 notopleural and 

2 (pairs of) subequal scutellar setae; 2 anepisternal, 1 katepisternal and 1 anepimeral 
seta. 

Legs. Entirely yellow; hindtibia anterodorsally with a row of 4-7 outstanding 
setulae. 

Wing male (Fig. 12). Very wide, about 1.8 times as long as wide, apically 
broadly rounded, posteriorly with large anal lobe; Rl dorsally setulose except for a gap 
opposite subcostal break; R4+5 ventrally with 1-6 setulae proximally R-M, dorsally 
with 3-5 black setulae to level of R-M. Pattern orange brown, covering most of surfa- 
ce distal to subcostal break and interrupted by about 20 hyaline and 10 slightly yello- 
wish spots; cell c hyaline with narrow brown stripes at base and tip; pterostigma with 
hyaline spot; rl with 3 hyaline spots with the basal two spots larger, extending into cell 
r2+3; r2+3 with 6-8 small hyaline or pale yellowish spots; br with 1-2 hyaline spots; 
r4+5 with 4-5 spots, apex largely black; dm with 2-3 hyaline spots; m with 4 spots 
which are separated from each other by distance greater than their diameter; cu2 and 
anal lobe with about 4 spots. Halter and calypteres yellow. Lower, ventral calypter 
smaller than upper, dorsal. 

Wing female (Fig. 13). Shape and chaetotaxy as male; pattern darker brown, 
with hyaline spots clearly separated from dark area and only few yellowish spots pre- 
sent; only basal hyaline spot of rl occasionally penetrating into r2+3, second hyaline 
spot of rl always isolated from hyaline spot in r2+3. The female from Kainuk (Kenya), 
the smallest specimen (wing length: 2.70 mm; all other specimens > 3.0 mm) has only 
2 hyaline spots in r2+3, remaining pattern as in other specimens. 

Abdomen male. Strongly reddish-brown, subshining; basal two tergites mainly 
white setulose, other tergites covered with short black setulae. Epandrium reddish; 
structure not studied. 

Abdomen female. Preabdomen as male; oviscape black, shining, pale brown 
setulose, about as long as length of 2.5 preceding tergites combined; eversible mem- 
brane with numerous denticles; aculeus evenly narrowed, apically sharply pointed. 

Distribution. Eastern Afrotropical Region (Kenya, Tanzania) and Arabian 
Peninsula (Saudi Arabia, Yemen). 

Host plants. Unknown. The specimens from the Arabian Peninsula were col- 
lected in Malaise traps. 

Pest status. Not a pest species. 



998 B. MERZ & H. A. DAWAH 

Remarks. The generic position of this species may be subject to discussions. In 
the key of Munro (1947) it runs with some problems to Platensina Enderlein because 
of the rather thick, ash-grey microtrichose mesonotum and the chaetotaxy (couplet 21 
in Munro, 1947). However, some species of Dicheniotes Munro, such as D. polyspila 
(Bezzi), have also an ash-grey microtrichose mesonotum and a very similar type of 
wing pattern and head shape (some 30 specimens of both sexes from Kenya, along road 
from Nairobi to Mombasa between Hunters Lodge and Mtito Andei studied, MHNG). 
Its inclusion in Dicheniotes (3 frontal setae, 2 orbital setae, 2 pairs of scutellar setae, 
postocular setae mixed white and black, wing of Tephrella type of Munro, 1947) seems 
therefore justified. 

Within Dicheniotes the new species differs from all congeners in wing pattern 
with 3 hyaline spots in cell rl, of which the apical spot is smaller than the two basal 
spots, but still clearly present, and the more numerous hyaline spots on the remaining 
wing surface. Because of the presence of a hyaline spot in r2+3 below tip of R2+3 
D. multipunctatus runs to couplet 9 in Munro (1947, p. 178) next to D. polyspila which 
is morphologically similar with the same reddish abdomen and the thick, ash-grey 
microtrichose mesonotum and thus differing from the much more subshining mesono- 
tum and shining black abdomen of other species, such as D. angulicornis , D. erosa 
(Bezzi) or D. katonae (Bezzi). The new species differs from D . polyspila by the follo- 
wing set of characters in addition to those given above: wing with less conspicuous di- 
morphism (very prominent in D. polyspila: male with basal half of rl and r2+3, and 
cell br entirely hyaline, and r2+3 at tip with two broadly confluent hyaline spots; fe- 
male with more isolated spots), hyaline spots often not sharply separated from black 
area (usually very sharply separated in D. polyspila), and oviscape much shorter than 
preabdomen (in D. polyspila about as long as preabdomen). 

Gymnaciura austeni (Munro, 1935) 

Material. 2: 1 $ . 

Distribution. Entire Afrotropical Region, Yemen (Merz et al., in press). New for 
Saudi Arabia. 

Host plants. Unknown. 

Pest status. Not a pest species. 

Metasphenisca hazelae (Munro, 1947) 

Material. 2: 1 6 , 3 $ 9 . 

Distribution. Eastern Africa. New for Saudi Arabia. 

Host plant. Barleria rehmanni (Acanthaceae) (Munro, 1947). 

Pest status. Not a pest species. 

Remarks. Hancock (in litt.) found another specimen from Saudi Arabia in the 
unsorted material of the NHML: 1 S , Dhamar, 3. IX .62 (or 42), G . Popov (NHML). 

Metasphenisca negeviana (Freidberg, 1974) 

Material. 2: 1 S . 

Distribution. Near East. Recorded from Saudi Arabia by Freidberg & Kugler 
(1989). 



FRUIT FLIES FROM SAUDI ARABIA 999 

Host plant. Blepharis attenuata (Acanthaceae) (Freidberg & Kugler, 1989). 
Pest status. Not a pest species. 

Oxyaciura tibialis (Robineau-Desvoidy, 1830) 

Material. 2:29?. 

Distribution. Western Palaearctic Region to Afghanistan, Ethiopia. New for 
Saudi Arabia. 

Host plants. Lavandula spp.,Nepeta septemcrenata (Lamiaceae) (Freidberg & 
Kugler, 1989; Hendel, 1927; Merz, 1992). 

Pest status. Not a pest species. 

Paraspheniscoides binarius (Loew, 1861) 

Material. 2: 1 9 . 

Distribution. Afrotropical Region and islands of the Indian Ocean, Yemen 
(Merz et al., in press). New for Saudi Arabia. 

Host plants. Lippia spp. and Lantana spp. (Verbenaceae) (Munro, 1947). 
Pest status. Not a pest species. 

Paraspheniscus debskii (Efflatoun, 1924) 

Material. 2: 1 9 . 

Distribution. Described from Egypt. New for Saudi Arabia. 

Host plant. Stachys aegyptiaca (Lamiaceae) (Efflatourn, 1924; Freidberg & 
Kugler, 1989; Hendel, 1927). 

Pest status. Not a pest species. 

Tribe Tephritini 
Campiglossa genus group 

Campiglossa ignobilis (Loew, 1861) 

Material. 2: 34 S S , 41 9 9 , 1 without abdomen. 

Distribution. Eastern Afrotropical Region, Yemen. New for Saudia Arabia. 
Host plant. Sonchus oleraceus (Asteraceae) (Munro, 1957a). 
Pest status. Because its host is a weed in some parts of the world, this species 
may be useful in the natural regulation of S. oleraceus. 

Desmella sp. nr myiopitoides (Bezzi, 1908) 

Material. 2: 29 9. 

Distribution. D. myiopitoides is known from the Eastern Afrotropical Region. 
Recently, a specimen of the D. myiopitoides-group was also found in Oman (Jebel 
Shams, 2500m, 7.V.1989, leg. M. J. Ebejer, MHNG, new record). New for Saudi 
Arabia. 

Host plant. D. myiopitoides was reared in South Africa from flowers of 
Chrysocoma tenui/olia (Asteraceae) (Munro, 1925). 

Pest status. Not a pest species. 

Remarks. As Munro (1957a) pointed out this genus is in bad need of a revision. 
No further progress has been carried out since then. The two females at hand key out 
as D. myiopitoides (Bezzi), but this may be a complex of different species (Munro, 



1000 B. MERZ & H.A. DAWAH 

1957a). One female has a wing pattern similar to Fig. 137, and the other is similar to 
Fig. 139 in Munro (1957a). No males are available which may have good characters 
(Munro, 1957a). For these reasons the specimens from Saudi Arabia cannot be iden- 
tified unambiguously. The genus is newly recorded for Saudi Arabia. 

Dioxyna sororcula (Wiedemann, 1830) 

Material. 2:599. 

Distribution. Tropics and Subtropics of the Old World, Australia, introduced to 
Hawaii. New for Saudi Arabia. 

Host plants. Bidens spp., Guizotia sp., Lactuca sativa (Asteraceae) (Freidberg 
& Kugler, 1989; Merz, 1992; White & Elson-Harris, 1992). 

Pest status. A potential or minor pest on some cultivated plants {Lactuca in the 
Philippines). On the other hand it may be important in the natural regulation of Bidens 
spp. which are weeds in some parts of the world (White & Elson-Harris, 1992). 

Tanaica Munro, 1957 

Type species: Emina hyalipennis Bezzi, 1924 

Diagnosis. Within the Campiglossa group of genera Tanaica (Munro, 1957a) 
may be recognized by the flattened head (Fig. 24) which is longer than high in profile 
and the narrow gena; by the long labella which are longer than heigth of head and 
which are projecting distinctly beyond anterior vibrissal corner; by the wing, which is 
either entirely hyaline or which has a faint reticulate pattern (Figs 15, 25); by the legs, 
with the femora black except for the apical quarter; and by the male terminalia: 
preglans with two setulose areas and glans with two sclerotized plates of uneven length 
emerging from acrophallus, with the longer plate strongly concave (Figs 29-30). 

Redescription. Head (Figs 14, 24). In profile flattened, longer than high; gena 
narrow, less than 0.8 times as wide as width of postpedicel; heigth of gena less than 
0.15 times the heigth of compound eye; frons bare; frontal stripe (as defined by Munro, 
1957a) indistinct, almost invisible; face concave; vibrissal corner distinctly projecting 
beyond frontofacial corner; mouthparts strongly geniculate with labella longer than 
heigth of head and projecting beyond vibrissal corner; palpus thin, parallel-sided, as 
long as labella; scape and pedicel black setulose; postpedicel about 1 .5 times as long 
as wide, dorsoapically rounded; arista almost bare. Chaetotaxy: 2 black frontal setae; 
2 orbital setae, anterior black, posterior short, white; ocellar seta black; medial vertical 
seta black; lateral vertical seta and postocellar seta white; paravertical seta absent or 
short and white; medial postocellar seta absent; postocular setae mixed white and 
black. 

Thorax. Scutum white setulose, more or less uniformly grey microtrichose, 
without distinct brown longitudinal stripes; scutellum apically with small yellow patch. 
Chaetotaxy (all setae black): 1 postpronotal seta; 2 notopleural setae, the posterior 
shorter; 1 presuturai supra-alar seta; 1 postsutural supra-alar seta; 1 intra-alar seta; 1 
postalar seta; 1 dorsocentral seta distinctly anterior to level of postsutural supra-alar 
seta; 1 acrostichal seta; 2 (pairs of) scutellar setae, the apical less than half as long as 
the basal; 1 anepisternal seta and 1 katepisternal seta, both black; 1 white, lanceolate 
anepimeral seta. 



FRUIT FLIES FROM SAUDI ARABIA 1001 

Legs. Mat yellow, but basal 0.7-0.8 of all femora black; coxae and trochanters 
black or mixed black and yellow; midtibia with 1 apicoventral black seta. 

Wing (Figs 15, 25). Hyaline or with faint reticulate pattern; pterostigma hyaline 
or brown with round hyaline spot medially; posterodistal extension of cell bcu short, 
but distinct; R-M in distal third of cell dm; distance between crossveins equal to length 
of DM-Cu; veins bare, only Rl with usual dorsal row of setulae, which is interrupted 
opposite level of tip of Sc, and ventrally with 0-5 setulae in distal half of pterostigma. 
Calypteres yellow, lower, ventral calypter strip-like, much smaller than upper, dorsal; 
halter yellow. 

Abdomen male. Shining black or grey microtrichose and then with indistinct 
black paired spots on tergites; sternite 5 (Figs 16, 26) smoothly concave apically; epan- 
drium (Fig. 18) ovoid, with distinct epandrial process; medial surstylus with 2 prensi- 
setae of very unequal length, medial prensiseta much larger (Fig. 19); preglans area 
with two areas of fine setulae (Fig. 20); glans with strongly sclerotized, tube-like acro- 
phallus; vesica twice as long, with two sclerotized plates, the longer twice as long as 
the shorter, concave; shorter plate distally truncate. 

Abdomen female. Preabdomen as male; oviscape black, fine black setulose; at 
most as long as preceding 3 tergites combined; aculeus (Figs 21-22) evenly pointed 
apically; two spherical, pyrifrom spermathecae with small papillae (Fig. 23). 

Remarks. Tanaica was described by Munro (1957a) in the Ensina series of 
genera for a species with entirely hyaline wings and a shining black abdomen. As it 
was already admitted by Munro (1957a), the genera placed in this group do not belong 
to a homogenous group, but are merely an assemblage of taxa which cannot be placed 
elsewhere. In fact, Ensina Robineau-Desvoidy is now placed in the tribe Noeetini, 
whereas the other genera in the tribe Tephritini (Norrbom et al., 1999a). Ptosanthus 
Munro, Sphenella Robineau-Desvoidy and Telaletes Munro belong to the Sphenella 
genus group (Freidberg, 1987; Norrbom et al., 1999a), whereas the relationships of the 
two monotypic genera Deroparia Munro and Namwambina Munro are still not satis- 
factorily known. Tanaica, the last of these genera, was correctly associated with the 
Campiglossa genus group (Norrbom et ai., 1999a) because of the presence of setulose 
preglans area (absent only in Antoxya Munro and Dioxyna Frey), the flattened head and 
the mostly black setae on head and thorax. 

The only species included in Tanaica by Munro (1957a) exhibits two unusual 
characters for the Campiglossa genus group and may have influenced the placement of 
the genus in the Ensina series: the entirely hyaline wing and the shining black 
abdomen. However, with the inclusion of the two species described below the concept 
of the genus is modified, including also species with faint reticulate wing pattern and 
with grey microtrichose abdomen. Superficially they resemble species of Desmella 
Munro and Dioxyna. Whereas the latter genus is somewhat isolated by the lack of 
setulae on the preglans area, it is possible to separate safely Desmella from Tanaica 
only by the structure of the glans. More careful comparative studies of all included 
species are, however, needed to establish whether both genera may be maintained or 
eventually be synonymized. 

In order to place Tanaica in the Campiglossa genus group, a key to the 
Afrotropical and Near Eastern genera is provided. Excluded are Homoeotricha Hering 



1002 B. MERZ & H.A. DAWAH 

(6 species in Central and Eastern Palaearctic, to the west to Kazakhstan and Kyrghyzia) 
and Oxyparna Korneyev (2 species in mountains between Kyrghyzia and China). 

1 One pair of frontal setae present (rarely on one side 2-3 frontal setae 
present, in these cases lunule very high and scape white setulose) 2 

1* At least 2 pairs of subequal frontal setae present; scape always black 

setulose 3 

2 Scape black setulose; frons setulose anteriorly; lunule normal; posterior 
notopleural seta black; postpedicel about twice as long as wide 

Antoxya Munro 

[1 Afrotropical species; A. oxynoides (Bezzi)] 

2* Scape white setulose; frons bare; lunule very high, a semicircle; poste- 
rior notopleural seta usually white; postpedicel barely longer than wide 

Oxyna Robineau-Desvoidy 

[23 species in Nearctic and Palaearctic Regions] 

3 Apical scutellar setae at least 0.75 times as long as basal scutellar setae 4 

3* Apical scutellar setae at most 0.6 times as long as basal scutellar setae, 

or entirely absent 5 

4 Labella long geniculate, longer than heigth of head (as in Fig. 24) 

Mesoclanis Munro 

[8 Afrotropical and 1 Oriental species; 1 species introduced to Australia] 

4* Labella short geniculate, much shorter than heigth of head .... Scedella Munro 
[15 Afrotropical and 2 Oriental/ Australasian species] 

5 Apical scutellar setae entirely absent 6 

5* Apical scutellar setae present, though sometimes only little longer than 

surrounding setulae 7 

6 Head in profile higher than long; labella about 0.8 times as long as 
heigth of head; gena 0.2 times as high as eye and about as high as width 
of postpedicel; abdominal tergites dull, with indistinct paired spots on 

tergites 3-5; male: preglans area setulose Lethyna Munro 

[7 Afrotropical species, 1 unidentified species in Yemen (Merz et al., in 
press)] 

6* Head in profile wider than long; labella longer than heigth of head; ge- 
na at most 0.15 times as high as eye; about 0.7 times as high as width of 
postpedicel; paired spots on abdominal tergites distinct; male: preglans 

area bare Dioxyna Frey, part 

[10 species, worldwide distribution] 

7 Posterior notopleural seta white Campiglossa Rondani, part 

[almost 200 species worldwide, most diverse in Afrotropical, Nearctic 

and Palaearctic Regions] 
7* Posterior notopleural seta black 8 

8 Dorsocentral seta distinctly posterior to level of postsutural supra-alar 

seta; head flattened Campiglossa Rondani, part 

[here only C. martii (Becker) from the Canary Islands and Southern Spain] 
8* Dorsocentral seta closer to suture than to postsutural supra-alar seta; 

shape of head variable 9 



FRUIT FLIES FROM SAUDI ARABIA 1003 

9 Head in profile higher than long; labella usually shorter than heigth of 
head; wing pattern usually reticulate, rarely banded; male: preglans area 

setulose Campiglossa Rondani, part 

[see couplet 7] 

9* Head in profile flattened, longer than high; labella longer than heigth of 

head, usually projecting beyond vibrissal corner; other characters variable . . 10 

10 Wing pattern strong, usually of banded appearance, with clear contrast 
between brown and hyaline areas; pterostigma uniformly brown without 
central hyaline spot; femora at least in basal half black; male: preglans 

area setulose Desmella Munro 

[3 Afrotropical and 2 Palaearctic species] 
10* Wing pattern weak, reticulate, or wing entirely hyaline; border between 
brown and hyaline areas indistinct; pterostigma uniformly hyaline or 
brown with central hyaline spot; femora yellow or partly black; male: 
preglans area setulose or bare 11 

11 At least midfemur mostly yellow. Male: preglans area bare . Dioxyna Frey, part 
[here only the Palaearctic species D. bidentis (Robineau-Desvoidy) with 

short apical scutellar setae] 
1 1 * All femora at least in basal 0.7 black. Male: preglans area setulose (Figs 

20, 29) Tanaica Munro 

[3 Afrotropical species, see below] 

Key to species of Tanaica Munro, 1957 

1 Wing with faint, but distinct reticulate pattern (Fig. 15); abdomen grey 
microtrichose, not shining T. maculata sp. n. 

1* Wing entirely hyaline, at most pterostigma yellowish (Fig. 25); 

abdomen variable 2 

2 Abdomen strongly microtrichose, mat T. pollinosa sp. n. 

2* Abdomen shining black, only very thinly microtrichose 

T. hyalipennis (Bezzi, 1924) 

Tanaica hyalipennis (Bezzi, 1924) 

Ensina hyalipennis Bezzi, 1924: 549. Holotype 9: SOUTH AFRICA: Cape, Cedarbergen, 
Clanwilliam, 4000-5000 ft, IX. 1923, leg. K. H. Barnard (SAMCT) (not examined). 

Material. 1 3 , 2 $ $ , NAMIBIA: Keetmanshoop, 12.IX.2003, leg. A. Freidberg 
(TAU). 

Remarks. The species was extensively described by Bezzi (1924), and redes- 
cribed by Munro (1929, 1957a) by adding new characters. Munro (1929) stated that the 
abdomen is shining black in the holotype, whereas it was described as «black, grey 
dusted» by Bezzi (1924). Because it was not possible to study the holotype we assume 
here that Munro correctly interpreted the colour of the abdomen. On the other hand, the 
illustration of the glans by Munro (1957a) is misleading because the distal two sclero- 
tizations were not illustrated nor mentioned in the description. It is possible that he 
studied a teneral specimen with the sclerotization not yet fully developed. The above 



1004 B. MERZ & H. A. DAWAH 

mentioned male was dissected and it was found that the terminalia are the same as in 
T.pollinosa (Figs 27-30). According to available information (Munro, 1957a; Norrbom 
et al., 1999b) T. hyalipennis has a rather narrow distribution in the Western part of 
South Africa and Namibia. This species was reared from Chrysanthemoides monilife- 
ra (Asteraceae) (Munro, 1957a). 

Tanaica maculata sp. n. Figs 14-23 

Material. Holotype 3: SAUDI ARABIA: Abha, Madenate Ameer Sultan, 
22.II.-25.V.2002, H. A. Dawah (MHNG). Paratypes: SAUDI ARABIA: 2c? S , 3 9 9 , 
same data as holotype (CHD, MHNG, NMWC). ETHIOPIA: 15 c? c?, 92 9, Mojo, 
75 km SE Addis Abeba, 20 .XII. 1989, A. Freidberg & F. Kaplan (MHNG, TAU). 
KENYA: 1 6 , W-Nariobi (ILRAD), 1900 m, 1. III. 1993, leg. B. Merz; 9c? c?, 5 9 9, 
same data, but 2. III. 1993; 1 9 , same data, but 3. III. 1993; 1 9 , same data, but 5. III. 1993; 
1 c? , 2 9 9 , same data, but 8 .III. 1993; 1 6 , 2 9 9 , same data, but 16.III.1993; 2 c? 6 , 40 
km SE Nairobi, [on road] A 109, 1650 m, 17.III.1993, leg. B. Merz; 3c? c? , 75 km SE 
Nairobi (A 109), 1700 m, 17.III.1993; 19, 135 km SE Nairobi (A 109), 1200 m, 
17.III.1993, leg. B. Merz (MHNG); lc?,Rt. 104, 15 km SE Nairobi, 29 .IV.- 15. V. 1991, 
A. Freidberg & F. Kaplan (TAU). The holotype is double-mounted on a minutien pin 
on a polyporus block and is in good condition with the right postocular seta and the 
right dorsocentral seta broken off and scutum slightly sunken medially. 

Etymology. The name refers to the patterned wing, which is unique among 
known species of the genus. 

Diagnosis. Within Tanaica this species is easily recognizable by the presence of 
a reticulate, weak wing pattern (Fig. 15). Cell rl with 3 hyaline spots. The other two 
species of the genus have an entirely hyaline wing. The abdomen is microtrichose as 
in T.pollinosa. 

Description. Wing length. Male: 2.1-3.4 mm; female: 2.0-3.5 mm. 

Head (Fig. 14). Ground colour mat yellow; face, gena and parafacial whitish 
yellow; frons and antennae orange-yellow. In profile slightly wider than high, with 
swollen occiput; gena about 1/8 as high as eye, the latter about 1.3 times as high as wi- 
de; vibrissal corner strongly projecting beyond anterior base of antennae; frons bare, 
sides slightly converging towards lunule; interfrontal stripe indistinct, only visible as 
whitish line under certain angle; paravertical seta white, short, not longer than setulae 
on ventral half of occiput. 

Thorax. Mesonotum black, densely ash-grey to silvery grey microtrichose, 
with traces of 3 brown longitudinal stripes over lines of dorsocentral setae and in midd- 
le; postpronotum and tip of scutellum yellowish; scutum covered with very dense, whi- 
te setulae; setulae on pleura less numerous; prosternum white setulose. Chaetotaxy as 
described for the genus. 

Legs. As described for the genus. 

Wing (Fig. 15). With a weak reticulate pattern; cell c often with a pale brown 
mark in middle; pterostigma brown with a hyaline spot in middle or uniformly yellow; 
rl with 3 hyaline spots; preapically on level of tip of R2+3 with 4 large spots in one 
line; apex of wing with small hyaline spot; remaining pattern faint, especially in anal 
lobe. 



FRUIT FLIES FROM SAUDI ARABIA 



1005 







Vi fu 'à \,\}iJs 



18 




Figs 14-23 
Tanaica maculata sp. n. (16-20, male; 21-23, female, paratypes from Abha): 14, head, lateral 
view; 15, wing; 16, abdominal sternites 4 and 5; 17, epandrium, lateral view; 18, same, caudal 
view; 19, medial surstylus with prensisetae enlarged; 20, aculeus; 22, tip of aculeus; 23, sper- 
matheca (only one illustrated). 

Abdomen male. Preabdomen as in T. pollinosa. Terminalia (Figs 16-20). 
Epandrium (Fig. 18) ovoid, black, but epandrial process brown; the latter with serrate 
border, visible in lateral view (Fig. 17); medial surstylus with two unequal prensisetae 
present on prominent plate, the lateral prensiseta much larger than the medial (Fig. 19); 
pregi ans area with 2 rows of short setulae; glans (Fig. 20) with well sclerotized basal 



1006 B. MERZ & H. A. DAWAH 

acrophallus and with 2 sclerites in vesica, of which one is long, apically pointed and 
medially curved and the other sclerite is short, widening apically with concave distal 
margin. 

Abdomen female. In all characters, including terminalia (Figs 21-23) as in 
T.pollinosa (see below). 

Distribution. Ethiopia, Kenya, Saudi Arabia. 

Host plants. Unknown. The specimens from Kenya were swept from various 
herbs in grassland areas, whereas the specimens from Saudi Arabia were collected in a 
Malaise trap. 

Pest status. Not a pest species. 

Remarks. This species is easy to recognize within Tanaica by the reticulate wing 
pattern. The species is somewhat variable in wing pattern. Specimens from Ethiopia 
and Kenya tend to have a weaker pattern, with notably cell c entirely hyaline (without 
central darker spot) and with pterostigma uniformly yellow (in Saudi Arabian spe- 
cimens usually brown with central hyaline spot). However, male and female termina- 
lia are the same for populations from Kenya and Saudi Arabia, and few specimens from 
Kenya have the same wing pattern as those from Saudi Arabia. At the present moment 
it is not possible to decide whether these populations represent different species. 
Further studies, in particular the knowledge of host plants, would give further eviden- 
ce about their status. 

T. maculata has the same type of wing pattern as do some Dioxyna Frey. 
However, all Dioxyna lack the two rows of setulae on the preglans area (present in 
Tanaica). Moreover, they lack, except for D. bidenti s , the apical scutellar setae. The 
latter species differs from T. maculata by the almost entirely yellow legs (femora 
blackish in T. maculata). 

Tanaica pollinosa sp. n. Figs 24-33 

Material. Holotype a: SAUDI ARABIA: Abha, Madenate Ameer Sultan, 
22.II.-25.V.2002, H. A. Dawah (MHNG). Paratypes: SAUDI ARABIA: 1 S , 2 9 9 , sa- 
me data as holotype (CHD, MHNG). SOUTH AFRICA: 3<? 6, 2$ 9 , N Transvaal, 
Sautpansberg. 1500 m . 15km NW Louis Trichardt, 25 .XII. 1994, A. Freidberg. The 
29 9 with additional label: ex ? seeds Tripteris auriculata, 1.1.1995 (TAU, MHNG). 
The holotype is double-mounted on a minutien pin on a polyporus block and is in good 
condition with few setae on head and thorax broken off. 

Etymology. The name refers to the grey microtrichose abdomen. 

Diagnosis. This species differs from the other species from Saudi Arabia by its 
entirely hyaline wing without traces of a pattern (Fig. 25). The abdomen is densely 
grey microtrichose; tergites 2-5 each with a pair of indistinct brown spots. 

Description. Wing length (both sexes). 2.4-2.6 mm. 

Head (Fig. 24). Ground colour mat yellow, but occiput and ocellar triangle 
black; face and gena whitish yellow; frons and antenna orange-yellow. Head in profile 
wider than high, with prominent occiput; gena narrow, less than one sixth as high as 
compound eye; the latter ovoid, about 1 .25 times as high as wide; vibrissal corner pro- 
jecting beyond level of antennal base; frons bare; interfrontal stripe indistinct, narrow, 
almost white; paravertical seta white, shorter than posterior orbital seta. 



FRUIT FLIES FROM SAUDI ARABIA 



1007 




Figs 24-33 
Tarmica pollinosa sp. n. (24-30, male; 31-33, female, paratypes from Abha): 24, head, lateral 
view; 25, wing; 26, abdominal sternites; 27, epandrium, lateral view; 28, same, caudal view; 29, 
glans, lateral view; 30, glans, frontal view; 31, aculeus; 32, tip of aculeus; 33, spermathecae. 



1008 B. MERZ & H. A. DAWAH 

Thorax. Mesonotum black, densely silvery grey microtrichose; postpronotal 
lobe and posterior half of scutellum yellow; scutum densely covered with white, 
lanceolate setulae; pleura slightly darker grey, setulae on anepisternum, katepisternum 
and anepimeron less numerous and more acuminate; prosternum grey, white setulose. 
Chaetotaxy as described for the genus. 

Legs. As described for the genus. 

Wing (Fig. 25). Entirely hyaline, only stigma faintly yellowish. 

Abdomen male. Tergites darker grey than mesonotum, but still distinctly 
golden-grey microtrichose; each tergite with a pair of almost parallel-sided spots which 
are forming a more or less uniform band from base to tip of abdomen; setulae short, 
dense, lanceolate; sternites (Fig. 26) darker grey, sparsely setulose; sternite 5 shallowly 
concave with smooth apicolateral corners; epandrium (Figs 27-28) black, but epandrial 
process brown; the latter prominent and distinctly projecting, with serrate border; inner 
terminalia as usual in tribe; aedeagus (Figs. 29-30) with 2 rows of setulae on preglans 
area; glans with strong basal sclerotization and 2 curved, unevenly long plates forming 
an open tube; the longer apically pointed and strongly curved medially; the shorter 
plate distinctly widening distally, forming an open rim; vesica as broad as acrophallus, 
distally truncate, rather small. 

Abdomen female (Figs 31-33). Coloration and chaetotaxy of pregenital 
segments as male; oviscape black, fine setulose, without white setae, as long as pre- 
ceding 3 tergites combined; aculeus apically evenly pointed, without step; 2 almost 
spherical spermathecae, covered with short papillae; aculeus length: 0.77 mm (n = 1). 

Distribution. Saudi Arabia, South Africa. 

Host plant. Tripteris auriculata (Asteraceae). 

Pest status. Not a pest species. 

Remarks. This species differs from T. hyalipennis (Bezzi), the other species with 
entirely hyaline wings, only in the colouration of the abdomen. It is shining black in 
the latter, but grey microtrichose in T. pollinosa. All other external characters and the 
terminalia of both sexes are the same in both species. 

Spathulina genus group 

Spathulina acroleuca (Schiner. 1868) 

Material. 2: 2 9 9 . 

Distribution. Widespread in the Afrotropical, Southern Palaearctic, Oriental and 
Australasian Regions. Recorded from Yemen by Merz et al. (in press). New for Saudi 
Arabia. 

Host plants. Various Asteraceae, such as Bidens, Ceruana, Matricaria and 
Spilanthes (Munro, 1938). 

Pest status. The species was recorded as minor pest on Mexican sunflower 
(Tithonia diversi/olia) in the Philippines (White & Elson-Harris, 1992). 

Sphenella genus group 

Sphenella marginata (Fallen. 1814) 
Material. 2: 1 6 , 1 9 . 
Distribution. Palaearctic and Afrotropical Regions. New for Saudi Arabia. 



FRUIT FLIES FROM SAUDI ARABIA 1009 

Host plants. Senecio spp. (Asteraceae) (Freidberg & Kugler, 1989; Merz, 1994). 

Pest status. May be important in the natural regulation of those Senecio spp. 
which are weeds in some parts of the world (White & Elson-Harris, 1992). 

Remarks. After the study of large series from the Palaearctic and Afrotropical 
Regions we conclude that the differences given to separate S. marginata s.str. 
(Palaearctic subspecies) from S. marginata austrina Munro, 1957 (Afrotropical sub- 
species) (Munro, 1957b) are unreliable with many intermediate specimens known. The 
two taxa are considered here to belong to the same species and they are herewith 
synonymized (syn. n.). 

Sphenella setosa sp. n. Figs 34-41 

Material. Holotype 6: «SAUDI ARABIA: Aseer, Abha, Farm Centre, 14.IV. 
2001 , H. A. Dawah»; «Tephritis sp. cT , lower angle anal cell more prominent than any 
sp. photo in Die Fliegen» [J. C. Deeming's handwriting] (MHNG). Paratype: lo*, 
SAUDI ARABIA: Aseer, Abha, Farm Centre, 14.IV.2001, H. A. Dawah (MHNG). The 
holotype is glued laterally on a card point, and is in good condition with left arista and 
few setae missing (left ocellar seta, few postocular setulae, left postpronotal seta, both 
left notopleural setae, left presuturai supra-alar seta; left postsutural supra- alar seta; 
both right scutellar setae). 

Etymology. The name refers to the strongly setulose legs. 

Diagnosis. This species differs from all described Sphenella by the strongly 
setulose hindfemur and hindtibia (Figs 36-37), the wing (Fig. 35) with the ill-defined 
pattern of paler and darker areas on the entire surface, the strongly modified sterni te 5 
with 2 medial tubercles (Fig. 38), and the very large, black, globose epandrium (Fig. 
40). The wing pattern resembles superficially species of Paratephritis Shiraki, Tela- 
letes Munro and Orotava Frey, but the combination of bare R4+5, flat scutellum, 
presence of antero ventral setae on hindfemur and presence of only one long, pointed 
prensiseta on the medial surstylus are typical for Sphenella. Other diagnostic characters 
of this species are the mixed white and black postocular setulae, the yellow legs with 
only hindfemur black in basal half, the yellow postpedicel, and the sclerotization of the 
glans. 

Description (male only). Wing length. 3.50-3.75 mm. 

Head (Fig. 34). Mat yellow, but ocellar triangle and occiput grey microtricho- 
se. In profile 1.25 times as high as long, gena 0.15 times as high as compound eye; 
fronto-facial angle about 110°; frons white setulose anteriorly; frontal plate rather 
densely white setulose; face distinctly concave, without carina; vibrissal corner white 
setulose; gena dorsally black and ventrally white setulose. Antenna yellow, scape with 
white setulae, pedicel with black setulae; postpedicel apicodorsally slightly pointed; 
arista virtually bare. Mouthparts short geniculate, labella fleshy, not extending beyond 
vibrissal corner. Chaetotaxy: 2 black frontal setae; 1 black anterior and 1 white, shor- 
ter posterior orbital seta; 1 black ocellar seta; 1 black medial vertical seta; 1 white la- 
teral vertical seta; 1 white postocellar seta; 1 white paravertical seta; postocular setu- 
lae mixed white and black. 

Thorax. Scutum densely golden grey microtrichose, in dorsal and posterior 
view with 3 faint brown longitudinal stripes on lines of dorsocentral setae and in 



1010 



B. MERZ & H. A. DAWAH 




FlGS 34-41 

Sphenella setosa sp. n. (maie paratype): 34. head, lateral view; 35, wing; 36, hind leg, anterior 
view; 37, hind leg, posterior view; 38, abdominal sternites 4 and 5; 39, epandrium, lateral view; 
40, same, caudal view; 41, glans. 

middle; entire surface covered densely with white setulae; scutellum flat, mat yellow, 
white setulose laterally; pleura slightly more ash-grey than scutum; anepisternum and 
anepimeron white setulose; prosternum densely white setulose. The following setae are 
black: 1 postpronotal seta; 2 notopleural setae (posterior notopleural seta about half as 



FRUIT FLIES FROM SAUDI ARABIA 1011 

long as anterior notopleural seta); 1 dorsocentral seta, which is inserted well posterior 
of suture, about at level of anterior wing base; 1 presuturai supra-alar seta; 1 post- 
sutural supra-alar seta; 1 intra-alar seta; 1 postalar seta; 2 (pairs of) scutellar setae 
which are of same length; 1 katepisternal seta. Anepisternal seta and anepimeral seta 
white. 

Legs (Figs 36-37). Ground colour yellow, forecoxa whitish yellow, other parts 
brownish yellow; midfemur posteriorly at base with black stripe; hindfemur in anterior 
half black. All femora rather stout, conspicuously setulose; hindfemur with a row of 
long, black setae anteroventrally (Fig. 36), one strong, black anterodorsal seta, and 
with numerous soft, whitish, very long setulae (about as long as diameter of femur) 
postero ventrally in basal third (Fig. 37). Hindtibia anterodorsally and posterodorsally 
each with a row of conspicuous, outstanding, black setulae. 

Wing (Fig. 35). R4+5 bare on both sides; Rl setulose dorsally, with gap on level 
of subcostal break; ventrally with few setulae in distal half of pterostigma. Pattern 
brown to yellow brown, indistinctly striped, with darker areas along C, at wing tip, and 
around crossveins; other parts with a pattern of ill-defined subhyaline spots on pale 
brown surface. 

Abdomen. Mat ash-grey microtrichose with yellow hind margins of tergites 
2-5; tergite 5 about twice as long as tergite 4; all tergites white setulose, only tergite 
5 posteriorly with a row of black setulae. 

Male terminalia. Sternite 5 (Fig. 38) greatly modified, much larger than pre- 
ceding sternites; apically with two medial bulbous projections, which are separated by 
a cleft and which bear apically about 12 stout, black setulae; other setulae rather soft, 
pale. Epandrium (Figs 39-40) very large, globose, black; epandrial process small, on- 
ly little projecting; lateral surstylus separated from epandrium by a fold, strongly bent 
medially and pointed apically; medial surstylus with one very long, spine-like black 
prensiseta, which is directed posteriorly; hypandrium as in other species of the genus; 
glans (Fig. 41) rather narrow, tapered apically and truncated, with a strongly sclero- 
tized ductus. 

Distribution. Saudi Arabia. 

Host plants . Unknown. 

Pest status. Not a pest species. 

Remarks. This species keys out readily in Munro (1957b) in couplet 4 as 
Sphenella with the setulose hindfemur anteroventrally, the bare vein R4+5, and the flat 
scutellum. In addition, only one long, pointed prensiseta is developed. Sternite 5 
carries apicomedially two tubercles as in other species of Sphenella but they are much 
more developed and they are covered with more conspicuous black setae posteriorly. 
Wing pattern, however, is very different, as other species of Sphenella have large 
hyaline areas, and the black areas are usually clearly delimited. In this respect S. seto- 
sa resembles some species of Paratephritis Shiraki, Tela! êtes Munro, and Orotava 
Frey, which have the same indistinct reticulation with the ill-defined hyaline spots and 
the brown stripe over DM-Cu. Within Sphenella the new species exhibits, in addition 
to the wing pattern and the structure of sternite 5, a number of unique characters: hind- 
femur with row of black setulae anteroventrally on entire length (in other species of 
Sphenella only near apex with few setulae) with white setulae in basal third postero- 



1012 B. MERZ & H. A. DAWAH 

ventrally (not present in other Sphenella); epandrium shining black and globose (much 
more elongated and less conspicuous in other Sphenella). 

Telaletes ochraceus (Loew, 1861) 

Materiali: 26 <?; 2: ló\ 1 ? . 

Distribution. East Africa. New to Saudi Arabia. 

Host plants. Unknown. 

Pest status. Not a pest species. 

Tephritis genus group 

Arabodesis gen. n. 

Gen. nr. Trupanodesis Merz - Merz et al. (in press). 

Type species. Arabodesis reductiseta sp. n. 

Etymology. The name refers to the distribution in Arabia and its resemblance to 
Trupanodesis Merz and Tephrodesis Merz. Gender is feminine. 

Diagnosis. The new genus belongs to the Tephritis group as defined by Merz 
(1999). The absence of medial postocellar setae (Fig. 43), the presence of only white 
postocular setae (Fig. 42), two (pairs of) subequal scutellar setae (Fig. 45) and the 
rather simple structure of the glans with short vesica (Fig. 50) place Arabodesis close 
to Tephrodesis and Trupanodesis. It differs from these genera by the black posterior 
notopleural seta, the dorsocentral seta situated well behind the transverse suture at le- 
vel of postsutural supra-alar seta (Fig. 45), the convex lower calypter, the rather conspi- 
cuous posterodistal extension of cell bcu (Fig. 46), and the aculeus with an apical step 
(Fig. 53). 

Description. Head (Figs. 42-44). In profile at least 1.5 times as high as long, 
gena and parafacial narrow; vibrissal corner projecting anteriorly; frons parallel-sided, 
narrow, longer than wide, with few tiny, fine, white setulae on interfrontal stripe; ocel- 
lar triangle with few white setulae between ocelli; scape white setulose and pedicel 
black setulose; postpedicel (Fig. 44) about twice as long as wide; arista almost bare; 
proboscis capitate; palpus not projecting beyond anterior margin of vibrissal corner. 
Chaetotaxy: 1-2 frontal seta, 1 orbital seta, 1 ocellar seta and 1 long medial vertical se- 
ta black, acuminate; postocellar seta, lateral vertical seta and row of postocular setae 
white, lanceolate; medial postocellar seta absent. 

Thorax (Fig. 45). Shape as in other genera of the Tephritis-group; scutum 
covered with white setulae. Chaetotaxy: scapular setae absent; setae on thorax black, 
including posterior notopleural seta, only anepimeral seta white; 1 postpronotal seta; 
2 notopleural setae, the posterior shorter; 1 presuturai supra-alar seta; 1 postsutural 
supra-alar seta; 1 intra-alar seta; 1 postalar seta; 1 acrostichal seta; 1 dorsocentral seta 
well behind transverse suture, about at level of postsutural supra-alar seta; 2 (pairs of) 
subequal, strong scutellar setae; 1 anepisternal seta; 1 katepisternal seta; 1 anepimeral 
seta; prosternum white setulose. 

Legs. Hindfemur anteroventrally with 1-2 outstanding, longer setulae at tip; 
midtibia apicoventrally with one black seta. 

Wing (Fig. 46). Stigma about twice as long as wide; R-M crossvein situated in 
distal third of cell dm; distance between crossveins about 0.8 times as long as DM-Cu; 



FRUIT FLIES FROM SAUDI ARABIA 1013 

posterodistal extension of bcu distinct; veins bare, but Rl dorsally setulose except for 
gap on level of tip of Sc, ventrally with 0-2 setulae at tip of pterostigma. Both calypters 
with convex distal border, of subequal width. 

Abdomen male. Densely white setulose, without longer setae; sternite 5 (Fig. 
47) over 1.3 times as wide as sternite 4, distally smoothly concave; epandrium (Figs 
48-49) ventrally very wide, bulging, fused with lateral surstyli, the latter unmodified; 
epandrial process present but not very prominent; medial surstylus with 2 subequal 
prensisetae and 2-3 setulae on plate; hypandrium and associated structures as usual for 
the Tephritis group, with unpaired hypandrial arms; distiphallus long, coiled, preglans 
area bare; glans (Fig. 50) with rather simple sclerotization; sclerotized ductus apically 
with few small spines; vesica short. 

Abdomen female. Preabdomen as male; tergites 5 and 6 subequal; oviscape 
(Fig. 51) white setulose, at distal margin with numerous soft setulae; aculeus (Figs 52- 
53) with apical step; 2 spermathecae, covered with small papillae (Fig. 54). 

Remarks. Ar abode sis is a puzzling genus which exhibits an unusual combi- 
nation of characters rendering its placement within the family difficult. Lack of 
scapular setae, the presence of only white postocular setae, and the gap of setulae on 
Rl opposite tip of Se show that it belongs to the Tephritini. Based on the revisions of 
some genus groups within the tribe by Freidberg (1987, Sphenella group), Korneyev 
(1990, Campiglossa group) and Merz (1999, Tephritis group) it can be concluded that 
the new genus shares the synapomorphies of the Tephritis group as defined by Merz 
(1999): glans rather simple, weakly sclerotized and the presence of capitate 
mouthparts. Within the Tephritis group, Arabodesis exhibits a number of plesio- 
morphic character states, such as (1) the absence of medial postocellar setae (apomor- 
phic state (= A): medial postocellar setae present), (2) the black posterior notopleural 
seta (A: white), (3) the position of the dorsocentral seta well behind the suture (A: 
almost on level of suture), (4) the rather large lower calypter with convex margin (A: 
small, stripe-like lower calypter with straight margin), and (5) the apical step of the 
aculeus (A: aculeus evenly pointed), although the polarity of the latter character is 
unresolved. A black posterior notopleural seta is known in the Tephritis group only 
from few Tephritis (arnicae (Linnaeus), conura (Loew)). The position of the dorso- 
central seta is comparable only with Euare stella Hendel, which belongs to the 
Goniurellia clade (medial postocellar setae present; in addition, this genus has 2+1 
frontal and 2 orbital setae). The large lower calypter is only known from Tephritis 
Latreille (but this genus has 2 frontal and 2 orbital setae; head shape is different, and 
the dorsocentral seta is almost on level of suture). The apical step on the aculeus is 
developed in the Tephritis group only in some Tephritis (pulchra group) but an apical 
step is widespread in many genera of Tephritini. 

Arabodesis shares with Tephrodesis Merz a similar type of glans of aedeagus, 
which is evidence for a sister-group relationship and this character may be a good 
synapomorphy for these two genera. However, the position of Tephrodesis within the 
Tephritis group is not well understood, as its sister-group relationships are unknown 
(Merz, 1999). Tephrodesis differs from Arabodesis by the development of the apo- 
morphic character states of characters (2) to (5) mentioned above, but also by the 
patterned wing, the 2+1 frontal and 2 orbital setae, and a black oviscape. Further, it 



1014 B. MERZ & H.A. DAWAH 

lacks the small spinules on the inner side of the ductus of the glans which are charac- 
teristic for Arabode sis. 

Autapomorphies of Arabodesis are probably the presence of (a) only 1 orbital 
seta (in Tephritini very rare, only known from Actinoptera Rondani, but this genus 
differs in addition to the apomorphic character states (2) - (5) listed above by the very 
small pterostigma and a patterned wing), (b) the very high head (otherwise only in 
Hyalotephritis Freidberg, but this genus has the apomorphic character states (1) - (5), 
and different male terminalia) and (c) the large posterodistal extension of cell bcu (less 
conspicuous in other species of the Tephritis group). 

In the key of Merz (1999) the new genus runs to couplet 11 which needs to be 
modified as follows: 

1 1 One black orbital seta present; posterior notopleural seta black; dorso- 
central seta almost at level of postsutural supra-alar seta; wing entirely 
hyaline Arabodesis gen. n. 

11* Two orbital setae present; posterior notopleural seta white; dorsocentral 
seta closer to suture than to postsutural supra-alar seta; wing usually 
with pattern, though sometimes weak 11a 

11a Two concolourous frontal setae present 12 

11a* Three frontal setae, the anteriormost seta sometimes white 13 

This genus was first mentioned, but not formally described from Yemen based 
on three badly preserved specimens belonging to two species differing chiefly in the 
chaetotaxy of the head (Merz et al., in press). 

Arabodesis reductiseta sp. n. Figs 42-54 

Gen. nr. Trupanodesis, sp. 1 - Merz et al. (in press). 

Material. Holotype 9: SAUDI ARABIA: Abha, Madenate Ameer Sultan, 
22.II.-25.V.2002. H. A. Dawah (MHNG). Paratypes: SAUDI ARABIA: 56 S , 15 9 9 , 
same data as holotype (CHD, MHNG, NMWC, TAU). YEMEN: 1 6 , 1 9 , San'a, light 
trap, 3 .IV. 1999, leg. A. von Harten (MHNG, ZMUA); 1 9 , San'a, ca. 7900 ft., 10.- 
15.X. 1937. D. Carl Rathjens, B.M. 1939-396, from Lucerne (NHML). The holotype is 
glued laterally on a cardpoint, and is in good condition but both wings folded over each 
other. 

Etymology. The species is named for the small number of setae on the head. 

Diagnosis. Easy to recognize by the presence of only 1 frontal and 1 orbital seta, 
both short, fine and black (Fig. 42); ocellar seta short; postpedicel with distinct dorso- 
apical point (Fig. 44); wing entirely hyaline (Fig. 45); male: epandrium medially 
strongly convex (Fig. 49), epandrial process in lateral view hardly visible (Fig. 48); 
glans with few small papillae near tip of ductus (Fig. 50); female: aculeus with apical 
step (Fig. 53). 

Description. Wing length. Male: 2.05-2.30 mm; female: 2.20-2.60 mm. 

Head (Figs 42-43). Mat yellow, but ocellar triangle and occiput dorsal to occi- 
pital foramen ash grey microtrichose; head in profile 1.5-1.6 times as high as long; 
compound eye about 1.75 times as high as wide; gena about 1/6 as high as compound 
eye; only one weak frontal seta almost at level of antennal bases; 1 weak, reclinate 



FRUIT FLIES FROM SAUDI ARABIA 



1015 




Figs 42-46 
Arabodesis reductiseta sp. n. (paratype from Abha): 42, head, lateral view; 43, head, frontal 
view; 44, antenna; 45, thorax, dorsal view; 46, wing. 

orbital seta very high on head near vertex; all other setae also weak except for long 
medial vertical seta; postocular setae all white, short, in one row; pedicel without 
strong dorsal seta; postpedicel about twice as long as wide, with conspicuous dorso- 
apical point (Fig. 44). 

Thorax (Fig. 45). Scutum, pleura and subscutellum mostly ash grey micro- 
trichose; only postpronotal lobe, dorsal part of anepisternum, posterior half of noto- 
pleuron, most of prosternum and entire scutellum mat yellow; scutum covered with 
short, weak, white setulae; scutellum bare, but white setulose along margin; chaetotaxy 
as described for genus; halter yellow; both calypters yellow. 

Legs. Yellow, but hindfemur ventrally in middle with grey patch. 

Wing (Fig. 46). Entirely hyaline, only pterostigma slightly yellowish. 

Abdomen male (Figs. 47-50). Mat ash grey microtrichose; posterior margin of 
tergites with yellow transversal stripes, broader on syntergite 1+2 than on posterior 



1016 B. MERZ & H. A. DAWAH 

tergites; sternites yellow, each with a pair of grey spots (Fig. 47); sclerotization of glans 
compact (Fig. 50); acrophallus with two internal ducts, which are crossing in distal 
half; near tip with few small papillae on inner side; vesica small, apically broadly 
rounded. 

Abdomen female (Figs 51-54). Oviscape about as long as preceding 1-1.5 
tergites combined, about 0.5 mm long; orange on surface, black at both sides and along 
a medial longitudinal stripe: aculeus with slightly concave margin in basal half and 
abruptly narrowing in apical third, tip with apical step; length about 0.45 mm. 

Distribution. Saudi Arabia, Yemen. 

Host plants. Unknown. All specimens were collected in Malaise traps. 

Pest status. Not a pest species. 

Remarks. At first glance A. reductiseta resembles species of Hyalotephritis , 
which is the only genus of the Tephritis group with species having entirely hyaline 
wings and a similar head shape. Chaetotaxy (especially absence of medial postocellar 
setae), position of the dorsocentral seta, colour of posterior notopleural seta, and 
presence of 1-2 anteroventral setulae on hindfemur near tip indicate immediately that 
the new species does not belong to the Goniurellia clade in the Tephritis-group (see 
also above). A second species of Arabodesis is known from Yemen (Merz et al., in 
press). It differs from A. reductiseta by the stronger setae on the head, the presence of 
2 frontal setae, and the postpedicel being dorso-apically rounded. As only one female 
is available, which is slightly shrivelled, this species is not formally described here. 

Capitites augur (Frauenfeld, 1857) 

Material. 2: 26 6 . 

Distribution. Egypt, Israel. Sudan. Recorded from Saudi Arabia by Al-Ahmadi 
& Salem (1999, as Trupanea augur) and by Norrbom et al. (1999b, as Paradesis au- 
gur). It should be noted that this species was misidentified in the past, and its status was 
resolved by Freidberg & Kugler (1977). Therefore, it may be possible that old records 
refer to other species (Goniurellia spp.). 

Host plant. Pulicaria crispa (Asteraceae) (Freidberg & Kugler, 1989, as 
Dectodesis augur). 

Pest status. Not a pest species. 

Dectodesis auguralis (Bezzi, 1908) 

Material. 2:2 OS, 19. 

Distribution. Eastern Afrotropical Region from Eritrea to Zimbabwe, Yemen 
(Merz et al., in press). New for Saudi Arabia. 

Host plants. Unknown, but other species of the genus live in flowerheads of 
Erigeron, Helichrysum and Gnaphalium (Asteraceae). 

Pest status. Not a pest species. 

Dectodesis sp. 1 

Material. 2: 1 cT . 

Remarks. This male lacks the head and is also otherwise in bad condition. It is 
very similar to the previous species, but the apical scutellar setae are absent. Judging 
from wing pattern and colour of mesonotum this species may belong to D. confluens 



FRUIT FLIES FROM SAUDI ARABIA 



1017 




Figs 47-54 
Arabodesis reductiseta sp. n. (47-50, male; 51-54, female, paratypes from Abha): 47, abdominal 
sternites 4 and 5; 48, epandrium, lateral view; 49, same, caudal view; 50, glans; 51, oviscape; 
52, aculeus; 53, tip of aculeus; 54, spermatheca (only one illustrated). 



1018 B. MERZ & H.A. DAWAH 

(Wiedemann, 1830), a very widespread and abundant species in Eastern and Southern 
Africa. Because of the poor condition of the specimen we are not naming it here. 

Freidbergia mirabilis Merz, 1999 

Material. 2: 1 S . 

Distribution. Described from Kenya and Ethiopia (Merz, 1999). New for Saudi 
Arabia. 

Host plants. Pluchea spp. (Asteraceae) (Merz, 1999). 

Pest status. Not a pest species. 

Goniurellia persignata Freidberg, 1980 

Materiali: le?; 2: Id, 22 9. 

Distribution. From Sri Lanka and China in the East through Middle Asia and 
the Southern Mediterranean Region to Morocco. Also known from Ethiopia. New for 
Saudi Arabia. 

Host plant. Pulicaria arabica (Asteraceae) (Freidberg, 1980). 

Pest status. Not a pest species. 

Goniurellia spinifera Freidberg, 1980 

Material. 2: 1 6 , 1 2 . 

Distribution. Near East and North Eastern Afrotropical Region. Recorded from 
Saudi Arabia by Freidberg (1980). 

Host plant. Pulicaria desertorum (Asteraceae) (Freidberg, 1980). 

Pest status. Not a pest species. 

Hyalotephritis Freidberg, 1979 

This genus was proposed by Freidberg (1979) for two species which are 
morphologically very similar: H. complanata (Munro) and H '. planiscutellata (Becker). 
He provided a key and illustrated the main differences between the two species (head 
shape, structure of glans, aculeus tip). Larvae of both species live in flowerheads of 
Conyza dioscoridis (Asteraceae). The comparison of the 35 specimens from Saudi 
Arabia with numerous specimens of H '. planiscutellata from Israel, Kenya and Oman, 
and two pairs of H. complanata from South Africa showed that both species are present 
in Saudi Arabia. It was observed that the structure of the glans is the only reliable 
character separating the two species, whereas head and aculeus shape are more 
variable. The glans of the Saudi Arabian specimens of H. planiscutellata fits the illus- 
tration of Freidberg (1979), but a notable difference could be found in the sclerotiza- 
tion of the glans in all specimens of H. complanata (Fig. 55), including one male from 
South Africa, compared with the illustration of Freidberg (1979, Fig. 13): The distal 
part of the sclerotized tube is longer and stronger, undulating, ending in a small apical 
sclerite which is darker than the previous part. 

Hyalotephritis complanata (Munro, 1929) Fig. 55 

Material. 2: 2<?<?,32 2 ; 3: 7Jc?,62 2. 
Distribution. South and East Africa. New for Saudi Arabia. 
Host plant. Pluchea dioscoridis (Asteraceae) (Freidberg, 1979). 



FRUIT FLIES FROM SAUDI ARABIA 1019 

Pest status. Not a pest species. 

Remarks. For the structure of the glans of the aedeagus see above (Fig. 55). 

Hyalotephritis planiscutellata (Becker, 1903) 

Material. 3: \\6 6, 6? 9. 

Distribution. Near East south to Ethiopia, Yemen (Merz et al., in press). New 
for Saudi Arabia. 

Host plant. Pluchea dioscoridis (Asteraceae) (Freidberg & Kugler, 1989). 

Pest status. Not a pest species. 

Tephritomyia despoliata (Hering, 1956) Figs 56-58 

Acanthiophilus {Tephritomyia) despoliatus Hering, 1956: 88. Holotype 6: «IRAN (Chorassan), 
Birdjant, 23 .VII. -2. VIII. 1954, Richter & Schäuffele», «Acanthiophilus despoliatus m. 
Type, det. M. Hering 1955» (SMNS). Paratype 6, same data as holotype (SMNS) 
(examined). 

Material. 2: ISO, 10??. 

Distribution. So far only known from Iran. New for Saudi Arabia. 

Host plants. Unknown. All other species of the genus with known host plants 
live in flowerheads of Echinops spp. (Cardueae, Asteraceae) without causing galls. 

Pest status. Not a pest species. 

Remarks. Tephritomyia is a difficult Palaearctic and Afrotropical genus of 
morphologically similar species and is in need of revision. Based on the original 
description the specimens from Saudi Arabia resemble Acanthiophilus despoliatus 
Hering, which was described from 3 specimens from Iran (Hering, 1956). The compa- 
rison of the holotype and a paratype of the latter and the specimens from Saudi Arabia 
shows a general resemblance. The differences are: 

1. Wing length: The two specimens from Iran have longer wings (4.60-4.81 
mm, compared to 3.6-4.2 mm in Saudi Arabian specimens), but the specimens from 
Saudi Arabia were collected in alcohol and dry mounted, and have the wings usually 
slightly folded. The real length may be longer. 

2. Leg colour: The types from Iran have entirely yellow legs, whereas the colour 
of the Saudi Arabian population is more variable, with specimens having entirely 
yellow legs to others with black marks or a black ring in basal third of one to all 
femora. 

T. despoliatus is morphologically quite similar to the widely distributed Palae- 
arctic species Tephritomyia lauta (Loew) (compare head shape, Fig. 56 with Fig. 133 
in Freidberg & Kugler, 1989, and male terminalia, compare Fig. 58 with Fig. 24.12.C 
in Merz, 1999, for T. lauta). The two species differ by the much paler wing pattern 
without clearly delimited hyaline spots in T. despoliatus (Fig. 57). In T. lauta, the 
pattern is stronger with the hyaline spots more regular and entirely surrounded by the 
dark pattern (see Fig. 24.8 A in Merz, 1999). In general, T. despoliatus is much paler, 
with the scutellum either entirely yellow or only grey at base (usually entirely grey in 
T. lauta). The abdominal tergites have a more extensive yellow pattern (usually almost 
entirely grey in T. lauta), but intermediate specimens occur in both species. The length 
of the oviscape is more variable in T. lauta (as long as preceding 2.5-4 tergites 
combined, but at most as long as preceding 3 tergites combined in T. despoliatus). 



1020 



B. MERZ & H. A. DAWAH 




Figs 55-58 
Hyalotephritis complanata (Munro, 1929, specimen from Abha) (55) and Tephritomyia despo- 
liata (Hering, 1956, specimen from Abha) (56-58): 55, 58, male, glans; 56, head; 57, wing. 

Hering (1956) considered the shape of the postpedicel as the best discriminating cha- 
racter between T. despoliatus and T. lauta. The study of numerous specimens of T. lau- 
ta from Israel, Cypus and Crete shows, however, that there is no clearcut gap between 
the two species for this character. It seems that the shape of the postpedicel in the dif- 
ferent specimens is the result of the drying process. 

At the present state of knowledge we prefer to consider the two populations 
from Iran and Saudi Arabia as conspecific. Their status should be revised, however, as 
soon as more specimens become available, and/or biological data about the host plants 
are known. 



Trupanea amoena (Frauenfeld, 1857) 
Material. 2: 3SS 6,319 9. 

Distribution. Southern Palaearctic Region, East Africa, Oriental Region, 
Australia. Recorded from Saudi Arabia by Al-Ahmadi & Salem (1999). 

Host plants. This is one of the most polyphagous Tephritinae, attacking capitu- 
la of species of various Asteraceae {Achillea, Carthamus, Ethulia, Lactuca, Launaea, 
Leontodon, Picris, Sonchus and others) (Freidberg & Kugler, 1989; Merz, 1994; 
Munro, 1964). 



FRUIT FLIES FROM SAUDI ARABIA 1021 

Pest status. Recorded as minor pest on Calendula officinalis (Asteraceae) in 
India (White & Elson-Harris, 1992). 

Trupanea pseudoamoena Freidberg, 1974 

Material. 2:229. 

Distribution. Near East. Recorded from Saudi Arabia by Freidberg & Kugler 
(1989). 

Host plant. Pulicaria crispa (Asteraceae) (Freidberg & Kugler, 1989). 

Pest status. Not a pest species. 

Trupanea repleta Bezzi, 1918 Figs 59-69 

Trypanea aucta var. repleta Bezzi, 1918: 45. Holotype 6: [ERITREA] «Ghinda, Mochi, VI.16» 
(handwritten, white paper), «repleta» (in Bezzi's handwriting, white paper), «Holotypus 
6 , Trupanea aucta var repleta Bezzi 1918. desig. B. Merz 2004» (handwritten, red label) 
(MCSNM) (examined). 

Material. 2: 10ó*ó\ 209 9. Further specimens studied. CAPE VERDE 
ISLANDS: 3Ô6, S. Tiago, Lagoa, 15.11.1954, Lindberg; 40*0*, 1$, Sol, Terra Boa, 
21.1.1954, Lindberg; 1 6 , Maio, Pedro Vaz, 3.II.1954, Lindberg; 1 S , Maio, Morrinho, 
3. IL 1954, Lindberg; lo*, S. Vicente, Rib. Juliäo, 9. -11. III. 1954, Lindberg; lo\ 
Nicolau, Rib. Brava, 6.-19.XII.1953, Lindberg (all FMNH). YEMEN: see Merz et al. 
(in press). 

Diagnosis. Within Trupanea this is a typical species of the former Urelliosoma, 
with crescent hyaline apical wing pattern (Figs 60-63) and with a pale brown abdomen 
in both sexes. No sexual dimorphism in wing pattern; wing with narrow basal cross- 
band with an obtuse angle on level of M; this band often broken around M; small 
hyaline spots in r2+3 and r4+5 sometimes present; head with slightly produced 
vibrissal corner (Fig. 59); glans with typical sclerotized hook (Fig. 67). 

Redescription. Wing length. Male: 2.5-3.05 mm; female: 2.5-2.85 mm. 

Head (Fig. 59). Mat yellow, upper frons, ocellar triangle and occiput partly thin 
grey microtrichose; compound eye 1.3-1.5 times as high as long, gena 1/10 as high as 
eye; frons about as long as wide, anteriorly with few pale setulae; lunule rather high, 
conspicuous; face slightly concave; vibrissal corner projecting in profile slightly beyond 
base of antennae; antennae yellow, scape white setulose, pedicel with black setulae; 
postpedicel dorsally concave, dorso-apically slightly pointed; mouthparts yellow, capi- 
tate; palpus not projecting beyond vibrissal corner. Chaetotaxy: frontal setae, anterior 
orbital seta, ocellar seta and medial vertical seta pale brown, other setae yellow to whi- 
te; 3 frontal setae, 2 orbital setae, 1 ocellar seta, 1 medial and 1 lateral vertical seta, 1 
postocellar seta; row of postocular setae white; no medial postocellar seta. 

Thorax. Scutum, scutellum and pleura ash-grey microtrichose; ground colour 
of notopleura, postpronotum and pleura partly yellow; scutum covered with short, 
white setulae; all setae pale brown, only posterior notopleural seta white; dorsocentral 
seta almost at level of suture; chaetotaxy as usual in the genus. 

Legs. Yellow; foretarsus without modifications or modified setulae in both 
sexes. 

Wing (Figs 60-63). R4+5 bare on both sides; posterodistal extension of cell bcu 
indistinct; pattern brown, but basal crossband often paler, almost yellow; no bulla; 



1022 B. MERZ & H. A. DAWAH 

basal crossband narrow, stripe-like, sometimes interrupted on one or both sides of 
obtuse angle on level of M; both crossveins bordered by brown ray; hyaline spot at tip 
of R2+3 present or absent; small hyaline spots in dark patch of r2+3 and r4+5 some- 
times present (Fig. 61); hyaline spot in r4+5 just anterior to DM-Cu always present; at 
tip of R4+5 sometimes with small dark, isolated spot; brown ray in dm just distal to of 
R-M reaching Cu Al (Figs 62-63) or reduced (Fig. 61), rarely absent. No sexual 
dimorphism in wing pattern. Halter and calypter yellow. 

Abdomen male. Mat orange-brown, in middle rarely grey microtrichose. 
Sternite 5 (Fig. 64) concave distally, with 1 lateroapical seta; epandrium yellow, ovoid 
(Figs 65-66); prensisetae subequal, claw-shaped; glans (Fig. 67) with sclerotized hook 
dorsally and small sclerotization basally; vesica small. 

Abdomen female. Oviscape as long as last two tergites of preabdomen 
combined, black, at base with white setulae; aculeus as in Figs 68-69, apically rounded, 
without step. 

Distribution. Cape Verde Islands, Eritrea, Yemen. New for Saudi Arabia. 

Host plants. They are unknown for T. repleta. Taking into account the very 
limited host plant range of species of the group of species formerly placed in the genus 
Urelliosoma which is restricted to Launaea (Asteraceae, Cichorioideae) it is assumed 
that T. repleta attacks the same plant genus. According to Kilian (in litt., 17.V.2004) a 
possible host is Launaea intybacea, whose distribution matches the distribution of the 
fly. This plant forms a monophyletic species group with Launaea nudicaulis, which is 
the only known host plant of T. pulcherrima, probably the sister species of T. repleta 
(see below). 

Pest status. Not a pest species. 

Remarks. The specimens from Saudi Arabia are superficially very similar to 
specimens of Trupanea pulcherrima (Efflatoun, 1924) from Israel, but they key out as 
T. repleta in Munro (1964). The latter is a very poorly known species. The original 
description and illustration of Bezzi (1918) are insufficient and misleading because 
they are based on one male only and compared with an Indian species which belongs 
to another species group. Frey (1958) reports on the variability of the wing pattern in 
specimens from the Cape Verde Islands, and Munro (1964) who did not see the holo- 
type, did not add further characters. Judging from the illustration of the wing in Bezzi 
(1918) it seems reasonable to assume that T. repleta belongs to the group of species for- 
merly placed in Urelliosoma Hendel, 1927, a genus which was synonymized with 
Trupanea by Merz (1999) and which includes also T. pulcherrima. In order to clarify 
the status of the two taxa, the holotype of T. aucta var repleta Bezzi and the specimens 
mentioned by Frey (1958) were studied, as well as numerous specimens of T. pulcher- 
rima (see below). It is concluded here that both taxa represent distinct species which 
differ in morphology, distribution, and probably also in their biology. 

Trupanea pulcherrima (Efflatoun, 1924) 

Tephhtis pulcherrima Efflatoun, 1924: 100. Syntypes: 7c? c?, 29 9, EGYPT: Wadi Hoff; 7th 
Tour, Suez Road; Ezbet-el-Naghl (Plant Protection Department, Ministry of Agriculture, 
Dokki, Cairo, Egypt; Entomological Society of Egypt, Cairo; for details see Norrbom et 
al., 1999a) (not examined). 



FRUIT FLIES FROM SAUDI ARABIA 



1023 




Figs 59-63 
Trupanea repleta Bezzi, 1918: 59, head, lateral view; 60, wing male (holotype); 61 wing male 
(Abha); 62, wing female (Abha); 63, wing male (Abha). 



Material. ISRAEL: 26 6, 19, Hameshar, 16.VI.1986, A. Freidberg; 19, 
Mishor Paran, 70 km N Elat, 17.111.1995, Merz; 46 6, 29 9, Nahal Hiyyon, 
13.IV.1992, Merz & Freidberg; 1 6 , Nahal Zofar, 11.IV1992, Merz & Freidberg; 1 6 , 
1 ? , Dead Sea, Qalya (= Kallia), 7.VI.1996, Merz & Freidberg (all MHNG). TUNISIA: 
1 9 , 24 km S Gabès, 33.42N/10.00E, 8.VI.2000, Schmid-Egger; 1 9 , 5 km W Douz, 
33.29N/8.59E, 21. in .2001, Schmid-Egger (both MHNG). 

Distribution. Iran, Tunisia, Egypt, and Israel. Recorded from Saudi Arabia by 
Freidberg & Kugler (1989). This last record should be revised because of its confusion 
with T. repleta. 

Host plants. Launaea nudicaulis (Asteraceae) (Freidberg & Kugler, 1989). The 
distribution of the plant corresponds perfectly with the known distribution of T. pul- 
cherrima. 

Remarks. The descriptions of Efflatoun (1924), Hendel (1927) and Freidberg & 
Kugler (1989) are very comprehensive and the species is therefore not redescribed 
here. Colouration and structure of head, thorax, abdomen and terminalia of both sexes 
are the same for T. pulcherrima and T. repleta. Differences between the two species 
concern the wing pattern. In T. repleta the basal dark crossband (from the tip of Sc to 
the tip of Al+CuA2) is narrow, often paler than the remaining pattern, and in some 



1024 



B. MERZ & H. A. DAWAH 



64 



•v 









Figs 64-69 
Trupanea repleta Bezzi, 1918 (64-67, male; 68-69, female, specimens from Abha): 64, abdo- 
minal sternites 4 and 5; 65, epandrium, lateral view; 66, same, caudal view; 67, glans; 68, 
aculeus; 69, tip of aculeus. 



specimens broken in the middle. Usually this band is oblique and directed towards R- 
M until M, forming a very obtuse angle and parallel to DM-Cu until wing margin. In 
T. pulcherrima this basal crossband is much wider and its basal border is almost 
straight from C to the hind margin of the wing without distinct angle at level of M. 
Moreover the enitre pattern is uniformly brown, or only slightly paler in the stigma. 

Trupanea sp. 1 

Material. 2: 3<?d,299. 

Remarks. Trupanea is one of the largest genera of Tephritidae and is known 
from all major biogeographical regions, with 40 species known from the Afrotropical 
Region (Munro, 1964) and about 12 species from the Palaearctic Region (Norrbom et 
al., 1999b). The 5 specimens belong probably to a species close to the T. stellata sub- 
group as defined by Munro (1964). However, they do not fit the description of any 
species in this monograph nor any other description available to us. Taking into 
account the large size of the genus, the small differences between species and the 
unknown variability in many species we are not describing this species here. 



FRUIT FLIES FROM SAUDI ARABIA 1025 

Trupanodesis sp. 1 

Material. 2: 1 S . 

Remarks. The genus was proposed by Merz (1999) for Trupanea aurea Bezzi, 
1924 (type species) and an undescribed species from Kenya based on a cladistic ana- 
lysis. The specimen from Saudi Arabia may be the same species as the undescribed 
species from Kenya. However, as only one specimen is available, this identification is 
tentative. 

Distribution and host plants. This is an Afrotropical genus with both species 
assigned to it living in flowerheads of Vernonia (Asteraceae, Vernonieae) (Merz, 1999). 

DISCUSSION 

In the course of a survey of the insect fauna of the Aseer (= ' Asir ) area in south- 
western Saudi Arabia 51 species of Tephritidae were found, of which 42 species could 
be named (Tab. 1). Six of them are new to science, and 9 species could not be identi- 
fied safely to species level. Previously, 22 species were recorded from this country, 
and the presence of half of them could be confirmed in this study. For 1 1 species we 
could not find specimens in the Malaise traps during this survey. In total, 62 species of 
Tephritidae are currently known to occur in Saudi Arabia. This diversity matches well 
with other countries of the Near and Middle East except for Israel which is much better 
known through the collecting efforts of A. Freidberg in the last 35 years in all parts of 
the country (Freidberg & Kugler, 1989). No comparative effort has been conducted for 
other countries in the region, and it is thus likely that their real species number is much 
higher. For instance, 17 species recorded from the southernmost part of Israel and the 
Sinai mountains in Egypt have not yet been found in Saudi Arabia, but most of them 
should also occur in the mountains next to the Gulf of Aqaba in the northwestern part 
of Saudi Arabia. Examples are Capparimyia savastani (Martell), Euarestella kugleri 
Freidberg and E. pninae Freidberg, Goniurellia lacerata (Becker) and G. longicauda 
Freidberg, Notomma mutilum (Bezzi) and Trupanea desertorum (Efflatoun). 

The present study is interesting from a biogeographical point of view: Over two 
thirds of the species of the Aseer area are of Afrotropical origin. This means that these 
species have a predominantly Afrotropical distribution, or most species of the genus 
are Afrotropical. Four species are typical for desert areas in the Near and Middle East 
(e. g., Metasphenisca negeviana, Trupanea pseudoamoena) , and only one species, 
Ensina sonchi, is a typical Palaearctic species. Four species are widespread in the Old 
World (e. g., Dioxyna sororcula, Spathulina acroleuca), and one species, Bactrocera 
zonata, is a recent introduction from the Oriental Region. On the other hand, typical 
Oriental species are not present in the Aseer area nor elsewhere in Saudi Arabia. 
Interestingly enough, some genera and tribes which are widespread in the Palaearctic 
Region are not yet known in Saudi Arabia, such as Terelliini, Myopitini, or the large 
genus Tephritis. They are all known from desert localities in Israel, Jordan and Syria 
(Freidberg & Kugler, 1989; Korneyev & Dirlbek 2000), and their occurrence in the 
North of Saudi Arabia may be expected. In contrast, Israel has a higher proportion of 
Palaearctic species, with only about 20% of Afrotropical elements (Freidberg, 1988). 

Although a remarkable diversity of Tephritidae was collected by a Malaise trap 
it is generally known that traps are not the best collecting method for Tephritidae. 



1026 B. MERZ & H.A. DAWAH 

Sweeping over potential host plants and rearing Tephritids from infested plants yield 
usually a much higher number of species. Consequently, it may be assumed that the 
real number of species in Saudi Arabia may be doubled by using better collecting 
techniques and by collecting in the northern, palaearctic part of the country. 

It can be seen from Tab. 1 that 10 species recorded from Saudi Arabia are pest 
species according to White & Elson-Harris (1992) and require special attention in fruit 
plantations. Only 4 of these species were found in the present survey: Carpomyia 
incompleta, Bactrocera zonata, Dacus frontalis, and D. vertebratus , and all of them 
only in small number of specimens. No new pest species was discovered. But other 
pest species attacking Cucurbitaceae, such as Bactrocera Cucurbitae or Dacus ciliatus, 
should also occur in the Aseer area. Probably this is again a question of the collecting 
method. Monitoring those species needs specific collecting methods (e. g., traps with 
cue lure or methyl eugenol for Dacini; see Munro, 1984). 

ACKNOWLEDGEMENTS 

Our sincerest thanks go to John Deeming (Cardiff, Wales) who initiated this 
study, established the contact between the authors, arranged the loans of specimens to 
BM, gave important information, and reviewed the manuscript. The generous help of 
Amnon Freidberg (Tel- Aviv, Israel) is thankfully acknowledged. He was loaning and 
donating numerous specimens, helped us with fruitful discussions and was reviewing 
an earlier version of the paper. For help with identifications and useful comments we 
are indebted to Marc De Meyer (Tervuren, Belgium), David Hancock (Cairns, 
Australia), and Ian White (London, United Kingdom). For loan of specimens we 
express our sincerest thanks to Fabrizio Rigato (Milano, Italy), Gunilla Stahls-Mäkäla 
(Helsinki, Finnland) and Hans-Peter Tschorsnig (Stuttgart, Germany). A special thank 
is sent to Norbert Kilian (Botanical Garden and Botanical Museum, Free University, 
Berlin, Germany) for information about the plant genus Launaea. Finally, we would 
like to thank Florence Marteau (Geneva) for technical help. 

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944 pp. 



Revue suisse de Zoologie 112 (4): 1029-1044; décembre 2005 



Proxiandrena subgen. nov. und Re vision der west- und 
zentralpaläarktischen Arten der Andrena proxima-Gruppe 
(Hymenoptera, Apidae) 

Christian SCHMID-EGGER 

Kirchstr. 1,D-82211 Herrsching, Deutschland. E-mail: christian@schmid-egger.de 

Proxiandrena subgen. nov. and revision of the Andrena proxima group 
in the western and central Palaearctic Region (Hymenoptera, Apidae). - 

The former Andrena proxima group is revised for the western and central 
Palaearctic Region. Proxiandrena subgen. nov. is described for the here 
treated species with Andrena proxima as type species. Proxiandrena 
includes 5 valid species, which had been treated as subspecies of Andrena 
proxima s. 1. before: Andrena ampia Warncke in Morocco, Spain, southern 
France, the western Alpes, and - separated from this area - in Central Asia 
north of Turkmenistan; Andrena alutacea Stoeckhert with a range from 
southeast Germany and eastern Switzerland to the eastern Mediterranean 
area and Turkmenistan; Andrena aspericollis Perez from Algeria; Andrena 
bermela Warncke from Turkey; Andrena proxima (Kirby) from Central 
Europe and Italy. A key to the species and diagnostic characters are given 
and distribution of the species is discussed. The female of A. bernicla is 
unknown and its status as a species remains doubtfull. 

Keywords: Apidae - Andrena proxima - new subgenus - taxonomy - 
revision - faunistics - systematics. 

EINLEITUNG 

Andrena proxima (Kirby, 1802) ist eine in Mitteleuropa weitverbreitete und 
stellenweise häufige Sandbienenart. Stoeckhert (1942) beschrieb die sehr ähnliche alu- 
tacea aus Mittel- und Südosteuropa. Neben morphologischen Unterschieden berief 
sich Stoeckhert dabei hauptsächlich auf die unterschiedliche Flugzeit der beiden Arten. 
Ihr Status ist seither jedoch umstritten, da alutacea in der Folge von verschiedenen 
Autoren nicht anerkannt wurde (Warncke, verschiedene Arbeiten; Dylewska 1987, 
Gusenleitner & Schwarz 2002). Erst Schmid-Egger & Scheuchl (1997) führten die Art 
wieder in ihrem Bestimmungsschlüssel der mitteleuropäischen Andrena- Arten auf. Die 
Autoren wiesen außerdem auf ein drittes Taxon der Artengruppe aus den Westalpen 
hin, welches sie ebenfalls für eine eigenständige Art hielten. 

In Südeuropa sind drei weitere Formen aus der proxim a-Gruppe beschrieben, 
nämlich proxima ssp. bernicla Warncke, 1975 aus der Türkei, proxima ssp. ampia 
Warncke, 1967 aus Spanien und proxima ssp. aspericollis Perez, 1895 aus Algerien. 
Eine weitere Form, die von Warncke als Unterart ausgezeichnet ist, aber nicht 
beschrieben wurde, befindet sich in seiner Sammlung im Oberösterreichischen Landes- 
museum Linz. 



Manuskript angenommen am 02.03.2005 



1030 C- SCHMID-EGGER 

Ziel der vorliegenden Revision ist es, die genannten Formen zu untersuchen und 
ihren Status abzuklären. So war insbesondere die Frage zu klären, ob die beschriebe- 
nen Taxa Unterarten, Arten oder infrasubspezifische Formen sind. Gleichzeitig erwies 
es sich als notwendig, für die Artengruppe um Andrena proximo, eine neue 
Untergattung zu beschreiben, da die bisherige Lösung, die Arten in die Untergattung 
Micrandrena Ashmead, 1988 zu stellen, mehr als unbefriedigend ist. Die Klärung der 
verwandschaftlichen Beziehungen innerhalb der untersuchten Artengruppe war nicht 
möglich, da die sichtbaren Unterscheidungsmerkmale dafür nicht ausreichend sind. 

UNTERSUCHTES MATERIAL 

Das untersuchte Material stammt aus den folgenden Sammlungen (mit den im 
Text verwendeten Abkürzungen): 

ETH Sammlung der Eidgenössischen Technischen Hochschule in Zürich, 

Schweiz 
GRÜ coli Grünwaldt, in der Zoologischen Staatssammlung München, 

Deutschland 
Kraus Privatsammlung Dr. M. Kraus, Nürnberg, Deutschland 
MHNG Muséum d'histoire naturelle, Genève, Schweiz 
OLL coli. Öberösterreichisches Landesmuseum Linz (vor allem coli. Warncke), 

Österreich 
ORT Privatsammlung Sabine Oertli, Zürich, Schweiz 

Scheuchl Privatsammlung Erwin Scheuchl, Velden, Deutschland 
SE Privatsammlung Dr. Christian Schmid-Egger, Herrsching, Deutschland 

ZSM Zoologische Staatssammlung München, Deutschland 

Für die Schweiz werden die Kantone gemäss den Autokennzeichen folgender- 
massen abgekürzt: BE = Bern; GR = Graubünden; TI = Tessin; VS = Wallis. 

ERGEBNISSE 

PROXI ANDREN A SUBGEN. N. 

Typus art. Melitta proxima Kirby, 1802 (= Andrena proximo). 

Diagnose und Beschreibung: Diagnostische Merkmale sind hervorgehoben: 

Weibchen: Behaarung und Körpergröße: 9-12 mm. Körperfarbe schwarz, 
Tarsen manchmal rötlich aufgehellt. Körperbehaarung grau, weißlich oder gelblich, 
Tergitendbinden schneeweiß, grau oder schwach gelblich. Kopf und Thorax struppig 
abstehend behaart, Haare im Mittel doppelt so lang wie Scapus. 

Kopf: Labrum doppelt so breit wie lang, unten seicht ausgerandet. Clypeus auf 
gesamten Oberfläche dicht und grob quer gerunzelt. Gesicht zwischen Ocellen und 
Clypeus längs gerunzelt, bzw. wabenartig skulpturiert. Foveae schmal, auf Höhe der 
Fühlereinlenkung etwa 0,28 x so breit wie halbe Gesichtsbreite (Gesichtsbreite ohne 
Auge gemessen). Fühlerglied IV halb so lang wie Fühlerglied IQ. 

Thorax: Mesonotum im vorderen Teil quer runzlig, auf der übrigen Oberfläche 
grob und dicht punktiert (Punktabstand = 0,5-1 Punkdurchmesser), Punktzwischen- 
räume glänzend. Mesopleuren mit grob wabenartiger Skulptur. Oberseite des Prodo- 



PROXIANDRENA SUBGEN. NOV. 1031 

deum grob netzartig skulpturiert, das dreieckige Mittelfeld ist mehr oder weniger 
deutlich mit einer feinen Linie von den ansonsten gleich skulpturierten Seitenfeldern 
abgegrenzt. Propodeumseitenf eider bedeckt mit sternförmig zusammenlaufenden 
Runzeln, die in der Mitte ein Haar tragen. Oberkante und Hinterkante der Seitenfelder 
mit mittellanger Sammelfranse, die Haare sind kaum gebogen. Hintertibia schlank, mit 
grauer Sammelfranse. Die Haare auf der Außenseite sind etwa doppelt so lang wie der 
Durchmesser der Tibia. Flügeladerung dunkelbraun, Flügel leicht graubraun getönt. 

Abdomen: Tergite spiegelglatt oder fein chagriniert, matt oder glänzend, manch- 
mal mit öligem Schein. Tergite II-IV mit Endbinden, die durchgehend oder breit unter- 
brochen sind. Endfranse gelblich oder graubraun, von weißen oder gelblichen längeren 
Haaren überdeckt. Sternitscheiben grob skulpturiert und punktiert, Sternitdepressionen 
fein skulpturiert. Sternite II-IV mit abstehenden Endbinden (Winkel von ca. 40 Grad). 

Männchen: Behaarung und Körpergröße: 8-11 mm, ansonsten wie Weibchen. 

Kopf: Labrum quer rechteckig, unten seicht ausgerandet. Clypeus dicht und 
grob quer gerunzelt. Der übrige Kopf grob gerunzelt. Fühlergeißel dunkel oder unten 
rötlich aufgehellt. Fühlerglied III etwa 1 ,3x so lang wie Fühlerglied IV. 

Thorax: Mesonotum vollständig gerunzelt oder punktiert und mit glänzenden 
Punktzwischenräumen. Mesopleuren mit grob wabenartiger Skulptur, Propodeum mit 
sternförmigen Runzeln (ähnlich wie beim Weibchen). 

Abdomen: Tergite glänzend, punktiert oder mit feiner Runzelung. Tergite bis auf 
eine Art mit grauweißen oder gelblichen Endbinden, die unterbrochen oder durch- 
gehend sind. Sternite mit undeutlichen Endfransen, Sternit VIII schmal, am Ende 
verbreitert, seitlich mit langen Haaren, die länger als der Durchmesser des Sternites 
sind und sich zum Ende zu verkürzen. Genital. Genital von einfachem Bau, Penisvalve 
sehr schmal, Gonostylus schmal, ohne Besonderheiten. 

Diskussion: Andrena proximo wurde von Stoeckhert (1942) in die minutula- 
Gruppe und von Warncke (1968) in die entsprechende Untergattung Micrandrena 
Ashmead, 1899 gestellt. Dylewska (1987) schloß sich dieser Ansicht nicht an, sondern 
plazierte proximo aufgrund der besonderen Skulptur der Mesopleuren in eine eigene 
Gruppe, die Andrena proxima-Gruppe. Dieser Ansicht schließen sich auch Dubitzky & 
Schönitzer (200 1 ) an und berufen sich dabei vor allem auf die sternförmig zusammen- 
laufenden Runzeln der Propodeumseiten. Dieses Merkmal betrachten sie als Autapo- 
morphie der Artengruppe . Eine nähere Verwandtschaft sehen die Autoren vor allem zur 
labialis Gruppe (subg. Holandrena Perez 1890). Da sich meine Kenntnisse der Gattung 
Andrena lediglich auf die mitteleuropäischen Arten beziehen, kann ich diesen Fall 
nicht selbst abschliessend beurteilen, sondern schließe mich dieser Meinung an. 

Ich stimme mit Dylweska (1987) und Dubitzky & Schönitzer (2001) darin über- 
ein, daß proximo und ihre Verwandten eine eigene monophyletische Artengruppe 
innerhalb der Gattung Andrena bilden. Ihre Stellung rechtfertigt die Beschreibung 
einer eigenen Untergattung: Proxiandrena subgen. n. Stoeckherts und Warnckes An- 
sicht, daß proximo zur Untergattung Micrandrena gehört, kann ich nicht folgen. Die 
Übereinstimmung ist eher oberflächlicher Natur (schwarze Arten mit grauer Körper- 
behaarung und teilweise glänzenden, bzw. punktierten Körperoberflächen, weiße Ter- 
gitendbinden). Zwei wesentliche Merkmale von Micrandrena, die geringe Körper- 
größe (< 8 mm) und die dem Stigma stark angenährte Cubitalader I finden sich bei 



1032 C. SCHMID-EGGER 

proxima nicht. Auf der anderen Seite sind die beiden charakteristischen Merkmale von 
proxima, die typische Skulptur der Mesopleuren und des Propodeums, nirgends bei 
Micrandrena ausgeprägt. 

Bestimmungsschlüssel zu den Andrena- Arten der Untergattung Proxiandrena 

Weibchen 

1 Tergite ohne weiße Endbinden. Verbreitung Algerien aspericollis Perez 

Tergite mit deutlichen weißen, grauen oder gelblichen Endbinden. Ver- 
breitung: Marokko, Europa, Asien 2 

2 Mindestens Scheiben und Depressionen der Tergite II-III, für Tiere aus 
Mitteleuropa auch von Tergit IV gleichmäßig gerunzelt und chagriniert, 
ohne Glanz. Tergite unpunktiert oder bei manchen Tieren Tergite II-IV 
seitlich punktiert. Tergitendbinde IV durchgehend (nur bei frischen 
Tieren sichtbar!). Thorax (in lateraler Sicht) mit einer kurzen schwarzen 
Unterbehaarung zwischen den langen hellen Haaren. Von Süd- 
deutschland, Ostschweiz, Südosteuropa, Türkei bis Zentralasien. In der 
Türkei auch bernicla, die im weiblichen Geschlecht nicht von alutacea 

zu trennen ist alutacea Stoeckhert 

Mindestens die Tergitbeulen und Teile der hinteren Tergite glänzend. 
Tergite häufig punktiert, mindestens auf Tergit III und IV seitlich. 
Tergitendbinde IV bei proxima auch bei frischen Tieren in der Mitte 
breit unterbrochen, bei ampia können die Tergitbinden durchgehend 
sein. Mit oder ohne schwarze Unterbehaarung auf dem Thorax 3 

3 Tergitscheiben und Tergitdepressionen fein gerunzelt und mehr oder we- 
niger glänzend, zum Teil mit öligem Schein. Tergitscheibe II-IV ohne 
deutliche Punktierung. Tergitbeulen spiegelglatt, glänzend. Thorax (in 
lateraler Sicht) meist ohne kurze schwarze Unterbehaarung, manchmal 
ist diese schwach ausgebildet. Dreieckiges Mittelfeld des Propodeum 
undeutlich zu den Seitenfeldern abgegrenzt. Binden gelblich. Kleiner, 
9-10 mm. Verbreitung: Europa außer Iberische Halbinsel, fehlt an- 
scheinend in den zentralen und westlichen Alpentälern proxima (Kirby) 

Tergitdepression spiegelglatt, Tergitscheiben II-IV mit deutlicher 
Punktierung, die der Tergite III und IV gröber als die von Tergit II. 
Tergitscheiben zwischen den Punkten ebenfalls spiegelglatt, im basalen 

Teil mit schwachen Runzeln. Thorax (in lateraler Sicht) zwischen der 
langen hellen Behaarung mit dichter und kurzer schwarzer Unterbe- 
haarung. Dreieckiges Mittelfeld des Propodeum bei den meisten Tieren 
mit deutlichen, aber feinem Kiel zu den Seitenfeldern zu abgegrenzt. 
Binden schneeweiß. Größer, 10-12 mm. Zentrale und westliche Alpen: 
Aostatal, Südschweiz, Durancetal, Iberische Halbinsel (Verbreitung in 
Südfrankreich noch unklar), Zentralasien ampia Warncke 

Männchen 

1 Tergite ohne weiße Endbinden. Mesonotum auf der ganzen Fläche grob 

gerunzelt, dazwischen chagriniert. Verbreitung: Algerien .... aspericollis Perez 



PR0X1ANDRENA SUBGEN. NOV. 1033 

Tergite mit deutlichen weißen, grauen oder gelblichen Endbinden. Tiere 
aus Marokko: Mesonotum mit glänzenden Punktzwischenräumen. Ver- 
breitung: Marokko, Europa, Asien 2 

2 Gesicht dunkelbraun behaart. Türkei bernicla Warncke 

Gesicht hell behaart 3 

3 Mesonotum vollständig chagriniert und wabenartig gerunzelt, keine ein- 
gestochenen Punkte oder glänzende Punktzwischenräume erkennbar. 
Tergite I-IV: Basis fein gerunzelt, Mittelteil glänzend mit zerstreuten 
Mikropunkten, Depression fast punktfrei, im apikalen Teil sehr feine 
Mikropunkte. Tergitendbinden II-IV in der Mitte breit unterbrochen. 
Körperbehaarung gelblich. Europa außer Iberische Halbinsel, Zentral- 
asien, fehlt anscheinend in den zentralen und westlichen Alpentälern 

proxima (Kirby) 

Mesonotum mit glänzenden Punktzwischenräumen, die im hinteren Teil 

0,5-1 Punktdurchmesser betragen (bei ampia aus der Schweiz vereinzelt 
stärker chagriniert). Tergite I-IV: Basis und Mittelteil glänzend mit deut- 
lich eingestochenen Punkten, die im direkten Vergleich mindestens dop- 
pelt so groß wie bei proxima sind. Depression unterschiedlich. Ter- 
gitendbinden unterschiedlich. Körperbehaarung weißlich. 
Die beiden folgenden Arten sind sehr ähnlich und ohne zugehörige 
Weibchen nur aufgrund ihrer geografischen Verbreitung sicher zu trennen ... 4 

4 Verbreitung: Süddeutschland, Ostschweiz, Südosteuropa, Türkei. 
Tergite im Mittel weniger dicht punktiert, Punkte im direkten Vergleich 
kleiner als bei ampia. Tergitendbinden III und VI bei frischen Tieren 

durchgehend alutacea Stoeckhert 

Verbreitung: Zentrale und westliche Alpen: Aostatal, Südschweiz, 
Durancetal, Spanien (Verbreitung in Südfrankreich noch unklar, in der 
Schweiz Überschneidung mit dem Areal von alutacea), Zentralasien. 
Tergitendbinden in der Mitte breit unterbrochen, selten Binde IV durch- 
gehend (bei asiatischen Tieren) ampia Warncke 

Vergleichende Beschreibung der Andrena- Arten der Untergattung Proxiandrena 

Nachfolgend werden die für die Artunterscheidung wichtigen Merkmale be- 
schrieben und die Arten dabei gegenübergestellt. Bei der nachfolgenden Besprechung 
der Arten wird lediglich eine kurze Diagnose der Arten gegeben. 

Behaarung des Mesonotum, Weibchen 

alutacea Weißlich, in der Länge unregelmäßig, in der Tendenz etwas kürzer 

als bei proxima. Auf der ganzen Länge des Tergites mit feiner, kurzer 
dunkler Unterbehaarung. 

ampia Gelblich, bei spanischen Tieren auch weißlich, die übrigen Merk- 

male wie bei alutacea. 

aspericollis Gelblich, lang, ähnlich proxima. 

proxima Gelblich, in der Länge gleichmäßig, im Mittel etwas länger als bei 

alutacea und ampia. 



1034 



C. SCHMID-EGGER 



Tergitskulptur, Weibchen 

alutacea I Mitteleuropa Tergite gleichmäßig fein gerunzelt (chagriniert), matt. Tergitbeulen 
leicht glänzend, ebenfalls gerunzelt. Glanz nimmt auf Tergit IV 
leicht zu. Auf den Tergitscheiben undeutliche Punkte erkennbar. 
Tergitdepressionen in der Tendenz schmaler und weniger stark bo- 
genförmig als bei proximo., außerdem wirkt der Niveauunterschied 
zwischen Scheibe und Depression niedriger als bei proxima. 

alutacea I Türkei Die feine Runzelung der Tergite verschwindet nahezu, Tergite wir- 

ken glatt. Scheiben und Tergitdepressionen mit identischer Skulptur. 
Auf den Tergiten I-II ist noch eine feine Chagrinierung erkennbar, 
die auf den Endtergiten nahezu verschwindet. Tergitscheiben II und 
IV sehr schwach, klein und zerstreut punktiert. 

ampia Tergite spiegelglatt und stark glänzend, auf den Tergitscheiben II und 

III leichte Chagrinierung erkennbar. Tergitscheiben II-IV mit deut- 
lich eingestochener und teilweise grober Punktierung, Punkte im 
Mittel 2-3 Punktdurchmesser voneinander entfernt. Bei den spani- 
schen Tieren kann die Punktierung weniger stark und die Chagri- 
nierung stärker ausgeprägt sein. An der Tergitpunktierung sind 
solche Tiere aber immer von proxima und alutacea zu unterscheiden. 

aspericollis Tergitskulptur entspricht der von alutacea. 

proxima Tergite chagriniert, mehr oder weniger stark glänzend, besonders auf 

den hinteren Tergiten. Die Tergitdepressionen stärker glänzend als 
die Scheiben, Depression IV manchmal spiegelglatt. Tergitbeulen 
fast immer spiegelglatt. 



Tergitendbinden auf den Tergiten II-IV, Weibchen 

alutacea Schneeweiß, anliegend. Sie entspringen etwas oberhalb des 

Tergitendrandes, so dass sie den Tergitendrand um ihre halbe Länge 
oder 2/3 ihrer Länge überragen. Binde auf Tergit IV durchgehend, 
auf Tergit II und III breit unterbrochen. Unterbrechung auf Tergit III 
entspricht etwa der Breite der Binde (Breite: vom Tergitseitenrand 
Richtung Tergitmitte), auf Tergit II etwa der doppelten Bindenbreite. 

ampia Schneeweiss, etwa im Winkel von 30° abstehend (bei den asiatischen 

Tieren anliegend, und nach hinten gerichtet). Die Binde entspringt 
etwas tiefer als bei alutacea, aber weiter zur Tergitbasis als bei pro- 
xima. Binde auf Tergit IV schmale (Ozellenbreit) bis breiter (Länge 
des Fühlergliedes III) unterbrochen, die anderen Binden breit unter- 
brochen. Bei asiatischen Tieren Binde auf Tergit IV durchgehend. 



aspericollis 
proxima 



Keine Binden vorhanden. 

Gelblich, etwa im Winkel von 30 abstehend. Die Binde entspringt 
fast unmittelbar am Tergitendrand. Binde IV breit unterbrochen 
(etwa um Bindenbreite), die übrigen Binden sehr kurz (etwa ein 
Viertel der Tergitbreite). 



Skulptur des Mesonotum, Männchen 

alutacea Deutlich eingestochen punktiert, mit glänzenden Punktzwischen- 

räumen, die 0,2-1 Punktdurchmesser betragen. Basal zum Teil auch 
chasriniert, bzw. ohne erkennbare Punkte. 



ampia 



Wie alutacea, allerdings mit großer Variationsbreite. Tiere aus dem 
Aostatal sind sehr deutlich punktiert mit glänzenden Zwischen- 
räumen, bei Tieren aus der Schweiz ist das Mesonotum manchmal 
vollständig chagriniert, ohne erkennbare Punkte. 



PROX/ANDRENA SUBGEN. NOV. 



1035 



aspericollis 

hemic la 
proxima 



Chagriniert, mit körnigen Erhabenheiten oder Runzeln, keine Punkte 
oder glänzende Flächen erkennbar. 

Wie aspericollis . 

Wie aspericollis. 



Skulptur der Tergite II-IV (Punktzwischenräume bei allen Arten glänzend), Männchen 

alutacea Tergitscheiben fein eingestochen punktiert, Punktdurchmesser ca. 

0,3-0,5 x wie Punktdurchmesser auf Mesonotum, Punktabstand 1-2 
Punktdurchmesser. Tergitbasis fein quer chagriniert. Tergitdepres- 
sion mit zerstreuten, sehr feinen Punkten, die vordere Hälfte der 
Tergitdepression teilweise punktfrei. 

ampia Wie alutacea, Punktdurchmesser im Mittel größer (ca. 0,5-0,8 x 

Punktdurchmesser auf Mesonotum), meist bis zur Tergitbasis glän- 
zend. 



aspericollis 
bernicla 

proxima 



Tergitscheibe mit sehr undeutlichen und kleinen Punkten, basal fein 
quergerunzelt. Tergitdepression nahezu glatt. 

Tergitscheiben mit feinen Punkten (in der Größe ähnlich alutacea), 
Punktabstände 1-3 Punktdurchmesser, Tergitdepression mit sehr 
wenigen zerstreuten Punkten. 

Tergitscheiben mit sehr feinen und zerstreuten Punkten, Punkt- 
abstände 2-5 Punktdurchmesser, Tergitdepressionen punktlos oder 
mit vereinzelten sehr feinen Punkten. 



Endbinden Tergite II-V Männchen (kann nur bei frischen Tieren beurteilt werden) 

alutacea Tergit III und IV mit (fast) durchgehender Endbinde. Tergit V mit 

lockerer Endbinde, die aus langen Haaren besteht. Binden schnee- 
weiß. 



ampia 

aspericollis 

bernicla 
proxima 



Körpergröße 

alutacea 

ampia 

aspericollis 

bernicla 

proxima 

Verbreitung 

alutacea 

ampia 



Tergite II-IV mit breit unterbrochenen Binden, Binde auf Tergit IV 
kann jedoch auch durchgehend sein (bei zentralasiatischen Tieren). 
Tergit V mit undeutlicher durchgehender Endbinde. Binden schnee- 
weiß, dichter als bei proxima. 

Die beiden untersuchten Männchen ohne erkennbare Binden. Dies 
mag jedoch vielleicht auf den Erhaltungszustand der Tiere zurück- 
zuführen sein. 

Wie bei proxima, Binden jedoch weiß. 

Tergitendbinden II-IV immer breit unterbrochen (mindestens auf 1/3 
der Tergitbreite). Tergit V apikal ohne erkennbare Endbinde. Binden 
gelblich. 



Weibchen 

10-11 mm 
10-12 mm 
11-12 mm 

9-10 mm 



Männchen 

9-11 mm 
9,5-11 mm 
8 mm 
8-9 mm 
8-10 mm 



südöstliches Deutschland, Ostschweiz bis Südosteuropa, Lettland. 
Türkei, Iran und Turkmenistan. 

Marokko, Iberische Halbinsel, Frankreich, Südfrankreich. 
Durancetal; Südschweiz, Italien: Aostatal, Aserbeidschan bis 
Zentralasien. 



1036 C. SCHMID-EGGER 

aspericollis Algerien. 

bermela Türkei (nördliche Hälfte?). 

proximo. Mitteleuropa, Großbritannien, Italien, Südosteuropa bis Griechen- 

land. 

Andrena alutacea Stoeckhert, 1942 

Andrena alutacea Stoeckhert, 1942: 236-252 (Erstbeschreibung 3 und 9 , Holotypus 9 , 
Typusfundort: Eining/Donau, ZSM, untersucht) - Pittioni & Schmidt, 1943: 33 (Fauna 
von Österreich, Bulgarien).- Kocourek, 1966: 69 (Fauna Tschechoslowakei, nur in 
Mähren) - Schmid-Egger & Scheuchl, 1997 67-68, 142 (Schlüssel für Mitteleuropa, 
Angaben zur Verbreitung und Phänologie) - Mandery, 2001: 116 (Fauna für Franken, 
aktuelle Funde) . 

Andrena proximo: Warncke, 1967: 209, 250 (synonym voit proximo) - Warncke, 1986: 45 (syn- 
onym mit proximo) - Westrich, 1984: 23 (synonym mit proximo, mit Paratypus ver- 
glichen) - Dylewska, 1987: 547 (synonym mit proximo) - Schwarz et al., 1996 50 
(synonym mit proximo: ,A. alutacea ist entgegen bisweilen geäußerter Vermutungen 
keine eigene Art") - Gusenleitner & Schwarz, 2002: 610-611 (Literaturzitate, zu pro- 
ximo gestellt, bzw. als forma von dieser behandelt). 

Untersuchtes Material (PT = Paratypus): Holotypus: 9 mit drei Etiketten: [Eining 
a.D 24.6.1927 leg. Stoeckhert]; [Type]; [Andrena alutacea 9, det. E. Stoeckhert] (ZSM), 
Fundort: Eining an der Donau, Bayern, Deutschland. ASERBEIDSCHAN: 9 21.6.1957 
Nachitschewan, Chrusa, 2000 m. (OLL). BULGARIEN: 9 Pirin Geb. 1000-1800 m, Banska 
25.6.1938, PT (ZSM). 2 c? 2 9 30.5.1989 Slancev Briag (OLL). DEUTSCHLAND: Bayern: 2 
9 24.06.1932 Eining (1 9 PT); 9 24.6.1946 Markt Schwaben (GRÜ); 2 9 6.8.1933,27.7.1933 
Pappenheim, PT; 9 21.6.1883 Allach (ZSM); Eining 1.7.1938 (PT); 9 3.8.97 Ergersheim, 
Eschenau, Weinbergbrache, Daucus carota (Kraus); 9 17.7.2004 Altmühltal, 12 km NEE 
Eichstätt, Gundoldinger Heide (SE), Thüringen: 9 20.6.98 Kyffhäuser/Auleben/Solberg, Kalk- 
magerrasen auf Achillea sp; 1 9 23.6.01 Wanderslebener Gleiche b. Gotha, Apiaceae gestreift; 
6 20.6.98 Kyffhäuser/Badra/Feldrain, Apiaceae gestreift (Burger, die beiden letzten Tiere det. 
Burger); Baden Württemberg: 9 2.7.1995 Baar/Südostschwarzwald, 2 km N Geisingen, NSG 
Klause; S 10.6.1996 Hegau, Singen, Hohentwiel (SE). FÜRSTENTUM LIECHTENSTEIN: 9 
13.6.1997 Balzers, Elltal, ein frisches Tier, zusammen mit einer abgeflogenen proximo (ETH). 
GRIECHENLAND: 9 9.05.1973 Mistras; 9 29.06.1970 Vytina; 13.6.1956 Olymp, Prioni, 1000 
m; 9 19.05.1973 Mistria; Mazedonia. Treskaschlucht: 9 8.6.1973 Lehau (GRÜ). IRAN: 9 
20.5.1975 Kermanshahan, 80 km W Kermanshah, Paweh (GRÜ). ITALIEN: 16 9 23.7.- 
3.8.1933; 19.8.1926 Rovereto; 9 25.8.1913 Ritten (Südtirol) (ETH); 2 9 16.7.1925 Bozen, PT; 
bei Bozen, leg Kohl (MHNG). KROATIEN: 2 9 19.7.1971, 2.8.1972 Istria, Ucka (GRÜ). 
LETTLAND (Lativa): 9 9.08.1931 Meiten (GRÜ). MACEDONIEN: 9 Cakor-Pass 1969 
(ZSM). ÖSTERREICH: 9 26.6.1976 Dürnstein, Wachau (GRÜ), 4 9 22.7.1920 Innsbruck (PT, 
ZSM); 2 9 24.7.1998 Burgenland. Mühlgraben; 9 24.7.1998 Burgenland, Oberdrosen-Bergen; 
9 28.7.1999 Tauka; 9 27.7.1996 Steiermark, Katzelsdorf; 9 28.7.1996 Steiermark, Reith bei 
Hartmannsdorf; 9 9.7.1998 Tirol, Igls S. Innsbruck; 9 13.7.1991 Oberösterreich, St. 
Georgen/Gusen (OLL). SCHWEIZ: 2 9 26.7.1926 GR, Sent; 9 31.7.-3.8. GR, Brusio; 4 9 
23.7.1997, 8.8.1997 GR, Haldenstein, Böfel; 9 Juli 1915 TI, Lugano, Maroggia, PT (ZSM); 9 
29.7.1999 TI, Indemini, Paese (ETH) - 3 9 1 â TL Valle Maggia, Lodano, 340 m, an 
Anthriscus; 9 1 TI, Valle Maggia, Gordevivo, 310 m (SE). TÜRKEI: 9 8.8.1982 Pr. Hakkari, 
Sai Dag, Shagulut, SW Yüksekova 1700 m; 9 12.06.1978 Konya, Siile, 9 31.5.1959 Amasya, 
9 12.6.1976 Mus, 9 4.6.1972 Gürün, 9 20.05.1975 Karakurt, Arastal (GRU), 9 5.6.1945 
Gürün; 9 12.6.1978 Konya; 9 31.5.1959 Amaxya, 9 12.6.1976 Mut, 9 22.5.1975 Karakurt, 
Arastal; 3 9 Horosan, Arastal; 3 9 1.6.1980 35 km SW Hakkari, 6 Kars, 20 km W Karakurt 
1600 mNN, S 29.5.1980 Agri. 20 km N Patnos, â Nevshehir, Ürgüp 21.8.1971 (OLL). TURK- 
MENISTAN: 5 9 16.5.1975 Bacharden (GRU). 



PROXIANDRENA SUBGEN. NOV. 1037 

Diagnose: Weibchen: Andrena alutacea ist anhand der gleichförmigen 
Tergitchagrinierung, den matten Tergiten, der schwarzen Unterbehaarung auf dem 
Mesonotum und der durchgehenden Tergitendbinde IV von den verwandten Arten zu 
unterscheiden. Männchen: Das Männchen ist im direkten Vergleich kaum von ampia 
zu trennen. Von den anderen Arten läßt sich alutacea anhand der glänzenden 
Punktzwischenräume auf dem Mesonotum und der weißlichen Körperbehaarung sowie 
den - bei frischen Tieren - durchgehenden hinteren Tergitendbinden abtrennen. 

In Mitteleuropa läßt sich alutacea ebenfalls anhand der späten Flugzeit von pro- 
xima abgrenzen. Männchen treten in der Regel ab Mitte Juni, Weibchen ab Anfang Juli 
auf. Um diese Zeit gibt es praktisch keine frischen proxima mehr. 

Variation: Weibchen aus der Türkei weisen teilweise eine viel schwächer aus- 
geprägte Tergitchagrinierung auf als die mitteleuropäischen Tiere. Sie erscheinen 
dadurch stärker glänzend und sehr viel feiner skulpturiert. Allerdings ist die Variations- 
breite bei türkischen Tieren relativ hoch. Diese Merkmalsausprägung ist auch bei ein- 
zelnen Tieren aus Griechenland zu beobachten, so dass hier wahrscheinlich eine 
kontinuierliche Merkmalsverschiebung stattfindet, die der Variationsbreite der Art 
zuzurechnen ist. Allerdings sind Weibchen aus Turkmenistan wiederum stärker chagri- 
niert und weisen außerdem zum Teil eine relativ grobe, fast schon körnige Tergit- 
punktierung auf den Tergiten II-IV seitlich auf. 

Diskussion: Andrena alutacea wurde von Stoeckhert (1942) unter besonderem 
Hinweis auf die unterschiedliche Flugzeit von proxima abgetrennt und als eigene Art 
beschrieben. Dennoch zog Warncke (1967) die Art ohne jede Begründung ein, weitere 
Autoren folgten ihm (Dylewska, 1987; Gusenleitner & Schwarz, 2002). Erst Schmid- 
Egger & Scheuchl (1997) betrachteten alutacea mit Hinweis auf die Stoeckhertsche 
Beschreibung und aktuellem Material wieder als selbständige Art. 

Eine genaue Prüfung des vorliegenden Materials zeigt, daß sich alle Weibchen, 
die in Mitteleuropa im Juli und August gefangen wurden, auch morphologisch gut von 
Tieren unterscheiden, die aus dem Mai und Juni stammen. Das bestätigt meiner 
Meinung nach die Ansicht von Stoe ckhert, daß alutacea eine valide Art ist. Die Unter- 
scheidungsschwierigkeiten zu proxima liegen im Bereich dessen, was bei vielen 
schwierigen' Andrena Artengruppen (z. B. Micrandrena) üblich ist. Probleme können 
bei abgeflogenen Tieren in Griechenland auftreten, da dort die Variation von alutacea 
beträchtlich zunimmt. 

Auch wenn man die bestehende Ähnlichkeit der beiden Arten in Betracht zieht, 
würde eine Synonymisierung von alutacea mit proxima die unterschiedliche 
Phänologie keinesfalls auf befriedigende Weise erklären. Für eine partielle zweite 
Generation ist der zeitliche Abstand beider Taxa sehr kurz (sie überlappen sich in der 
Flugzeit um 1-2 Wochen), außerdem müssten dann beispielsweise im wärmebegün- 
stigten und gut besammelten Baden- Württemberg mehr Tiere von alutacea auftreten, 
was aber nicht der Fall ist (eigene Beobachtung, Doczkal mundi.). Daher erscheint das 
Konzept zweier valider Arten die wahrscheinlichere Lösung für den vorliegenden Fall. 

Anders verhält sich die Situation in Südosteuropa und vor allem in der Türkei. 
Während in Griechenland alutacea etwas später als proxima fliegt, wurden alle alu- 
tacea aus der Türkei, aber auch in Südosteuropa, jeweils im Mai und Juni gefangen 
(aus der Türkei liegen keine Tiere vor, die proxima entsprechen). Einzige Ausnahme 



1038 C. SCHMID-EGGER 

bilden ein Männchen und ein Weibchen aus der Türkei, die aus dem August stammen. 
Das Männchen aus Ürgüp besitzt eine sehr weitläufige Mesonotumpunktierung, die 
viel weiter als bei allen anderen Männchen ist (Punktabstände zum Teil mehrere 
Punktdurchmesser groß). Das Weibchen aus der Provinz Hakkari ist völlig frisch, 
besitzt eine vollständige Endbinde auf Tergit IV und hat eine insgesamt etwas kürzere 
und wie geschoren wirkende Körperbehaarung. Ansonsten stimmt es vollständig mit 
alutacea überein. Ob hier eine partielle zweite Generation vorliegt oder sich hinter alu- 
tacea noch weitere Arten verbergen, muss derzeit offen bleiben. 

Verbreitung: Andrena alutacea ist von Mitteleuropa über Südosteuropa bis in 
die Türkei, den Iran und Turkmenistan verbreitet. In Deutschland kommt sie in Bayern 
und dem Osten von Baden- Württemberg vor, die meisten Funde stammen dabei aus 
dem Einzugsbereich der Donau. Ein Einzelfund stammt aus dem südlichen Rheintal 
(Grißheim; Stoeckhert, 1942). Außerdem liegen verschiedene Funde aus Thüringen vor. 
In der Schweiz ist alutacea im Tessin sehr häufig, weitere Funden stammen aus der 
Ostschweiz (Graubünden) und aus Liechtenstein. Der nördlichste Nachweis stammt aus 
Lettland. Das Verbreitungsgebiet von alutacea überschneidet sich in weiten Teilen mit 
proxima, an mehreren Orten konnten beide Arten sogar zusammen gefangen werden. 

Stoeckhert (1942) meldet sie aus Deutschland: Bayern; Thüringen: Jena, Erfurt; 
Baden- Württemberg: Grißheim in Südbaden. Österreich: viele Funde. Polen: Krakau, 
Lemberg (Galizien). Schweiz: Lugano. Italien: Bozen, Ligurische Alpen, Turin; Trient, 
Fiume. Kroatien. Rumänien: Szankesd (Siebenbürgen). Bulgarien. Griechenland: 
Parnass. Georgien: Kaukasus, Borsholm bei Tiflis (dieser letzte Fund bezieht sich 
höchstwahrscheinlich auf bermela). 

Ökologie: Nach bisherigen Erkenntnissen ist alutacea wie ihre Schwesterart 
proxima oligolektisch auf Doldenblütler spezialisiert (Westrich 1989, der die 
Oligolektie von proxima feststellte, trennt nicht zwischen den beiden Taxa). Andrena 
alutacea besiedelt in Deutschland vielfach trockenwarme Magerwiesen. Im Tessin 
(Schweiz) fing ich die Art auf einer typischen ,Fettwiese' auf Anthriscus sylvestris und 
im Altmühltal (Bayern) am Rande einer Wachholderheide auf Daucus carota. Auf- 
grund der wenigen Funde und der Spezialisierung auf trockenwarme Offenstandorte 
müßte die Art in Deutschland in den betreffenden Bundesländern in die Rote Liste 
aufgenommen werden. 

Andrena ampia Warncke, 1967, stat. nov. 

Andrena proxima ssp. ampia Warncke, 1967: 229, 250: Erstbeschreibung â und 9 , Holotypus 
â; Typenfundort: Spanien, Alberche, OLL, untersucht. - Warncke, 1974a: 11 (Fauna 
Marokko) - Warncke, 1976: 151: (Fauna Iberien) - Erlandsson, 1979: 125 (Fauna 
Spanien) - Gusenleitner & Schwarz, 2002: 611-612 (Katalog, S. 1145 Verbreitungskarte 
von ampia). 

Untersuchtes Material: Holotypus: 9 mit drei Etiketten: [Alberche V 1909 Arias], 
[Andrena proxima ssp. ampia War. det. Dr. Warncke], mit rotem Karton: [Holotype], in coli. 
OLL. ASERBEIDSCHAN: 9 21.6.1957 Nachitachawan, Chrusa (GRÜ). FRANKREICH: 9 
29.6.1978 Provence, Mt. de Lure. Cruis, ca. 500 m (GRÜ); 49 23.8.1998 Ht. Alpes, Aiguilles, 
le Lombard 1800 m; 12 9 6.6.1997 Ht. Alpes, 18 km S Briancon, Gouffre 900 m, 9 6.6.1997 
Ht. Alpes, 6 km SW Briancon, Prelles 1200 m (CS). ITALIEN: Westalpen, Valle d'Aosta: 22 9 
8 6 7.7.1995; 29 16.7.1995; 2 9 19 â 27.6.1996,4 9 2.8.1997 Cognetal, Lillaz, 1700-1800 
m, an Lasiopitium aff. suer L.; 3 16.5.1996, St. Pierre 8000 m; 1 9 10 km E Aosta, Trois Villes, 



PROXIANDRENA SUBGEN. NOV. 1039 



1400 m NN. 16.5.1998 (CS). KASACHSTAN: 9 16.6.79 Alma Ata, Alataugeb. 1900m (Kraus) 
- Axaj bei Alma Ata, 19.6.1981 (OLL) (Etikettiert als Andrena kocoureka ssp. n., det Warncke 
Paratypus]. KIRGISIEN: 9 9 15.8.1998 Alaj,G.K. Gultscha Tal, Ak-Bosogo (GRÜ) - 2 S 1 9 
7.6.1998 Sandalach-Gebirge, Kain-Su Schlucht. 2000 m NN (Scheuchl). MAROKKO: 9 
13.5.1965 Azrou {proxima det Warncke) (OLL). SCHWEIZ: 9 29-31.7.1933 GR, Braggio-S- 
Marina, Calanca, 1000-1600 m leg. Nadig; 9 24.6.1934 GR, Rabius; 9 24.6.1885 VS, Sierre; 
Ô 19.8.1943 VS, Follatères; 9 26.7.1991 VS, Visperterminen 1380 m; 9 1.6.1991 VS, 
Hohtenn; (ETH) - 65 9 47 9 , Fangjahre 2001-2002 VS, Erschmatt: Undere Chastler, 1210 m; 
Oberu Castler 1270 m; Oberi Zaig 1450 m; Chritzmatte 1220 m; Rossuwald 1370 m; 
Rüemetschbodu 1530 m; Chritzmatte 1120 m, Ritin 1190 m; Platten 890 m, S 15.5-2.6., 9 
16.5.-27.6. (ORT) - 9 21.6.1998 VS, Jeizinen 1500 mNN, felsige Bergwiese (Burger, Herrman); 
4 9, VS: Sierre, Martigny, St. Luc, Niouc, alle vor 1900 (MHNG). SPANIEN: 9 Sierra 
Guadarrama, Dusmet; S Alberche V 1909 (leg) Arias; Montarco (leg) Dusmet, alles Paratypen 
det. Warncke (OLL) - 9 Villaviciosa, 9 2 c? Escoriai (alle det K. Warncke als proxima ssp. am- 
pia), 9 11.7.1977 Pr. Guadalajara, Tierzo bei Molina, 1100 m; 9 10.7.1975 Malaga, Ronda 
(GRÜ). TADSCHIKISTAN: 9 Dusambe, 40 km vych. Javros, 24.6.1981 (OLL) (Etikettiert als 
Andrena kocoureka ssp. n., det Warncke Holotypus]. 

Diagnose: Andrena ampia ist im weiblichen Geschlecht wie folgt charakteri- 
siert: Tergite überwiegend spiegelglatt, Tergitscheiben II-IV deutlich und grob punk- 
tiert, Punkte gleichmäßig über die Tergitscheibe verteilt. Mit alutacea teilt die Art die 
schwarze kurze Unterbehaarung des Mesonotums. Andrena ampia ist durchschnittlich 
1 Millimeter länger als alutacea (11-12 mm). Andrena alutacea Weibchen aus 
Turkmenistan, die auf den hinteren Tergiten seitlich ebenfalls grob punktiert sind, unter- 
scheiden sich auch hier durch die feine Chagrinierung auf den Tergiten. Die Männchen 
sind sehr ähnlich wie alutacea. Von dieser unterscheiden sie sich durch eine geringfügig 
dichtere und stärkere Tergitpunktierung sowie Tergitendbinden, die auf den Tergiten III 
und IV in fast allen Fällen unterbrochen sind (bei alutacea durchgehend). Andrena pro- 
xima läßt sich im männlichen Geschlecht leicht aufgrund des chagrinierten 
Mesonotums und der nur sehr fein und zerstreut punktierten Tergite unterscheiden. 

Variabilität: Die Tiere aus Zentralasien stimmen mit den europäischen Indivi- 
duen fast vollkommen überein. Die Tergitendbinden sind bei zwei vollständig erhal- 
tenen Weibchen nach hinten gerichtet und anliegend, während sie bei den europäischen 
Individuen abstehend und nach außen gerichtet sind. Die Tergitendbinde IV ist bei den 
asiatischen Tieren durchgehend, während sie bei den europäischen Individuen fast 
immer unterbrochen ist. Außerdem ist die dichte Tergitpunktierung bei einigen der 
asiatischen Weibchen körnig-dicht, die Punkte sind nur von schräg hinten zu erkennen. 

Diskussion: Andrena ampia wurde von Warncke als Unterart zu proxima von 
der iberischen Halbinsel beschrieben. Laut seiner Verbreitungskarte (publiziert in 
Gusenleitner & Schwarz, 2002) stellen die Pyrenäen die Verbreitungsgrenze zwischen 
der ssp. ampia und der ssp. proxima dar. Durch die aktuellen Funde von ampia in den 
Alpen wird diese Einteilung hinfällig. Außerdem ist das Taxon von den beiden Arten 
alutacea und proxima morphologisch gut unterschieden. In der Flugzeit liegt ampia 
etwa zwischen proxima und alutacea. Somit halte ich seinen Status als Unterart zu 
proxima für unwahrscheinlich und betrachte ampia als valide Art. 

Andrena ampia ist außerhalb der iberischen Halbinsel nur aus den trocken- 
warmen großen Binnentälern der Süd- und Südwestalpen nachgewiesen (Wallis, 
Graubünden, Tessin, Aostatal, Durancetal, bei Genf). Weiterhin liegt ein Fund aus der 
südlichen Provence vor. Es ist zu vermuten, dass die Art in Südfrankreich weiter 
verbreitet ist. Wie sie sich dort zu proxima abgrenzt, ist noch unbekannt. 



1040 c - SCHMID-EGGER 

In der Schweiz überlappen sich die Areale der drei mitteleuropäischen Arten der 
Untergattung Proxiandrena Allerdings sind auch hier Unterschiede in der Verbreitung 
feststellbar. So ist das Wallis (wie das ummittelbar südlich daran angrenzende italie- 
nische Aostatal) nach den bisherigen Daten nur von ampia besiedelt. Diese dringt 
jedoch auch noch weiter nach Südosten bis Graubünden und dem Tessin vor und findet 
sich auch in der Umgebung von Genf. Andrena alutacea besiedelt vor allem die 
Ostschweiz, bzw. Lichtenstein, ihr Verbreitungsgebiet erstreckt sich dabei vom 
Rheintal bis ins Tessin. Andrena proxima hingegen ist vor allem auf die Nordschweiz 
beschränkt, kommt aber stellenweise auch in den südlichen Landesteilen vor. 

Eine ähnliche Verbreitung wie ampia weist Andrena afrensis Warncke, 1967, 
auf, die in Spanien, dem Wallis (Westrich et al., 1997) und im Aostatal (Schmid-Egger, 
unpubl.) vorkommt. Die Schwesterart von afrensis. Andrena agilissima Scopoli 1770, 
die in Mitteleuropa weit verbreitet ist. scheint zumindest im Wallis zu fehlen, während 
im Aostatal beide Arten zum Teil syntop fliegen. 

Interessant ist weiterhin das Vorkommen von ampia in West- und Zentralasien, 
welches auf ein disjunktes Areal hinweist. Das Verbreitungsgebiet überschneidet sich 
dort nach den vorliegenen Daten weder mit proxima noch mit alutacea. Die asiatischen 
Tiere sind geringfügig von den europäischen unterschieden (runzelig dichte 
Punktierung auf den Tergiten III und IV lateral, durchgehende Endbinde auf Tergit IV), 
allerdings rechtfertigen diese Unterschiede keine Abtrennung der östlichen 
Populationen als Subspecies. 

Ökologie: Im Aostatal wurde ampia auf trockenen, blütenpflanzenreichen, süd- 
exponierten Almwiesen in Höhen zwischen 1700 und 1800 m ü. NN angetroffen. Die 
Art flog dort zahlreich auf Laserpitium cf. siler und war unter anderem vergesellschaf- 
tet mit Osmia dalmatica Morawitz (Apidae), Bembix tarsata Latreille (Crabronidae) 
oder Parnassius apollo Linné (Lepidoptera) . Im Durancetal flog sie in 800-1000 m ü. 
NN an Wegrändern und aufwiesen mit magerer, trockenheitsliebender Vegetation. Im 
Wallis konnte Sabine Oertli die Art ebenfalls in Anzahl auf südexponierten mageren 
Wiesen und Weiden antreffen. Sie stellte Blütenbesuch auf Anthriscus sylvestris, in 
geringerem Mass auch auf Heracleum sphondylium und Pimpinella major fest. Auf 
den Weiden blühte Laserpitium siler (Oertli in litt.). Somit scheint die Art wie auch 
proxima oligolektisch an Doldenblütlern (Apiaceae) gebunden zu sein. 

Verbreitung: Disjunkt in Europa und Asien. Europa: Südliche Schweiz; 
Norditalien: Aostatal: Südwestfrankreich: oberes Durancetal; Südfrankreich: 
Provence; Spanien. Marokko. Nach Warncke (1976) in Spanien und Portugal weit ver- 
breitet. Asien: Aserbeidschan. Kasachstan, Kirgisien. Tadschikistan. 

Andrena aspericollis Perez. 1895, stat. rest. 

Andrena aspericollis Perez 1895: 37 (Beschreibung 2, â, keinen Holotypus festgelegt) - 
Paralectotypus untersucht. 

Andrena proxima ssp. aspericollis: Warncke 1967: 183, 252 (Lectotypus 2 festgelegt, Teniet. 
Algerien, coli. Perez/Paris, stat. nov. ) - Warncke 1974a: 11 (Fauna Algerien, Flugzeit 
Mai bis Juni) - Gusenleitner & Schwarz 2002: 610-61 1 , 1 145 (Katalog). 

Untersuchtes Material: ALGERIEN: 2 mit unleserlichem Etikett, 2 S mit gedruk- 
ktem Etikett .Algerien*, alle det Warncke als ,proxima ssp. aspericollis' , das 2 zusätzlich mit 
einem roten Etikett .Paratype" (OLL) - 2 mit unleserlichem Etikett und verschiedenen 



PROXI ANDREN A SUBGEN. NOV. 1041 

Etiketten: Rosa mit schwarzem Rand [Cotypus], [Andrena aspericollis det. E. Stoeckhert]. 
[Andrena proxima det K. Warncke] (ZSM). 

Diagnose: Andrena aspericollis ist laut Warncke (1967) dunkler gefärbt als 
proxima und besitzt keine Binden auf den Tergiten. Die Art ist in ihrer Verbreitung auf 
Algerien beschränkt. Die Weibchen sind auf den Tergiten sehr gleichförmig und fein 
chagriniert und zerstreut punktiert, das Mesonotum weist eine dichte grobe 
Punktierung mit kleinen, glänzenden Punktzwischenräumen auf {ampia mit 
glänzenden und auf dem Mesonotum größeren Punktzwischenräumen). Die Männchen 
besitzen wie proxima ein durchgehend chagriniertes Mesonotum und unterscheiden 
sich dadurch von der ebenfalls in Nordafrika vorkommenden ampia, deren Punkt- 
zwischenräume auf dem Mesonotum glänzen. 

Diskussion: Ich konnte nur zwei mangelhaft etikettierte Pärchen untersuchen, 
die vermutlich alle aus Algerien stammen und wahrscheinlich über 100 Jahre alt sind. 
Alle diese Tiere sind ohne Tergitbinden. Ob das von Perez und Warncke beschriebene 
Hauptmerkmal der Art, die fehlenden Tergitbinden, auf den schlechten Erhaltungs- 
zustand der Tiere zurückzuführen oder natürlich ist, kann ich nicht beurteilen. Da sich 
die Art aber skulpturell sowohl von ampia als auch von den europäischen Arten unter- 
scheidet, habe ich keine Zweifel, daß es sich hier um eine eigene Art handelt. Somit 
stelle ich den ehemaligen Status von aspericollis als eigene Art wieder her: Andrena 
aspericollis stat. rest. 

Verbreitung: Algerien. 

Andrena bermela Warncke, 1974, stat. nov. 

Andrena proxima ssp. bermela Warncke 1974b: 96. Auflistung. - Warncke 1975: 50 (Teil B der 
Arbeit von 1974): Beschreibung S , Holotypus S: Türkei, Ankara. (OLL) - Holotypus 
untersucht, dieser stammt nicht wie bei Warncke 1975 angegeben von Ankara, sondern 
von Sereflikochisar. Allerdings liegen Paratypen von Ankara vor. 

Andrena alutacea: Stoeckhert 1942: 236 ff. Ein Männchen mit schwarzer Gesichtsbehaarung aus 
Tiflis/Georgien, bezieht sich höchstwahrscheinlich auf bermela. 

Untersuchtes Material. Holotypus: 6 [TR Sereflikochisar 17.5.1970 leg. Warncke] 
[Andrena proxima ssp. bernicla War. det. Dr. Warncke ]; rotes Etikett [Holotypus] (OLL). 
TÜRKEI. 6 2 9 21.5.1972 Ankara leg. Warncke (alle Paratypen, det Warncke, OLL). Die 
Weibchen unterscheiden sich nicht von alutacea und werden zu dieser gerechnet. 

Diagnose: Die Männchen unterscheiden sich von allen anderen Arten der 
Untergattung durch die schwarzbraune Gesichtsbehaarung (bei den übrigen Arten 
hell). Das Mesonotum ist ähnlich wie bei proxima skulpturiert. Das Weibchen ist nicht 
bekannt, bzw. läßt sich nicht von alutacea unterscheiden. 

Diskussion: Warncke (1974b) basiert die Beschreibung seiner Unterart bernicla 
auf die schwarze Gesichtsbehaarung des Männchen, die laut dem Autor allerdings nur 
in der ersten Generation vorkommen soll. Weiterhin ging Warncke in einer Ver- 
breitungskarte (publiziert in Gusenleitner & Schwarz 2002) davon aus, daß in der 
Türkei nur ein Taxon aus der proxima-Gruppe vorkommt, nämlich die ssp. bernicla. 
Wie das vorliegende Material zeigt, ist die Hypothese einer ersten dunkelgefärbten 
Generation jedoch nicht zu halten, da ich verschiedene türkische Männchen mit heller 
Gesichtsbehaarung untersuchen konnte, die wie bernicla ebenfalls im Mai gefangen 
wurden (zum Teil von Warncke selbst gesammelt). Da sich diese auch skulpturell von 



1042 C. SCHMID-EGGER 

bermela unterscheiden, gehe ich vielmehr davon aus, daß in der Türkei mindestens 
zwei Taxa aus der proxima-Gruppe, nämlich alutacea und bermela, nebeneinander 
vorkommen. Andrena proximo wurde bisher in der Türkei nicht nachgewiesen. 

Allerdings ist die Situation um die beiden Taxa damit noch längst nicht befrie- 
digend gelöst. In der coli. Warncke (OLL) befinden sich zwei Weibchen von Ankara, 
bzw. Siile bei Konya, die Warncke als bernicla-Paiatypen auszeichnete (ohne daß diese 
in der Literatur Erwähnung fanden). Beide Tiere - ein Weibchen wurde zeitgleich und 
am selben Ort mit dem männlichen Paratypus gefangen - sind völlig identisch mit alu- 
tacea- Weibchen aus Mitteleuropa und aus der übrigen Türkei. Entweder lassen sich die 
beiden Taxa im weiblichen Geschlecht nicht unterscheiden oder das Weibchen von 
bermela ist noch unbekannt. 

Ein weiteres Problem besteht außerdem bei den Männchen. Stoeckhert (1942) 
erwähnt in der Originalbeschreibung von alutacea, daß die meisten Männchen ge- 
mischte schwarze und weiße Gesichtshaare mit Übergängen zu ganz weiß oder ganz 
schwarz besitzen sollen. Im von mir selbst untersuchten Material gab es nur entweder 
weiße oder schwarz gefärbte Tiere. Ich konnte keine einzige Mischform finden. Es 
bleibt offen, von wo Stoeckhert solche Männchen untersucht hat, da er keine explizi- 
ten Fundortangaben dafür macht. Ingesamt basiert er die Beschreibung von alutacea 
auf nur 7 Männchen und über 80 Weibchen, was einen Hinweis auf die Seltenheit der 
Männchen gibt. Stoeckhert erwähnt jedoch ausdrücklich ein Männchen aus Tiflis/ 
Georgien mit vollständig dunkler Gesichtsbehaarung, was höchstwahrscheinlich eine 
bermela ist. 

Vorerst schließe ich aufgrund der morphologischen Unterschiede zwischen 
beiden Taxa und dem zeitlich übereinstimmenden Auftreten aus, daß bermela eine 
schwarz gefärbte Form von alutacea ist und betrachte das Taxon als eigenständige Art: 
Andrena bermela stat. nov. Die Betrachtung des Taxon als Form von alutacea wäre 
nicht plausibel, da mir solche lokalen Formen bei Andrena nicht bekannt sind. Die 
Gesichtsbehaarung ist vielmehr zumindest bei den mitteleuropäischen Arten ein 
konstantes Artmerkmal. Um eine partielle erste Generation von alutacea kann es sich 
auch nicht handeln, da eine solche eher in einer wärmeren Region der Türkei als gerade 
Ankara zu erwarten wäre. Die Frage der Weibchen von bermela bleibt allerdings 
ungeklärt. 

Verbreitung: Türkei, Georgien (Stoeckhert 1942, als alutacea). 

Andrena proxima (Kirby, 1802) 

Melitta proximo Kirby, 1802: 146-147. 

Andrena proxima wird sehr häufig in der taxonomischen und faunistischen Literatur erwähnt. 

Die einzelnen Meldungen und alte Synonyme sind hier nicht einzeln aufgeführt. Siehe 

dazu Gusenleitner & Schwarz, 2002: 610-611. 

Untersuchtes Material: Tiere aus Deutschland und Österreich werden nicht einzeln 
aufgeführt. Hier standen etwa 200 Tiere aus den Sammlungen CS, GRÜ, ZSM u.a. zur 
Verfügung, außerdem untersuchte D. Doczkal sein mehrere hundert Tiere umfassendes Material 
aus Baden-Württemberg auf mögliche alutacea. FÜRSTENTUM LIECHTENSTEIN: 9 
13.6.1997 Balzers, Elltal, ein abgeflogenes Tier, zusammen mit einer frischen alutacea; 9 
24.4.1997 Vaduz, Möhliholzröfi (ETH). ITALIEN: 9 20.04.1971 Sizilien, Taormina; 9 06.1920 
Liguria, Vittoria (GRÜ) - 9 20.5.1922 Sizilien Taormina, 9 Puglia, 9 31.5.1993 Mt. Gargano, 
StrGiovanni, 9 16.5.1992 Mt. Gargano, S. Marco (ETH). GRIECHENLAND: 9 11.05.1968 



PROXIANDRENA SUBGEN. NOV. 1043 

Fiorina; 2 9 13.6.1959 Olymp, Prioni, 1000 m (sehr abgeflogene Tiere, zusammen mit einer 
frischen alutacea (GRÜ) -29 9.5.1976 Nea Agathoupolis / Makedonien (Kraus). UNGARN: 9 
28.05.1959 Tatatovaros, 9 04.06.1957 Roros, Ipoly; 9 8.06.1954 Bükk hegys; Hosszuberc; 
Tihany u.a. Fundorte (GRÜ). SCHWEIZ: 2 9 4.6.1997 BE, Lauperswil, Bluttengrat 900 m; 9 
12.4.1942 GR, Haldenstein; 9 6.6.1936 GR, Felsberg; 9 8.6.1924 GR,Mundaun; 9 27.05.1925 
(ETH); TI, Lugano, Mt. Areo (GRÜ) 

Diagnose: Andrena proxima ist im weiblichen Geschlecht charakterisiert durch 
die teilweise glänzenden Tergitflächen, die zumindest seitlich auch deutlich punktiert 
sein können. Alle Tergitendbinden sind in der Mitte breit unterbrochen, das Meso- 
notum zeigt keine schwarze kurze Unterbehaarung, oder diese ist nur im hinteren Teil 
schwach ausgeprägt. Siehe auch die Diskussion bei alutacea. Die Männchen lassen 
sich anhand des vollständig chagrinierten Mesonotums, der nur sehr fein und zerstreu- 
ten Tergitpunktierung und wie beim Weibchen an den breit unterbrochenen 
Tergitendbinden erkennen. 

Verbreitung: Andrena proxima ist in Mitteleuropa verbreitet und stellenweise 
häufig (Verbreitungskarte bei Gusenleitner & Schwarz, 2002). Im Norden erreicht sie 
Schleswig Holstein und das südliche Dänemark. Außerdem besiedelt sie den Süden 
von Großbritannien. Ihre Arealausdehnung im Osten ist unbekannt, allerdings scheint 
sie in Asien (auch in der Türkei) zu fehlen. Ein Tier in der Sammlung GRÜ aus 
Lettland erwies sich zu alutacea gehörend. Im Südwesten erreicht sie Mittel- und 
wahrscheinlich auch Südfrankreich, allerdings muss hier die Artzugehörigkeit von 
Belegmaterial zu ampia überprüft werden. Italien ist bis Sizilien von proxima besie- 
delt, im Südosten erreicht sie Griechenland und kommt dort zum Teil syntop mit alu- 
tacea vor. 

Ökologie: Andrena proxima besiedelt vor allem mageres Grünland, aber auch 
andere trockenwarme Offenstandorte. Die Art ist auf Doldenblütler (Apiaceae) als 
Pollenquelle spezialisiert (Westrich, 1989). Die Art ist in Deutschland nicht gefährdet. 



DANKSAGUNG 

Für die Bereitstellung von Sammlungsmaterial bedanke ich mich bei den fol- 
genden Personen: Frank Burger (Weimar), Dr. Andreas Müller und Sabine Oertli 
(Zürich), Dr. Bernard Merz (Genf), Johannes Schuberth und Dr. Stefan Schmidt 
(München), Dr. Manfred Kraus (Nürnberg), Fritz Gusenleitner (Linz), Erwin Scheuchl 
(Velden) und Dr. Mike Herrmann (Konstanz). Erwin Scheuchl und Prof. Dr. Klaus 
Schönitzer gaben wertvolle Hinweise zum Manuskript. Erwin Scheuchl stellte mir 
außerdem freundlicherweise ausführliche Literaturauswertungen zur Verfügung. 



LITERATUR 

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Mitteleuropa. Acta Zoolocia cracovil 30: 359-708. 
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countries II. Acta Entomologica 15: 125. 



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Gusenleitner, F. & Schwarz, M. 2002. Weltweite Checkliste der Bienengattung Andrena mit 

Bemerkungen und Ergänzungen zu paläarktischen Arten (Hymenoptera, Apidae, 

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Bordeaux. 37. 
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Österreichs, Deutschlands und der Schweiz (Hymenoptera, Apidae). Entomofauna, 

Supplement 8: 398 Seiten. 
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Andrena alutacea n.sp. (Hym. Apid.). Mitteilungen der Münchener Entomologischen 

Gesellschaft 32: 236-252. 
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(Hymenoptera, Apoidae, Andrena). Mitteilungen des Zoologischen Museums in Berlin 

50: 3-53. 
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Mitteilungen der Münchener Entomologischen Gesellschaft 65: 29-102. 
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Warncke, K. 1986. Die Wildbienen Mitteleuropas, ihre gültigen Namen und ihre Verbreitung. 

Entomofauna, Supplement 3: 45. 
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Mitteleuropa neue Bienen-Art (Hymenoptera, Apidae). Linzer biologische Beiträge 29: 

1167-1174. 



Revue suisse de Zoologie 112 (4): 1045-1069; décembre 2005 



African biodiversity hotspots: the reptiles of Mt Nlonako, 
Cameroon 

Hans-Wemer HERRMANN 1 - 2 , Wolfgang BÖHME 3 , Oliver EUSKIRCHEN 4 , Patricia 

A. HERRMANN 2 & Andreas SCHMITZ 5 

Center for Reproduction of Endangered Species (C.R.E.S.), Zoological Society of 

San Diego, RO. Box 120551, San Diego, CA 92112-055, U.S.A. 

E-mail: hwherrmann@sandiegozoo.org 
2 P.O. Box 3055, Messa, Yaounde, Cameroon. 
3 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 

D-53113 Bonn, Germany. 
4 Bismarckstrasse 10, D-56626 Andernach, Germany. 
5 Department of Herpetology and Ichthyology, Museum of Natural History, C.P. 6434, 

CH-1211 Geneva 6, Switzerland. 

African biodiversity hotspots: the reptiles of Mt Nlonako, Cameroon. - 

The reptiles of Mt Nlonako, a mountain at the southeastern edge of the 
Cameroon mountain range ("Dorsale camerounaise"), were inventoried 
continually over a six year period from 1998 to 2004. This area encom- 
passes 150 km 2 of lowland, submontane and montane rainforest with an 
elevation up to 1,825 m. Accounts of 89 species are provided based on 
collected and photo-documented material. This inventory proved Mt 
Nlonako to be the most species rich single-locality area in reptilian fauna in 
Africa. With 63 snake species Mt Nlonako exhibits the greatest number of 
species in Africa and possibly worldwide. Analysis showed the reptilian 
species composition to be most similar within Cameroon with that of Korup 
National Park followed by the Dja Faunal Reserve in the south. Relative to 
the snake composition the Korup NP and the Dimonika region in Congo- 
Brazzaville show the highest resemblance. In an African context the reptile 
fauna of Mt Nlonako is characterized by species which occur in both West 
and Central Africa as opposed to the mountain's amphibian species which 
more closely resemble Central African fauna. The high species richness and 
endemicity is discussed from a paleoclimatic perspective. Conservation 
status and threats to the reptiles are noted. 

Keywords: Amphisbaenia - Crocodylia - Chelonia - Sauria - Serpentes - 
species richness - endemicity - biogeography - conservation. 

INTRODUCTION 

Despite efforts in surveying the reptile (and amphibian) fauna of the rain forests 
of tropical central and west Africa (Perret, 1959, 1960, 1961; Leston & Hughes, 1968; 
Böhme, 1975; Joger, 1982; Trape, 1985; Böhme & Schneider, 1987; Ota et al., 1987: 



Manuscript accepted 15.12.2004 



1046 H .- W. HERRMANN ET AL . 

Trape & Roux-Esteve, 1990; Lawson, 1993; Böhme, 1994a, b; Rodel et al., 1995; 
Schmitz, 1998; Rodel et al, 1999; Böhme, 2000; Euskirchen et al, 2000; Schmitz et 
al., 2000; Gossmann et al., 2002; Branch & Rodel, 2003) the knowledge on African 
rainforest reptile faunas falls behind that of its equivalents in tropical Middle and South 
America and in tropical Asia (Dunn, 1949; Inger & Col well, 1977; Murphy et al., 
1994). Within the rainforest zones of central and west Africa however, Cameroon, 
Ghana and Ivory Coast have the best studied reptile faunas (Hughes, 1983; Rodel et 
al, 1995; LeBreton, 1999; Rodel et al, 1999). Lawson (1993) lists 83 reptile species 
for the Korup National Park (Korup NP), which makes this one of the most species rich 
areas for reptiles on the continent. Trape (1975) records 45 snake species from the 
Dinamika forest site in the People's Democratic Republik of Congo (Congo- 
Brazzaville). 

In Cameroon, the Western and Southwestern Cameroon highlands also known 
as "Dorsale camerounaise" extending from Mt Cameroon in the south to Tchabal 
Mbabo in the north are characterized by a high amphibian and reptile species richness 
and an extraordinary high proportion of endemic species; a veritable hotspot of African 
herpetofauna diversity (Duellman, 1999; Poynton, 1999; Herrmann et al., 2005). 

Contrary to the high biodiversity value of such areas, especially in West and 
Central Africa, most have no formal conservation status protection (Stuart et al., 
1990) and are endangered by habitat destruction, mainly by logging activities and 
human encroachment. Additionally, the natural history of most reptile species from 
tropical forests in Africa remains unknown. Quantitative ecological studies are entirely 
lacking with only the first steps taken towards quantitative studies of chameleons 
(Herrmann & Herrmann, 2005). Ultimately, only a comprehensive understanding of 
the reptiles of these areas and their natural history will provide the fundamental 
knowledge for their future survival. 

This paper presents a comprehensive reptile species list for Mt Nlonako, which 
was derived over a period of six years, in which we compile the largest number of 
reptile species (89), in particular snake species (63), for any single-locality on the 
African continent. 

STUDY SITES 

We surveyed the Mt Nlonako area (Fig. 1) which extends roughly from 4°49'- 
4°56'N and from 9°56'-10°0rE and encompasses ca. 15,000 ha. The western and 
northern flanks face the town of Nkongsamba, and the busy road between Douala and 
Bamenda. The slopes on this side are heavily cultivated with the forest destroyed up to 
an elevation of approximately 1 .100 m. To the south and east however, the forest slopes 
are much less influenced by human activities. A vast lowland rainforest, encompassing 
several thousand km 2 , extends from the foothills of Mt Nlonako reaching past 
Nkondjok in the East and past Yabassi in the South. This area is divided by some 
unpaved roads and settlements. Logging is or has been carried out in many places 
within this area. Although hunting pressure is imminent, forest elephants, gorillas, 
chimpanzees, drills and other large mammals persist. Mt Nlonako itself raises from 
about 400 m elevation on the southern side to 1 ,825 m on its peak. The highest and 
central part of the mountain forms a cuvette, ca. 1.5 km in diameter, with much 



THE REPTILES OF MT NLONAKO, CAMEROON 



1047 



A= study site 



1a 




1b 



I 




Nkebe waterfall 

Badjong 



Ekomtolo 



km 



5 km 



Fig. 1 
Survey locations (a) in Cameroon and (b) at Mt Nlonako. Shaded areas are rural communities. 



1 048 H .- W. HERRMANN ET AL . 

grass/bracken in its center and with forested rims at 1 ,600 m on the north, east and 
south sides and the peak on the western side (Dowsett-Lemaire & Dowsett, 1999). 

The forest above 1,100 m is pristine with a tall canopy (25-30 m). The forest 
here seems to be drier and warmer than forests on the close-by Manengouba and 
Bakossi mountains or Mt Kupe at comparable altitudes. Botanically, Guttiferae 
{Allanblackia sp.) and Burseraceae (Santina trimerai) are especially common. Caesal- 
piniaceae (incl. Tessmannia anomala), Ebenaceae (Diospyros), Meliaceae, Mimo- 
saceae (Albizia), Moraceae, Olacaceae (Strombosia), Sapotaceae (incl. Chrysophyllum 
albidum), Steruliaceae (Cola) and Apocynaceae (Tabernaemontana sp.) are recorded 
(Dowsett-Lemaire & Dowsett, 1999). Above 1,450 m some rare montane species such 
as Poly scias fulva can be found locally. Many small to medium-sized creeks, often fast 
flowing and rocky, can be found in the forest. Swamps and pools are rare. 

The climate is warm and humid. Over a period of 34 years Nkongsamba (882 
m elevation) received an average of 2,762 mm rainfall per year (table 1 , Amiet, 1975). 
During that period the peak dry season extended from December to February with less 
than 50 mm precipitation per month. The peak rainy season extended from July to 
September with up to 482 mm precipitation per month. For detailed data on climate 
and rainfall see Herrmann et al. (2005). 

Fieldwork initiated in November 1998 and continual sampling extended to June 
2004. Sampling occurred in all seasons. 

Our survey efforts concentrated on the northern, eastern and southern slopes of 
Mt Nlonako with the following localities being the main points of collecting: 

- Ekambeng (EKA): village at the foot of the northern slopes in the vicinity of 
Nkongsamba, many coffee plantations, no primary forest 

- Mekoum (MEK): village on northern side of the mountain, many coffee plan- 
tations, no primary forest 

- Nguéngué (NGU): village on the rim between northwestern and southeastern 
slopes, N 4°55'02", E 9°59'21", 1,140 m elevation, some coffee plantations, second- 
ary and predominately primary forest 

- Summit (SUM): eastern side of the cuvette, N 4°54'47", E 9°57'93", 1,660 
m elevation, rock outcrops, primary montane forest, some areas with grass and bushes 

- Eyimba (EYI): very small village on the eastern side of the mountain, N 
4°52'92", E 9°59'19", 710 m elevation, small cultivated areas, much primary forest 

- Nkebe waterfall (NWF): area between the villages Ekomtolo and Badjong (but 
also including those two villages), N 4°49'83", E 9°55'49", 470 m elevation, coffee, 
oil palm and food crop plantations, predominately secondary forest, area with previous 
and current logging. 

METHODS 

Sampling methods 

We used Y-shaped drift fence/pitfall trap arrays (Corn, 1994) with segments of 
5 m length during the initial phase of the project at several localities. Catching success 
was very low to nil. This method was abandoned after some weeks. 

Initially, gluetraps (Bauer & Sadlier, 1992; Glor et al., 2000) which were fixed 
on tree trunks at about 1.5 m height, were used to sample geckos and other arboreal 



THE REPTILES OF MT NLONAKO, CAMEROON 1049 

reptiles. Only a few skinks (Trachylepis) were caught using this method. The abundant 
ants tended to attack trapped reptiles immediately. This method was abandoned after 
the first two weeks. 

We applied quadrat sampling with 8 x 8 m quadrats (Jaeger & Inger, 1994) at 
several localities at various elevations. Quadrats worked satisfactorily to gain quanti- 
tative data but are work intensive. For the quantitative results see Herrmann et al. 
(2000). 

Visual encounter surveys along transects (Crump & Scott, 1994) or opportu- 
nistic searches during the day and at night were the dominant methods used. The 
number of persons surveying varied, but was usually two or three; this method yielded 
the best success and was carried out throughout the study periods. Acoustic monitoring 
was also applied but only species represented by at least one voucher specimen are 
included in the species list. Our data was complemented by donated specimens which 
were encountered by villagers during their daily movements. 

Voucher specimens are deposited in the herpetological collection of the 
Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn (ZFMK) and 
in the collection of the senior author (HWH) in the U.S. The latter collection will be 
transferred to the National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. (USNM). All specimens were either fixed in 10% buffered formal- 
dehyde or 75% ethanol and consecutively preserved in 75% ethanol. 

BlOGEOGRAPHIC ANALYSIS 

To compare the species composition of Mt Nlonako with other areas in West 
and Central Africa we calculated the "coefficient of biogeographic resemblence" 
(CBR) after Duellman (1990) with the formula 

CBR = 2C / (Nj+N 2 ) 

in which C is the number of shared taxa (here species) in two compared areas, 
Nj is the number of taxa (here species) in area one and N 2 is the number of taxa (here 
species) in area two. A CBR value of would mean that Mt Nlonako shares no rep- 
tilian species with the compared area, a CBR value of 1 would mean that the species 
inventory in both areas are identical (Jansen & Köhler, 2002). Jansen and Köhler 
(2002) excluded ubiquitous species from their analysis arguing that those species are 
not primary inhabitants of the mountain forest habitats they compared and thus do not 
bear any information on the biogeographic relationships of those areas. We do not 
follow this approach for (1) we do not know if ubiquitous species are or are not pri- 
mary inhabitants of the areas in question, (2) possible "noise" by such species should 
be similar over the areas analyzed and (3) they are low in number and thus have a 
limited effect on the analysis. 

RESULTS 
Species account 

In the following we present a systematic list of reptile species encountered 
during our survey. We list localities and voucher specimens for each taxon. Nomen- 
clature follows Uetz et al. (2004), unless otherwise noted. For voucher specimens with 
no exact locality data we state the locality as Mt Nlonako (MtNL). Otherwise col- 



1050 H.-W. HERRMANN ET AL. 

lecting localities are Nkebe waterfall area (NWF), Ekambeng (EKA), Mekoum 
(MEK), Eyimba (EYI), Nguéngué (NGU) and the Mt Nlonako summit area (SUM). 
For habitat we differentiate between forest (F) and farmbush (FB). Farmbush are the 
ecotones between forests and cultivated areas as well as the vicinities of villages and 
the dirt roads. We also include degenerated secondary forest patches in this category. 

Amphisbaenia 

Amphisbaenidae 

Monopeltis sp. (Duméril, 1859) 

Locality: EYI. Voucher specimen: ZFMK 68960. Habitat: FB. 
Remarks: Only two species, Monopeltis jugularis Peters, 1880 and M. galeata 
(Hallowell, 1852) are known from Cameroon. This is the first record of an am- 
phisbaenid from Western Cameroon. Since this specimen is a roadkill in a rather 
bad conservation state, an identification to the species-level was not possible. 

Crocodylia 

Crocodylidae 

Osteolaemus tetraspis Cope. 1861 

Locality: NWF, EYI. Voucher specimen: none. Habitat: F. 
Remarks: Regularly seen in streams and pools. This species is commonly found 
as bushmeat in local markets such as in Nkongsamba (see Akani et al., 1998 for 
the situation in Nigeria). Although under severe hunting pressure, this species 
seems to occur in stagnant river arms and pools at lower elevations throughout 
the study area. CITES appendix I. 

Chelonia 

Testudinidae 

Kinixys erosa (Schweiger, 1812) 

Locality: NGU. Voucher specimen: ZFMK 68982. Habitat: F. 

Remarks: Shells of this species can regularly be found in villages where this 

species is appreciated as bushmeat (Lawson, 2000). CITES appendix II. 
Kinixys homeana Bell, 1827 

Locality: NWF. Voucher specimen: none. Habitat: F. 

Remarks: Shells of two specimens of this species are kept in the compound of 

the chief of Ekomtolo. Like K. erosa, this species is commonly eaten by local 

people. CITES appendix IT. 

Sauria 

Agamidae 

Agama agama (Linnaeus. 1758) 

Locality: NGU. Voucher specimens: ZFMK 69017, 75376. Habitat: FB. 

Remarks: This species was found in all settlements of the study area where it is 

very abundant in villages and along forest roads. It was never observed in the 

forest. 



the reptiles of mt nlonako, cameroon 105 1 

Chamaeleonidae 

Chamaeleo camerunensis Müller, 1 909 

Locality: NWF. Voucher specimen: HWH 20. Habitat: F. 

Remarks: Only one 2 of this species was found and is treated in detail in 

Harbort & Herrmann (2002). CITES appendix II. 

Chamaeleo cristatus Stutchbury, 1837 

Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 68961 , 68980- 
1, 73063-6, 75146, 75394, HWH 31, 202-4, 617, 816-29, 893-6. Habitat: F, FB. 
Remarks: CITES appendix II. 

Chamaeleo montium Buchholz, 1874 

Locality: MtNL, NGU. Voucher specimens: ZFMK 69117-20, 72855, 73067- 
74, 73491-5, 75388-93, HWH 208-12, 830-44. Habitat: F, FB. 
Remarks: CITES appendix II. 

Chamaeleo oweni Gray, 1831 

Locality: MtNL, NWF. Voucher specimens: HWH 205-7. Habitat: F, FB. 
Remarks: CITES appendix II. 

Chamaeleo pfefferi Tornier, 1900 

Locality: SUM. Voucher specimens: ZFMK 73490, 74336, HWH 1013. 

Habitat: F. 

Remarks: CITES appendix II. 

Rhampholeon spectrum (Buchholz, 1874) 

Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 69101-16, 72853- 
4, 73075-8, 73496, 75287-95, HWH 16, 194-200, 618, 845-85. Habitat: F 

Gekkontdae 

Hemidactylus intestinalis Werner, 1897 

Locality: NWF. Voucher specimens: ZFMK 78684, 81713, HWH 213-4, 602. 
Habitat: F,FB. 

Remarks: Henle & Böhme (2003) provide a detailed species account, though 
using the name Hemidactylus ansorgii Boulenger, 1901, which has been shown 
to be a junior synonym of//, intestinalis Werner, 1897 by Perret (1975), and this 
synonymy has also been mentioned in Kluge (1991). 

Hemidactylus echinus O'Shaughnessy, 1875 

Locality: NGU. Voucher specimen: ZFMK 69040. Habitat: F, FB. 

Hemidactylus fasciatus Gray, 1842 

Locality: NWF. Voucher specimen: ZFMK 77893. Habitat: F. 

Hemidactylus mabouia (Moreau de Jonnès, 1818) 

Locality: NGU. Voucher specimens: ZFMK 69026-39, 75377-9, HWH 1056-7. 
Habitat: FB. 

Remarks: Like Agama agama, this species can be found in all human settlement 
in the study area. It is especially abundant on houses. 



1052 H .- W. HERRMANN ET AL . 

Hemidactylus sp. indet. 

Locality: NWF. Voucher specimen: ZFMK. Habitat: F, FB. 
Remarks: Juvenile specimen which we could identify to species level. 

Lygodactylus conraui Tornier, 1902 

Locality: EYI. Voucher specimen: ZFMK 68966. Habitat: F, FB. 

Lacertidae 

P oromera f ordii (Hallowell, 1857) 

Localities: NWF, NGU. Voucher specimens: ZFMK 69021-5, 78660, 81715, 
HWH 1058. Habitat: F,FB. 

SCINCIDAE 

Trachylepis affinis (Gray, 1838) 

Localities: EYI, NGU. Voucher specimens: ZFMK 68962-4, 69062-73, 75383- 
7. Habitat: FB. 

Remarks: Listed as Mabuya affinis in Uetz et al. (2004). We follow the nomen- 
clature proposed by Bauer (2003) and use the genus name Trachylepis. 
Common in villages, farms and along roads throughout the study area. 

Trachylepis maculilabris (Gray, 1845) 

Localities: NGU, SUM. Voucher specimens: 69041-61, 69434, 75576-7. 
Habitat: F, FB. 

Remarks: Listed as Mabuya maculilabris in Uetz et al. (2004). We follow the 
nomenclature proposed by Bauer (2003) and use the genus name Trachylepis. 
Found in human settlements but also in secondary forest and on rock faces at 
the summit of Mt Nlonako. Locally abundant. 

Lygosoma fernandi (Burton, 1836) 

Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 68959, 69018, 
73497, 78026-7, 78814. Habitat: F, FB. 

Remarks: In Uetz et al. (2004) as Mochlus fernandi . We prefer the genus name 
Lygosoma as molecular analyses reveal a closer relationship with this genus 
(A. S. unpublished results). 

Panaspis breviceps (Peters, 1873) 

Localities: NWF, NGU. Voucher specimens: ZFMK 69074-85, 72856, 75380, 
77889-92, 78700, 78711, HWH 601. Habitat: F. 

Lacertaspis reichenowi (Peters. 1874) 

Locality: EYI. Voucher specimen: ZFMK 68965. Habitat: F 

Remarks: Listed as Panaspis reichenowi in Uetz et al. (2004). We follow the 

nomenclature proposed by Schmitz et al. (2005) and use the genus name 

Lacertaspis. 

Lacertaspis rohdei (Müller, 1910) 

Locality: NGU. Voucher specimens: ZFMK69086-99, 75382, HWH 1052-5. 
Habitat: F. 



THE REPTILES OF MT NLONAKO, CAMEROON 1053 

Remarks: Listed as Panaspis rohdei in Uetz et al. (2004). We follow the nomen- 
clature proposed by Schmitz et al. (2005) and use the genus name Lacertaspis . 

Leptosiaphos vigintiserierum (Sjöstedt, 1897) 

Localities: NGU, SUM. Voucher specimens: ZFMK 69426-32, 69549. Habitat: 

F. 
Leptosiaphos spp. 

Locality: NGU. Voucher specimens: ZFMK 69529-31, 69550-4, 75381. 

Habitat: F. 

Remarks: Our material of hitherto undescribed or unidentifiable Leptosiaphos 

includes five morphologically distinct groups. One groups resembles L. pau- 

liani (Angel, 1940) (Fig. 2) from the Bamboutos Mtns, Cameroon, another L. 

vigintiserierum (Fig. 3). 

Varante» AE 

Varanus ornatus (Daudin, 1803) 

Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 69016, HWH 18, 
982. Habitat: F,FB. 

Remarks: A common species which is often found as bushmeat in villages and 
local markets. Large specimens sometimes are encountered in snare traps. 
CITES appendix II. 

Serpentes 

Typhlopidae 

Typhlops angolensis (Bocage, 1866) 

Localities: EKA, NGU. Voucher specimens: ZFMK 75126, 75178-9. Habitat: 

FB. 
Typhlops congestus (Duméril & Bibron, 1844) 

Locality: MtNL. Voucher specimens: ZFMK 77642, 78132-6, 78819-24, HWH 

587. Habitat: FB. 
Typhlops lineolatus Jan, 1864 

Locality: MtNL. Voucher specimens: HWH 588, 695. Habitat: FB. 
Typhlops punctatus (Leach in Bowdich, 1819) 

Locality: MtNL Voucher specimens: ZFMK 78815-8. Habitat: FB. 

BOIDAE 

Calabaria reinhardtii (Schlegel, 1848) 

Localities: MtNL, NGU. Voucher specimens: ZFMK 75780, HWH 690-1 , 808- 
15, 897-901, 1009. Habitat: F, FB. 

Remarks: This is one of the most abundant snakes in the area. Often found cros- 
sing roads or dead on the road (DOR). The species was observed in many dif- 
ferent localities in the study area. CITES appendix II. 

Python sebae (Gmelin, 1789) 

Localitiy: MtNL. Voucher specimens: HWH 890-1. Habitat: F, FB. 



1054 



H.-W. HERRMANN ET AL. 




Figs 2-3 
2. Leptosiapìws sp. cf. pauliani from the forest above Nguéngué. 3. Leptosiaphos sp. cf. vigiliti- 



THE REPTILES OF MT NLONAKO, CAMEROON 1055 

Remarks: This species is probably restricted to the lower elevations of the study 
area. People of NGU for example did not confirm the existence of the species 
in their area. On one occasion, villagers of Ekomtolo were observed preparing 
a medium sized P. sebae for consumption. The species is well known to the peo- 
ple of Ekomtolo and Badjong. It is not common. CITES appendix II. 

COLUBRIDAE 

Afronatrix anoscopus (Cope, 1861) 

Locality: MtNL, NWF. Voucher specimens: ZFMK 81716, HWH 400, 753. 

Habitat: F, FB. 
Boiga blandingii (Hallo we 11, 1844) 

Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 68987, 75119, 

75175-7, 75857-8, HWH 681, 749-51. Habitat: F, FB. 
Boiga pulverulenta (Fischer, 1 856) 

Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 75155-7, 

77913-4, 78085, 78852-6, 81717, HWH 423-32. Habitat: F, FB. 
Bothrophthalmus brunneus (Günther, 1863) 

Locality: MtNL, NWF. Voucher specimens: ZFMK 75121, 78839, HWH 418, 

450. Habitat: F 

Remarks: The former B. lineatus subspecies brunneus is considered to be a full 

species by Hughes (2000) and Gossmann et al. (2002). 
Chamaelycus fasciatus (Günther, 1858) 

Localities: MtNL, NWF, EKA, MEK, EYI, NGU, SUM. Voucher specimens: 

ZFMK 69002, 69433, 75224-32, 77631 , 77663, 78103, 78845-6, HWH 407-17, 

633. Habitat: F 
Dasypeltis fasciata Smith, 1849 

Locality: MtNL. Voucher specimens: HWH 404-5. Habitat: F. FB. 
Dasypeltis scabra (Linnaeus, 1758) 

Localities: MtNL, NGU. Voucher specimens: ZFMK 78118-9, 78864-7. 

Habitat: F,FB. 
Dipsadoboa duchesnei (Boulenger, 1901) 

Locality: MtNL, NWF. Voucher specimens: ZFMK 81718, HWH 435-7, 715. 

Habitat: F. 
Dipsadoboa underwoodi Rasmussen, 1993 

Locality: NWF. Voucher specimen: ZFMK 81719. Habitat: F 
Dipsadoboa unie olor Günther, 1858 

Localities; MtNL, NGU. Voucher specimens: ZFMK 75266-7, HWH 438, 716. 

Habitat: F 
Dipsadoboa viridis (Peters, 1869) 

Locality: MtNL. Voucher specimens: ZFMK 78862-3, 79031, HWH 433-4, 

717, 759-64. Habitat: F 
Dipsadoboa weileri (Lindholm, 1905) 

Localities: MtNL, NGU. Voucher specimens: ZFMK 75340, 78121-2, HWH 

439-40. Habitat: F 



1056 H. -W. HERRMANN ET AL. 

Gastropyxis smaragdina (Schlegel, 1837) 

Localities: MtNL, NWF, EKA, MEK, EYI, NGU. Voucher specimens: ZFMK 
68975, 68989-91 , 75134, 75152-4, 75262-4, 77657-60, 77906, 78077, 78093-4, 
79029-30, HWH 399, 626, 802. Habitat: F, FB. 

Gonionotophis brussauxi (Mocquard, 1889) 

Localities: MtNL, NWF, EYI. Voucher specimens: ZFMK 77636, 77915, 
78850-1. Habitat: F. 

Grayia caesar (Günther, 1863) 

Locality: NGU. Voucher specimen: ZFMK 78120. Habitat: F. 

Grayia smythii (Leach, 1818) 

Locality: NWF. Voucher specimen: HWH 1208. Habitat: F, FB. 

Hapsidophrys lineatus Fischer, 1856 

Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 75125, 75265, 
75373, 78078-9, 79022-8, HWH 387-93, 803-6. Habitat: F, FB. 

Hormonotus modestus (Duméril, Bibron & Duméril, 1854) 

Locality: MtNL. Voucher specimens: ZFMK 78859-61. Habitat: F. 

Hydraethiops melanogaster Günther, 1872 

Locality: MtNL. Voucher specimens: ZFMK 72729, HWH 694, 752. Habitat: F. 

Lamprophis fuliginosus-lineatus-complex 

Thorpe & McCarthy (1978) state, that the two species L.fuliginosus and L. lin- 
eatus are difficult to differentiate in some areas. Our material allowed the as- 
signment of specimens to either of the two species. 

Lamprophis fuliginosus (Boie, 1827) 

Locality: NGU. Voucher specimens: ZFMK 69010-5, 78219. Habitat: F, FB. 

Lamprophis lineatus Duméril, Bibron & Duméril, 1854 

Localities: MtNL, NWF, NGU. Voucher specimens: HWH 441-9, 628-30, 702- 
9, 778-94, 965-74, 976, 1008, 1163. Habitat: F, FB. 

Lamprophis olivaceus (Duméril, 1856) 

Localities: MtNL, NWF, MEK, NGU. Voucher specimens: ZFMK 68976, 
75122, 75240, 75343, 77920, 79037-9, HWH 590, 981. Habitat: F, FB. 

Lamprophis virgatus (Hallowell, 1854) 

Localities: MtNL, NWF, EKA, MEK, EYI, NGL. Voucher specimens: ZFMK 
68977, 69007-9, 75116-8, 75138-45, 75147-51, 75344-71, 77637-8, 77662, 
77921-2, 78203-18, 79040-66. 

Lycophidion laterale Hallowell. 1857 

Localities: MtNL, NGU. Voucher specimens: ZFMK 69398, 75233-5, 75374, 
78080-1, 78843-4, HWH 406, 754-5. Habitat: F. 

Mehelya guirali (Mocquard. 1887) 

Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 69001 , 75169- 
74, 75270, 77909, 78082-4, 78092, 78830-2. Habitat: F, FB. 

Mehelya poensis (Smith, 1847) 

Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 75268, 77910- 
2, 78825, 78827-9, HWH 419, 616, 713, 756-8. Habitat: F, FB. 



THE REPTILES OF MT NLONAKO, CAMEROON 1057 

Mehelya riggenbachi (Sternfeld, 1910) 

Localities: MtNL, NWF. Voucher specimens: HWH 420, 631, 718. Habitat: F. 
Mehelya stenophthalmus (Mocquard, 1887) 

Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 75269, 78131, 

78826, HWH 632, 984. Habitat: F, FB. 
Natriciteres fuliginoides (Günther, 1858) 

Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: 75220, 77643- 

56, 77916-7, 78104, 78178-202, 78985-79003, HWH 451-74, 610-1, 613-4, 

696-8,701,767-74,942-58, 1010. Habitat: F,FB. 
Natriciteres olivacea (Peters, 1854) 

Localities: MtNL, MEK, NGU. Voucher specimens: ZFMK 68974, 68992, 

68997-9, 75201-19, 75341-2, 75372, 78979. Habitat: F, FB. 
Natriciteres variegata (Peters, 1861) 

Localities: MtNL, NWF, NGU. Voucher specimens: ZFMK 68993-6, 75180- 

200, 77918-9, 78095, 78173-7, 78980-4, HWH 475-89, 698-700, 776-7, 959- 

64. Habitat: F,FB. 
Philothamnus carinatus (Andersson, 1901) 

Localities: MtNL, MEK, NGU. Voucher specimens: ZFMK 75236-9, 75339, 

78127, 79032-6, HWH 634-6, 712, 798, 985. Habitat: F. 

Remarks: In Uetz et al. (2004) not listed for Cameroon and in LeBreton (1999) 

as a subspecies of heterodermus . 
Philothamnus heterodermus (Hallowell, 1857) 

Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 77632-5, 78123-6, 

HWH 710-1, 795-7, 807. Habitat: F, FB. 
Philothamnus nitidus (Günther, 1863) 

Locality: MtNL. Voucher specimen: HWH 637. Habitat: F. 
Poecilipholis cameronesis Boulenger, 1903 

Locality: MtNL Voucher specimen: HWH 589. Habitat: F. 
Rhamnophis aethiopissa Günther, 1 862 

Locality: NGU. Voucher specimen: ZFMK 68979. Habitat: F, FB. 
Rhamnophis batesii (Boulenger, 1908) 

Localities: MtNL, NWF, EKA, NGU. Voucher specimens: ZFMK 75334, 

77661, 78105-10, 78833-8, 79067, HWH 401-2, 627. Habitat: F 
Thelotornis kirtlandii (Hallowell, 1844) 

Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK 68988, 

75271-5, 77664, 78073-4, 78871-9, HWH 421-2, 615, 625, 680, 975, 1006-7. 

Habitat: F, FB. 
Thrasops flavigularis (Hallowell, 1852) 

Localities: MtNL, NGU. Voucher specimens: ZFMK 78032-3, 78840-2, HWH 

403. Habitat: F,FB. 



1058 H. -W. HERRMANN ET AL. 



Atractaspididae 



Aparallactus modestus (Günther, 1859) 

Localities: NWF, EYI, MEK, NGU. Voucher specimens: ZFMK 68978, 69003- 

6, 75123-4, 75241-60, 75335, 75337-8, 77640-1, 77907-8, 78150-72, 79006- 

21, 79221-2, HWH 640-9, 977-80, 1011-2. Habitat: F, FB. 
Aparallactus niger Boulenger, 1897 

Locality: MtNL. Voucher specimen: ZFMK 79005. Habitat: F. 

Remarks: The specimen possesses a single prefrontal with the nasals in broad 

contact. This specimen represents the first record of this species for Cameroon. 
Atractaspis aterrima Günther, 1 863 

Locality: MtNL. Voucher specimen: HWH 766. Habitat: F, FB. 
Atractaspis boulengeri Mocquard, 1897 

Locality: EKA. Voucher specimen: ZFMK 75136. Habitat: F. 
Atractaspis irregularis (Reinhardt, 1843) 

Locality: EKA. Voucher specimen: ZFMK 75137. Habitat: F, FB. 
Atractaspis reticulata Sjöstedt, 1896 

Localities: MtNL, NGU. Voucher specimens: ZFMK75261, HWH 639, 714, 

765. Habitat: F 
Polemon collaris (Peters, 1881) 

Localities: MtNL, NGU. Voucher specimens: ZFMK 69000, 75221-3, 78137- 

49, 78847-9, HWH 364-5, 693. Habitat: F. 
Polemon gabonensis (Duméril, 1856) 

Locality: NWF. Voucher specimen: ZFMK 75120. Habitat: F. 
Polemon gracilis (Boulenger, 1911) 

Locality: NGU. Voucher specimen: ZFMK 78129. Habitat: F. 

Remarks: This is the first voucher specimen from Cameroon. 
Polemon notatus (Peters, 1882) 

Localities: NWF, EYI, NGU. Voucher specimens: ZFMK 77905, 78128, HWH 

17. Habitat: F 

Elapidae 

Dendroaspis jamesoni jamesoni (Traill , 1 843 ) 

Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK68957, 
75276, 75336, 77672-4, 77898. 78096-102, 78887-95, HWH 380-2, 719-24, 
996-8. Habitat: F,FB. 

Naja melanoleuca Hallowell, 1857 

Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK 68985- 
6, 69019-20. 75127-33, 75158-62, 75277-84, 75793, 77630, 77667-71, 77900- 
4, 78034-53, 78947-78, 79004, HWH 384-6, 604-5. 612, 619-22, 676-9, 727- 
36, 889, 925-41, 1000-5. Habitat: F, FB. 

Remarks: In the study area this is the most abundant snake in villages where 
specimens are often attracted by poultry. 



THE REPTILES OF MT NLONAKO, CAMEROON 1059 

Pseudohaje goldii (Boulenger, 1895) 

Localities: MtNL, NWF, EKA, EYI, NGU. Voucher specimens: ZFMK 68984, 
69598, 75163, 77629, 77665-6, 77897, 78028-31, 78880-6, HWH 370-9, 682- 
4, 725-6, 999. Habitat: F, FB. 

VlPERIDAE 

Athens squamigera (Hallowell, 1854) 

Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 75285-6, 77622-3, 
78111-7, 78868-70, HWH 394-5, 692, 1183. Habitat: F. 

Causus lichtensteinii (Jan, 1859) 

Localities: MtNL, EYI, NGU. Voucher specimens: ZFMK 68959, 75164, 
77624-8, 78075-6, 78896-901, HWH 396-8, 607-9. Habitat: F, FB. 

Causus maculatus (Hallowell, 1842) 

Localities: MtNL, NWF, EKA. Voucher specimens: ZFMK 75135, 77899, 
78902, HWH 687-9, 740-8. Habitat: F, FB. 

Bitis gabonica (Duméril, Bibron & Duméril, 1854) 

Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 68983, 75168, 
75331-3, 77894-6, 78070, 78086-91, 78903-9, HWH 367-9, 888, 902, 993-5. 
Habitat: F,FB. 

Bitis nasicornis (Shaw, 1802) 

Localities: MtNL, NWF, EYI, NGU. Voucher specimens: ZFMK 75165-7, 
75296-330, 75375, 77617-21, 78054-69, 78071-2, 78130, 78910-46, HWH 
366, 606, 623-4, 685-6, 737-9, 903-24, 987-92. Habitat: F, FB. 

DISCUSSION 

We recorded a total of 89 reptile species from Mt Nlonako (table 1 and 3). 
Thirty-six percent of all 249 reptile species (excluding sea turtles) and 42 percent of 
the 151 snake species recorded for Cameroon (LeBreton, 1999) occur at Mt Nlonako. 
Thus Mt Nlonako hosts the most species rich single-locality reptile and especially 
snake fauna in Africa (Böhme, 2000). For snakes Mt Nlonako at this stage possibly 
represents the most species rich single-locality in the word. The subsequent area of rep- 
tile species richness is represented by Korup NP approximately 130 km W of Mt 
Nlonako with 83 reptile species (Lawson, 1993). This is followed by the Dja Faunal 
Reserve in Cameroon (80 species, LeBreton, 1999). Relative to snakes Mt Nlonako is 
again followed by the Korup NP (55 species), Mt Nimba in Guinea (52 species, Ineich, 
2003), the Dja Faunal Reserve (49 species, LeBreton, 1999), the Dimonika forest site 
in Congo-Brazaville (45 species, Trape, 1975), the Comoe NP in Ivory Coast (44 
species, Rodel 1995 and 1999), the Ziama Forest in Guinea (41 species, Böhme, 2000) 
and 40 species found in the Tai NP in Ivory Coast (Rodel & Mahsberg, 2000; Ernst & 
Rodel, 2002). Four snake species assemblages, reported from Ivory Coast, Ghana, 
Gabon and the Central African Republic, ranged between 35 and 39 species (Böhme, 
2000). These results may not only represent the actual pattern of snake species richness 
in the rainforest areas of West and Central Africa but may also indicate survey activi- 
ties and gaps as many areas with a very high potential for exceptional species richness 
have not been sampled adequately. 



1060 



H.-W. HERRMANN ETAL. 



Table 1 . Reptile species richness of three different rainforest areas in southwestern and southern 
Cameroon. Numbers in parentheses are the number of species directly recorded by the authors 
for the areas. NP = National Park: FR = Faunal Reserve. From LeBreton's list only species found 
or mentioned from the Dja FR or adjacent areas were included in this analysis. 





Mt Nlonako 


KorupNP 


DjaFR 


area in km 2 


150 


1,240 


8,400 


reptile species 


89 


83 (69) 


80 (14) 


reptile species/km 2 


0.59 


0.07 


0.01 


snake species 


63 


55 (41) 


49 (10) 


snake species/km 2 


0.42 


0.04 


0.01 


source 


this study 


Lawson, 1993 


LeBreton, 1999 



Table 2. West and Central African rainforest reptile and snake faunas and their relationships 
with the Mt Nlonako reptile and snake fauna. 



country 


locality 


reptile 
species 


CBR 


snake 
species 


CBR 


source 


Cameroon 


Korup National Park 


83 


0.65 


55 


0.72 


Lawson, 1993 


Congo (RC) 


Dimonika region 


- 


- 


45 


0.70 


Trape, 1985 


Cameroon 


Dja Faunal Reserve 


80 


0.60 


49 


0.66 


LeBreton, 1999 


Guinea, Ivory Coast, 


Mt Nimba 


69 


0.51 


52 


0.57 


Ineich, 2003 


Liberia 














Gabon 


Mounts de Cristal 


48 


0.54 


32 


0.55 


Pauwels et al. 2002 


Gabon 


Massif du Chaillu 


46 


0.47 


29 


0.50 


Pauwels et al., 2002 


Guinea 


Zima forest 


46 


0.43 


41 


0.50 


Böhme, 2000 


Ivory Coast 


Tai NP 






40 


0.50 


Rodel & Mahsberg, 
2000, Ernst & Rodel, 
2002 


Cameroon 


Mt Kupe & Bakossi 
Mtns 


38 


0.47 


18 


0.35 


Schmitz, 1998 


Ivory Coast 


Haute Dodo 


17 


0.23 


9 


0.19 


Branch & Rodel, 
2003 



Within Cameroon, our data shows not only the highest species richness for a 
single locality, but also the highest number of species per area (km 2 ) as presented in 
table 1. 

Of the 89 species recorded for Mt Nlonako, one species is an amphisbaenid 
(1%), one is a crocodile (1%) and two are tortoises (2%). Of the 22 lizard species 
(25%) one is an agamid (5%), one a lacertid (5%), one a varanid (5%), five are geckos 
(23%), six are chamelons (26%), and eight skinks (36%). The 63 snake species (71%) 
include two boids (3%), three elapids (5%), four typhlopids (6%), five vipers (8%), ten 
atractaspidids (16%), and 39 colubrids (62 %). The lizard fauna appears depauperated 
compared to lizard faunas in arid or semiarid African environments. However, similar 
findings are presented by Böhme (2000) for the Zima Forest in Guinea and Branch & 
Rodel (2003) for areas in western Ivory Coast. 



THE REPTILES OF MT NLONAKO, CAMEROON 



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THE REPTILES OF MT NLONAKO, CAMEROON 1065 

The distribution of species along an elevational gradient shows that most 
species are found at lower elevations (table 3). Only a few species can be considered 
purely montane such as Chamaeleo pfefferi and three skink species {Lacertaspis 
rohdei, Leptosiaphos vigintiserierum, Leptosiaphos spp.). 

In a broader biogeographic context, 51 of the reptile species of Mt Nlonako can 
be considered West / Central African in distribution (table 3). Only two species have a 
pure West African distribution, of which one, Aparallactus niger is recorded here for 
the first time in Cameroon and Central Africa and had its eastern distribution border in 
Ivory Coast. Twenty-one are distributed in Central Africa only. Ten of the recorded 
species are endemic to Cameroon. Eight species, Agama agama, Hemidactylus 
mabouia, Trachylepis maculilabris, Dasypeltis scabra, Lamprophis fuliginosus , Natri- 
citeres olivaceus and N. variegata, and Naja melanoleuca have a wider distribution in 
Sub-Saharan Africa. 

Overall, Mt Nlonako is dominated by a West / Central African reptile fauna. 
This is supported by the relatively high coefficient of biogeographic resemblence 
(CBR) as demonstrated in table 2. This is different to the biogeographic relationships 
of the amphibian fauna which showed closer relations to the Central African amphi- 
bian fauna as compared to the west African amphibian fauna (Herrmann et al., 2005). 
Here, the reptile faunas of the southwest Cameroonian Korup NP and the south 
Cameroonian Dja Faunal Reserve show a high degree of resemblance. As for snakes 
the CBR's exhibited by the Korup NP, the Dimonika region in Congo-Brazzaville, the 
Dja Faunal Reserve and Mt Nimba in West Africa indicate the highest resemblance. 

The exceptional species richness and high degree of endemicity in lizards of Mt 
Nlonako and the West Cameroonian mountain range can be explained by palaeogeo- 
graphic events. Historically this area has served as a refuge during drastic climate fluc- 
tuations. During the Pleistocene circa 20,000 years ago, African wet forests were 
restricted to a few isolated areas (Livingstone, 1982). Those fluctuations and refuges 
played an important role in the evolution of the high number of (endemic) reptile 
species as Lawson (1993) also describes for the Korup NP. 

Osteolaemus tetraspis, Kinixys erosa and K. homeana, all five Chamaeleo 
species, Varanus ornatus, Calabaria reinhardtii and Python sebae are listed in CITES 
appendices for which international trade is regulated. The only species listed as 
threatened is Osteolaemus tetraspis (vulnerable, IUCN 2003). Akani et al. (1998) pro- 
vide further insight into the local trade and consumption of this species in Nigeria; the 
situation in Cameroon can be considered very similar. The two tortoise species Kinixys 
erosa and homeana are listed as data deficient by the IUCN (2003). Lawson (2000) 
provides data on their conservation status in Cameroon. Osteolaemus tetraspis, Python 
sebae and Naja melanoleuca are protected by Cameroonian law. Threats to the Mt 
Nlonako reptile fauna are (1) habitat destruction by logging in the eastern and south- 
ern parts of the area, (2) habitat destruction by human encroachment as notable on the 
northern and western slopes of Mt Nlonako adjacent to the town Nkongsamba and (3) 
the hunting and consumption as food of species like Osteolaemus tetraspis, the two 
Kinixys species and Python sebae. 



1 066 H .-W. HERRMANN ET AL . 

ACKNOWLEDGEMENTS 

We thank ALSCO (American Linen Supply Company) Germany, especially 
Horst Nobis, for their generous financial support during the initial phase of this project 
and the continuation thereafter. The Zoological Garden Cologne, Germany, supported 
the work of H.-W. H.. The Cameroon Ministry of Scientific and Technical Research 
(MINREST) issued research permits and the Ministry of Environment and Forestry 
(MINEF) issued collecting and export permits. The Bundesamt für Naturschutz, Bonn 
granted import permits. We thank the traditional authorities of Mt Nlonako for per- 
mitting us to work in their tribal areas. The Worldwide Fund for Nature (WWF) 
Cameroon and the WWF Mount Kupe Forest Project staff helped with logistics, 
literature and information on local issues. In the field we acknowledge the efforts of 
Mirco Plath, Otto Mesumbe, and many other Cameroonian assistants; their knowledge 
of their forests and the animals therein helped make this survey possible. 



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REVUE SUISSE DE ZOOLOGIE 
Tome 112 — Fascicule 4 

Pages 

Cerny, Milos. Additional notes on the fauna of Agromyzidae (Diptera) in 

Switzerland 771-805 

Golovatch, Sergei I. The millipede family Paradoxosomatidae in 
Paraguay, with descriptions of five new species (Diplopoda, Poly- 
desmida) 807-830 

Yoshizawa, K., Lienhard, C. & Idris, A. B. On the systematic position of 
Podopterocus Banks and Dinopsocus Banks, with a revised 
diagnosis of the genus Sigmatoneura Enderlein (Psocodea: 
'Psocoptera': Psocidae) 831-848 

Yang, Ding & Merz, Bernhard. Review of the species of Platypalpus 
Macquart from Guangxi, China (Diptera, Hybotidae, Tachydro- 
miinae) 849-857 

Ebejer, Martin J. A contribution to the knowledge of the Chyromyidae 
(Diptera) of Italy with description of a new species of Apha- 
niosoma Becker 859-867 

Monod, Lionel & Lourenço, Wilson R. Hemiscorpiidae (Scorpiones) 
from Iran, with descriptions of two new species and notes on 
biogeography and phylogenetic relationships 869-941 

Ivanova, Elena S. & Vaucher, Claude. New genera and species of 
nematode parasites (Drilonematoidea: Ungellidae) from coelomic 
cavity of Neotropic acanthodrilids deposited in the Natural History 
Museum of Geneva, Switzerland 943-961 

Salgado Costas, José Maria. New species of Leiodidae (Coleoptera) and 

new records from the Neotropical Region 963-982 

Merz, Bernhard & Dawah, Hassan A. Fruit flies (Diptera, Tephritidae) 
from Saudi Arabia, with descriptions of a new genus and six new 
species 983-1028 

Schmid-Egger, Christian. Proxiandrena subgen. nov. und Revision der 
west- und zentralpaläarktischen Arten der Andrena proxima- 
Gruppe (Hymenoptera, Apidae) 1029-1044 

Herrmann, Hans-Werner, Böhme, Wolfgang, Euskirchen, Oliver, 
Herrmann, Patricia A. & Schmitz, Andreas. African biodiversity 
hotspots: the reptiles of Mt Nlonako, Cameroon 1045-1069 



REVUE SUISSE DE ZOOLOGIE 

Volume 1 1 2 — Number 4 

Pages 

Cerny, Milos. Additional notes on the fauna of Agromyzidae (Diptera) in 

Switzerland 771-805 

Golovatch, Sergei I. The millipede family Paradoxosomatidae in 
Paraguay, with descriptions of five new species (Diplopoda, Poly- 
desmida) 807-830 

Yoshizawa, K., Lienhard, C. & Idris, A. B. On the systematic position of 
Podopterocus Banks and Dinopsocus Banks, with a revised 
diagnosis of the genus Sigmatoneura Enderlein (Psocodea: 
'Psocoptera': Psocidae) 831-848 

Yang, Ding & Merz, Bernhard. Review of the species of Platypalpus 
Macquart from Guangxi, China (Diptera, Hybotidae, Tachydro- 
miinae) 849-857 

Ebejer, Martin J. A contribution to the knowledge of the Chyromyidae 
(Diptera) of Italy with description of a new species of Apha- 
niosoma Becker 859-867 

Monod, Lionel & Lourenço, Wilson R. Hemiscorpiidae (Scorpiones) 
from Iran, with descriptions of two new species and notes on 
biogeography and phylogenetic relationships 869-941 

Ivanova, Elena S. & Vaucher, Claude. New genera and species of 
nematode parasites (Drilonematoidea: Ungellidae) from coelomic 
cavity of Neotropic acanthodrilids deposited in the Natural History 
Museum of Geneva, Switzerland 943-961 

Salgado Costas, José Maria. New species of Leiodidae (Coleoptera) and 

new records from the Neotropical Region 963-982 

Merz, Bernhard & Dawah, Hassan A. Fruit flies (Diptera, Tephritidae) 
from Saudi Arabia, with descriptions of a new genus and six new 
species 983-1028 

Schmid-Egger, Christian. Proxiandrena subgen. nov. and revision of the 
Andrena proxima group in the western and central Palaearctic 
Region (Hymenoptera, Apidae) 1029-1044 

Herrmann, Hans-Werner, Böhme, Wolfgang, Euskirchen, Oliver, 
Herrmann, Patricia A. & Schmitz, Andreas. African biodiversity 
hotspots: the reptiles of Mt Nlonako, Cameroon 1045-1069 

Indexed in Current Contents, Science Citation Index 



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Penard, E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp. 
Penard, E. 1889. Etudes sur quelques Héliozoaires d'eau douce. Archives de Biologie 9: 1-61. 
Mertens, R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp. 
Handley, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: Wenzel, R. L. & Tipton, V. J. 

(eds). Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. 

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