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The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. |. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address c/o National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Correspondence regarding membership of the society, subscriptions and possibilities for exchange of this journal should be addressed to: Nederlandse Entomologische Vereniging c/o Instituut voor Taxonomische Zoölogie Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands Subscription price per volume Hfl. 300.- (postage included). Special rate for members of the society. Please enquire. Instructions to authors May be obtained from the editors. Graphic design Ontwerpers B.V., Aad Derwort, ‘s-Gravenhage Tijdschrift voor Entomologie Contents of Volume 133 Articles 3 121 27 143 149 197 31 205 239 39 43 245 Asquith, A. & J. D. Lattin Nabicula (Limnonabis) propinqua (Reuter) (Heteroptera: Nabidae): dimorphism, phylogenetic relationships and biogeography. Aukema, B. Taxonomy, life history and distribution of three closely related species of the genus Calathus (Coleoptera: Carabidae). Bedos, A. & L. Deharveng New species of Superodontella Stach (Collembola: Odontellidae) from Thailand. Belle, J. Progomphus nigellus and Phyllocycla hamata, two new dragonflies from Brazil (Odonata: Gomphidae). Belle, J. & J. van Tol Anomalagrion hastatum (Say), an American damselfly indigenous to the Azores (Odonata, Coenagrionidae). Chaudhuri, P. K. & S. Chattopadhyay Chironomids of the rice paddy areas of West Bengal, India (Diptera: Chironomidae). Mastrigt, H. J. G. van New (sub)species of Delias from the central mountain range of Irian Jaya (Lepidoptera: Pieridae). Monk, K. A. & R. K. Butlin A biogeographic account of the grasshoppers (Orthoptera: Acridoidea) of Sulawesi, Indonesia. Nieukerken, E. J. van The Trifurcula subnitidella group (Lepidoptera: Nepticulidae): taxonomy, distribution and biology. Nieukerken, E. J. van Stigmella rolandi sp. n.: a widespread southern European species on Rosa (Lepidoptera: Nepticulidae). Pape, T. Two new species of Sarcophaga Meigen from Madeira and mainland Portugal (Diptera: Sarcophagidae). Pape, T. Revisionary notes on American Sarcophaginae (Diptera: Sarcophagidae). Pérez, T. M. & W. T. Atyeo New taxa of feather mites (Acarina, Pterolichidae) from Megapodes (Aves, Megapodiidae). Quicke, D. L. J. & C. van Achterberg The type specimens of Enderlein’s Braconinae (Hymenoptera: Braconidae) housed in Warsaw. 265 75 269 85 97 273 107 Schouten, R. T. A. A new species of Euchromius Guenée, 1845 (Lepidoptera: Pyralidae: Crambinae) from Australia. Sruoga, V. Seven new species of Elachistidae (Lepidoptera) from the USSR. Stock, J. H. A new species of Austrodecus (Pycnogonida) from New South Wales, Australia. Theowald, Br. & P. Oosterbroek Zur Zoogeographie der Westpaläarktischen Tipuliden. IX. Die Tipuliden des Vorderen Orients. Tol, J. van Key to the Malesian species of Leptogomphus Selys, with the description of a new species from Sabah (Odonata, Gomphidae). Descriptions and records of Malesian Odonata, |. Tol, J. van Zoological expeditions to the Krakatau Islands, 1984 and 1985: Odonata. Des- criptions and records of Malesian Odonata, 2. Wittmer, W. Die Familie Malachiidae (Coleoptera) auf Sulawesi. 43. Beitrag zur Kenntnis der indo-malaiischen Fauna. Book announcements and reviews 96 268 268 268 268 R. Johansson et al., The Nepticulidae and Opostegidae (Lepidoptera) of North West Europe. [R. T. A. Schouten] T. B. Larsen, The butterflies of Egypt. [R. T. A. Schouten] O. |. Merzheevskaya, Larvae of owlet moths (Noctuidae). P. W. Oman, W. J. Knight & M. W. Nielson, Leafhoppers (Cicadellidae): a bibli- ography, generic check-list and index to the world literature 1956-1985. D. J. Williams & G. W. Watson, The scale insects of the Tropical South Pacific region, 3. Referees for volume 133 B. Aukema (Renkum), M. F. Claridge (Cardiff), R. W. R. J. Dekker (Amsterdam), Th. S. van Dijk (Wijster), W. N. Ellis (Amsterdam), H.-J. Geiger (Bern), Th. Heijerman (Wage- ningen), P. J. van Helsdingen (Leiden), L. B. Holthuis (Leiden), O. Karsholt (Copenha- gen), O. A. Saether (Oslo), S. Shinonaga (Tokyo). Dates of publication Volume 133 (1), pages 1-120, 31 July 1990 Volume 133 (2), pages 121-280, i-viii, 14 December 1990 © Nederlandse Entomologische Vereniging, Amsterdam ISSN 0040-7496 iv Volume 133, no. I, 1999 AE Ti üdschrift | voor | Entomologie A journal of systematic and evolutionary [ritomalazy since 1858 Published Bane Nederlandse Ento Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address c/o National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Correspondence regarding membership of the society, subscriptions and possibilities for exchange of this journal should be addressed to: Nederlandse Entomologische Vereniging c/o Instituut voor Taxonomische Zoölogie Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands Subscription price per volume Hfl. 300.- (postage included). Special rate for members of the society. Please enquire. Instructions to authors May be obtained from the editors. Graphic design Ontwerpers B.V., Aad Derwort, 's-Gravenhage Editorial NINE te Jen) le A, The editors of the ‘Tijdschrift voor Entomologie’ en the first issue of volume 133, complete with a new cover design and a reshaped graphic page design. Both the cover and the lay-out were designed by Aad Derwort of the Dutch firm ‘Ontwerpers’, The Hague. The cover depicts a variety of insects from a collection of illustrations for graphic designers and illustrates the variety of topics covered. The logo of the Netherlands Entomological Society can still be found on the new cover, albeit somewhat smaller. This logo, in use since 1953, depicts the Large copper (Lycaena dispar batavus Oberthür) on top of its greatly enlarged egg. The logo was introduced on the cover of volume 96 when the late Alex Diakonoff became the managing editor. The lay-out of the articles has undergone relatively small changes compared with previous issues. ONTWERPERS he rchitectonische vormgeving The new design parallels changes in the editorial policy of the ‘Tijdschrift’ to transform it into an international journal of systematic and evolutionary ento- mology. Although the journal has been published for an international audience for several decades, the Dutch language was still used on the cover and elsewhere. This has now all been translated into English, except the journal's title which has become too well-known to be changed. Furthermore the editors now explicitly express preference for articles on the insect fauna of the Palaearctic and Indo- Australian regions. Papers on other regions are in principle only accepted if space allows or if the paper deals with a topic of broader interest. Some changes have gradually been introduced in the last three volumes. The system of publishing a number of issues per year with one article each was left in favour of two issues with various contents. This has reduced the expenses and has given the opportunity to accept also short papers and book reviews. The journal will usually be published around June Ist and December Ist. The editors regret that the present issue appears some weeks late. As can also be seen in this issue, the ‘Tijdschrift’ is certainly not only a medium for Dutch entomologists or for members of the Netherlands Entomological Society alone. Authors from all countries are cordially invited to submit their papers. Only quality and consistency with the scope play a role in acceptance, although we would certainly welcome new members of the society. It is general policy now to invite external referees to review the manuscripts with regard to their scientific quality and soundness. A new set of instructions for authors will be published in the next issue and a guide for preparation of manuscripts on diskette will be distributed on request. The ‘Tijdschrift voor Entomologie’ has a long tradition and is certainly one of the oldest entomological journals still being published. The journal has been published annually since 1858, accounting for a total of 132 volumes up to now. In the early days mainly the Dutch language was used. Later French, German and English gradually started to play a role. In the second half of this century Dutch lost its position as a scientific language and disappeared from the journal. From the start the journal has especially served as a medium for systematic entomology and, until the fifties, also for Dutch faunistics. Prior to 1953 it also included the proceedings of the meetings of the Netherlands Entomological Society. Faunistic papers, articles in the Dutch language and smaller papers of international interest are now being published in 'Entomologische Berichten’ (Entomological Reports), another society journal. Other publications of the Society are the international journal "Entomologia experimentalis et applicata’ and the monograph series ‘Monografieën van de Nederlandse Entomologische Vereniging’. The editors hope that the new face of the journal will please the readers and especially that the contents, which is after all the most important part of a journal, will continue to find their way to the professional and amateur entomol- ogist. Erik |. van Nieukerken Jan van Tol [editors] A. W. M. Mol R. T. A. Schouten [co-editors] A. ASQUITH & J. D. LATTIN Systematic Entomology Laboratory, Department of Entomology, Oregon State University NABICULA (LIMNONABIS) PROPINQUA (REUTER) (HETEROPTERA: NABIDAE): DIMORPHISM, PHYLOGENETIC RELATIONSHIPS AND BIOGEOGRAPHY Asquith, A. & J. D. Lattin, 1990. Nabicula (Limnonabis) propingua (Reuter) (Heteroptera: Nabidae): dimorphism, phylogenetic relationships and biogeography. - Tijdschrift voor Entomologie 133: 3-16, figs. 1-15, tabs. 1-3. [ISSN 0040-7496]. Published 31 July 1990. The taxonomy and distribution of the North American species Nabicula propinqua (Reuter), is reviewed. The macropterous form occurs in both sexes, but is much less common than the brachypterous form. Macropterous forms of both sexes have wider pronota. Males have disproportionately longer antennae than females. Male and female genitalia of N. propinqua are described and illustrated. The genitalia of individuals from isolated west coast populations differ only slightly from those of eastern populations. A cladistic analysis of the subgenus Limnonabis Kerzhner is presented and a new species-group classification sug- gested. A biogeographic hypothesis explaining the present distribution of Limnonabis is presented. We suggest that N. propingwa is a vicariant species that was isolated from a population contiguous with that in western Europe with the opening of the Atlantic Ocean. Correspondence: Adam Asquith, Systematic Entomology Laboratory, Department of Ento- mology, Oregon State University, Corvallis, OR 97331, USA. Key words. - Nabidae, sexual dimorphism, wing dimorphism, biogeography, phylogeny. INTRODUCTION Nabicula (Limnonabis) propinqua (Reuter), re- cognized by its large size and elongate, slender form, is one of the most conspicuous North Amer- ican species of Nabidae (fig. 1). Unlike some of the commonly encountered species of Nabis that occur on vegetation in a variety of habitats, N. propinqua is rare in collections and lives on the ground or low vegetation in marshy habitats (Blatchley 1926). In addition to N. propinqua, five other species in the genus Nabicula Kirby occur in North America (Henry & Lattin 1988). N. propinqua, however, is the only member of the subgenus Limnonabis Kerzhner in North America, the other five species occur in Europe and Eastern Asia. The most recent nomenclature (Kerzhner 1988) considers Nabicula a subgenus of Nabis, and does not recognize Limnonabis as a taxonomic group. In this paper, however, we use the nomenclature of Kerzhner (1981) because this classification has also been used in recent faunistic works for Europe (Pé- ricart 1987) and North America (Henry & Lattin 1988). In addition, species relationships within Na- bicula are still unresolved and we anticipate the nomenclature of Kerzhner (1988) to change again (unpublished data). Reuter (1872) described N. propinqua in Nabis Latreille from a single brachypterous female spe- cimen from Wisconsin, and described vicarius (in Nabis) from a single brachypterous male from Il- linois. Reuter (1880) synonymized vicarius with propinqua. Hart (1907) described elongatus (in Na- bis) from a single macropterous male from Havana, Illinois. This species was synonymized with propin- qua by Van Duzee (1916). In this paper, we summarize the distribution and habits of N. propingua and describe the internal genitalia for both the male and female. We also present a cladistic analysis of the subgenus Limno- nabis and provide a hypothesis of the biogeographic history of the taxon. DESCRIPTIVE PART Distribution Nabicula propinqua occurs along the eastern sea- board from Maryland to Maine and extends west- ward between 40 and 50 degrees N latitude throughout the Great Lakes region to North Da- kota (fig. 2). West of the 100th meridian, N. pro- pinqua occurs much farther north, with records N d TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 from Alberta, Manitoba, Quebec and above 60 de- grees N latitude along Great Slave Lake in the Northwest Territory. Essentially the distribution of N. propinqua follows the Saline Lakes and Forest Zone of Freshwater Lakes limnological regions of Northcate & Larkin (1966). There are at least two populations of N. propingua that are disjunct from the main distribution. We have collected N. propin- qua from two localities in the coastal marshes in Oregon and have examined a single specimen from Meade Co., Kansas. With the exception of the latter three records, most of the known distribution of N. propinqua occurs within the area occupied by the ice sheet of the Wisconsin Glaciation. If N. propinqua was present in North America during the Pleistocene, as we will argue, it must have been restricted to areas south of the ice sheet or in one of the far northern refugia (Matthews 1979). Because N. pro- Fig. 1. Nabicula propingua (Reuter). Brachypterous male. Dorsal habitus. pinqua is not yet known to occur in the areas of northern glacial refugia, it seems likely that N. propinqua moved into the areas it now occupies from areas south of the Wisconsin ice sheet some- time within the last 10,000 years. In this scenario, the Kansas specimen probably represents a Pleis- tocene relictual population in an area previously occupied by N. propinqua. Other such populations are likely to be located (e.g. the marshes in the Sand Hills of western Nebraska). The origin of the Oregon coast population is more perplexing. N. propingua has not yet been found in the arid Inter-mountain region, nor in the mesic Willamette Valley and Puget Trough areas of Oregon and Washington. It appears to be restricted to the coastal marshes of Oregon and possibly Washington. Lattin (1966) suggested that this po- pulation might be an introduction from the eastern United States. Now, however, we believe this un- likely, considering the restricted habitat of this spe- cies and the fact that the Oregon population dis- plays a slightly different genitalic structure than eastern populations (see below). If N. propinqua or its direct ancestor was present in North America by the Eocene as we hypothesize (see Discussion), there were few barriers to impede Fast-West movement and a contiguous distribution across the northern latitudes would have been pos- sible. Beginning in the Miocene, orogenic activity in western North America resulted in topographical geographic barriers and, perhaps more impor- tantly, increased aridity in the Intermountain re- gion. Because N. propinqua lives in moist, riparian habitats, this dramatic change in climate and terrain would have restricted mobility and could have led to the extinction of many, or all of the intervening populations. This is merely a hypothesis of course, and we cannot actually date the separation of the west coast population and it is possible that it is a much more recent event. The climate of the northern United States during the Pleistocene periods was much more mesic than the present, with abundant, large shallow lakes and marshes throughout the Great Basin province (Smith 1978). This would have pro- vided abundant habitats for N. propingua from the Rocky Mountains west. With the advent of the Hypsithermal and higher temperatures, these hab- itats in this area largely disappeared (Barnosky et al. 1987), which could have left the coastal popula- tion isolated from those north and east of the Rock- ies. This hypothesis would be corroborated if relic- tual populations of N. propingua were found in isolated marshes in the northern Great Basin. Dimorphism In many nabids, males and females often differ markedly in size, occurrence of wings and propor- ASQUITH & LATTIN: Nabicula propingua ? / SCALE \ \ | ° o__ioo 200 300 400 500 600 700 800 900 1000 MILES IS 6” 200° 400 600 800 1000 1200 1400 KILOMETERS | LAMBERT’S AZIMUTHAL EQUAL-AREA PROJECTION © \ | | | 120 | 110 fl 90 x WEST LONGITUDE Fig. 2. Distribution of N. propingua in North America. Circles represent specimens examined, triangles are literature records only. Solid line indicates maximum extent of Wisconsin ice sheet. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 2.2 PRONOTAL WIDTH (mm) 9 10 11 12 13 TOTAL LENGTH (mm) Fig. 3. Relationship between the posterior width of the pronotum and total length in N. propinqua. y = 0.093239(x) + 0.465369, r? = 0.565. Open squares are brachypterous specimens. Solid squares are macropterous specimens. 29 TONDE 2.8 n 241 2.6 2.5 2.4 2.3 LENGTH ANTENNAL SEGMENT II (mm) 2.2 Wo Lia cs a la. AI 9 10 11 12 13 TOTAL LENGTH (mm) Fig. 4. Relationship between the length of the second antennal segment and total length in N. propinqua. Open squares are males. y = 0.13069(x) + 1.318405, r? = 0.149. Solid squares are females. y = 0.077957(x) + 1.62609, r? = 0.101. 6 ASQUITH & LATTIN: Nabicula propingua Table 1. Comparison of six measurements among brachypterous and macropterous male and female Nabicula propinqua. Data are mean and (range) in mm. ALE. VERT PPW PL Al A2 male 9.8 0.405 1:39 152 1.66 2.60 (9.1-10.6) (0.38-0.44) (123-151) (1.41-1.69) (1.51-1.85) (2.28-2.86) Brach M 10.5 0.41 1.97 1.69 1.70 2.69 female 11.8 0.444 1.57 1.73 1.65 DSS (11.0-13.2) (0.39-0.48) (1.39-1.86) (1.61-1.86) (1.53-1.85) (2.36-2.82) Brach F 11.8 0.41 2.09 1.86 1.60 2.49 (10.3-12.1) (0.40-0.42) (2.04-2.14) (1.83-1.89) (1.59-1.60) (2.48-2.50) tions of body parts, but rarely are these differences quantified. We examined sexual dimorphism in six measurements, total length (TL), length of anten- nal segment one (A1), length of antennal segment two (A2), width of the vertex (VERT), posterior pronotal width (PPW) and pronotal length (PL). Measurements using an ocular micrometer were made of 21 males and 32 females from throughout the range of N. propinqua. We found differences between sexes in four of the six measurements. TL, VERT, PPW and PL, were greater in females than in males, but there were no differences between males and females for Al and A2 (table 1). The differences between sexes seen in the former char- acters are explained by the positive linear relation- ship between these characters and the absolute size of the individual. For example, PPW increases with TL, and because females achieve a greater TL than males, they also display a greater PPW (fig. 3). We do not consider these characters true sexual dimor- phisms therefore, but only size dependent charac- ters. The lengths of antennal segments also displayed an increase with TL, but only within a given sex. This is demonstrated by the fact that the slopes of the A2-TL regressions for the two sexes are the same (F = 0.415, P = 0.523), but the Y intercepts are different (F = 8.662, P = 0.005) (fig. 4). This indicates that although females are larger than males (greater TL), small males have the same length antennae as small females and large males have the same length antennae as large females. Thus, in males, the antennae are disproportion- ately longer than in females. The length of the antennae seems to be uncoupled from the general correlation with body size that other measurements show. This relationship does not appear to be com- mon in the Nabidae. In a preliminary analysis of four species of Nabis, males did not have dispropor- tionately longer antennae, but rather there was a linear relationship between antennal length and size across the sexes. Macroptery Nabicula propinqua usually occurs in the bra- chypterous state, with the apex of the fore wings reaching to the middle of the second abdominal ter- gite (fig. 1). Macropterous females have been re- ported by Harris (1928) and Froeschner (1971). On- ly 2 of 45 females we examined were macropterous. In these individuals, the fore wings reached the mid- dle of the seventh abdominal tergite. The hind wings are equally well developed, reaching to the anterior margin of the seventh abdominal tergite (fig. Sa). Fig. 5. N. propinqua A. Macropterous female. B. Macrop- terous male. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Fig. 6. Seminal depository of female N. propinqua. Dorsal view. A. Sclerotized ring. B. Membranous sac. C. Muscular lobe ventral to oviducts. We have examined 29 male specimens and have seen only one macropterous individual (fig. 5b). This specimen was the holotype of Nabis elongatus Hart and is deposited in the Illinois Natural History Survey collection (INHS). Although Harris exam- ined the material in the INHS (we have seen spe- cimens with his determinations), he apparently overlooked this specimen, because he stated (Harris 1928) that males may never occur in the macrop- terous state. In England and on the Continent, only the females of Nabicula lineata (Dahlbom) are known in the macropterous form (Southwood and Leston 1959). Macropterous females and males have been reported for N. ussuriensis (Kerzhner) in eastern Asia (Kerzhner 1968). Differences in the structure of the pronotum between brachypterous and macropterous speci- mens have been reported for both the latter two species. In N. /ineata, the pronota of brachypterous specimens are 1.1 times as long as wide and in the macropterous forms the length and width are equal (Pericart 1987). In N. wssurtensis, the pronotum in brachypterous specimens is 1.1-1.2 times as long as wide and only 0.94 times as long as wide in macrop- terous specimens (Kerzhner 1968). In N. propinqua, the ratio of PL to PPW is lower in the macropterous form (0.85-0.89) than the bra- chypterous form (1.05-1.15) in both males and fe- males. The lower ratio results not from a shorten- ing of the pronotum in the macropterous form, but from an increase in the width (fig. 3). In particular, the posterior lobe of the pronotum is greatly flared. 8 The increased width of the pronotum is probably an indirect effect of the development of flight muscles and phragmata in the pterothorax of the winged form (Darnhoffer-Demar 1969). Because the ante- rior end of the mesothorax is enclosed by the pos- terolateral angles of the pronotum, an increase in diameter of the mesothorax would cause a similar change in the posterior lobe of the pronotum. Female Genitalia The seminal depository is large and dome- shaped, consisting of a fleshy external portion and a partially sclerotized internal structure, the apex of which bears three to five blunt sclerotized teeth (fig. 6). A thick muscular structure lies between the base of the depository and the oviduct. The sclerotized ring is a single structure occupying the ventral sur- face of the left side of the depository, extending from the depository laterally and curving ante- riorly. It is surrounded by a large membrane also directed anteriorly. Male Genitalia Parameres Reuter (1872) and Harris (1928) provided lateral views of the left paramere of Nabicula propingua. Paramere morphology in some groups of Nabidae is very conservative (e.g. the genus Nabis) and N. propingua displays this general form (fig. 7). Within the subgenus Limnonabis, the paramere of N. propinqua most closely resembles that of N. pontica. In contrast, N. lineata displays an unusually elongate apex. In some specimens of N. propingua, the apex of the paramere is slightly bent laterally, a condition which is strongly developed in N. ussu- riensis (Kerzhner) and N. demisa (Kerzhner 1968, 1981). b Fig. 7. Right paramere of N. propingua A. Lateral view B. Medial view. C. Ventral view. Fig. 8. Male aedeagus of N. propinqua from Oregon. Sclerites indicated by roman numerals. IVa. sclerite IV of specimen from Illinois. Aedeagus The aedeagus of Nabicula propinqua is very sim- ilar to that of N. lireata and N. pontica. The mem- branous portion of the aedeagus has four sclerites (fig.8). No nomenclature has been proposed for the aedeagal sclerites of the Nabidae, and here we number the sclerites with Roman numerals begin- ning at the base of the aedeagus (sclerite I) and moving to the apex (sclerite IV). Sclerite I is linear and parallel to the longitudinal axis of aedeagus; it has a wide base and slightly tapering distalarm with a blunt, rounded apex. Some specimens may display a slight flange on this sclerite. Sclerite II is pistol- shaped, with a wide, arcuate base, a curved obtuse angle near its midpoint and a tapering, rounded distal arm. The distal arm bears flanges on both edges reaching from angle to apex. Sclerite III is roughly quadrangular and distinctly comb-shaped, with five short, blunt teeth directed toward the midline of the aedeagus. Sclerite IV is situated transversely in the aedeagus, with a broad, oval base. This sclerite exhibits geographic variation. In specimens from the Oregon coast, the sclerite has a thick, bluntly rounded arm (fig. 8 IV), and in specimens from east of the Rocky Mountains, the arm is longer, thinner, with a more pointed apex. (Fig. 8 IVa). There are two areas with sclerotized ASQUITH & LATTIN: Nabicula propinqua denticles along the folds of the membrane, one area distal to sclerite II and the other distal to sclerite IV. These denticulate areas are also present in identical form in related species. PHYLOGENETIC ANALYSIS To understand the origin and evolution of Nabz- cula propinqua in North America, it is necessary to identify its phylogenetic relationship to other taxa. Within Nabicula, Kerzhner (1981) recognized three subgenera, Limnonabis Kerzhner, Dolicho- nabis Reuter, and Nabicula Kirby. He distinguished the subgenus Limnonabis from Dolichonabis (sensu strictu) by the diverging posterior lobe of the head, recurved connexivum of the male and the presence of three or four sclerites in the aedeagus. Within Limnonabis, he identified two groups, the lineata group composed of N. lineata, N. pontica and N. propingua, identified by the large parameres and multiple rows of spines of Ekblom’s organ (see fig. 11). The wsswriensis group contains N. ussw- riensis (Kerzhner), N. demisa (Kerzhner) and N. sauteri (Poppius), united by the small parameres and the single row of spines (see fig. 10). To further clarify the relationships among these taxa, we conducted a cladistic analysis of the subge- nus Limnonabis using the computer program HENNIG 86 (Farris 1988). Because we were not able to examine specimens of all species in the subgenus, we used characters that were described and/or illus- trated for the other taxa by Kerzhner (1968, 1981) and Péricart (1987). We used structures of the male Anaptus flavomarginata limbata sauteri demisa ussuriensis propinqua lineata — pontica Limnonabis Fig. 9. Cladogram of Limnonabis species based on table 2. Nodes denoted by letters in circles. Characters and character states of synapomorphies given under each node. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 10 ines. , multiple rows of sp . propinqua BEN ines le row of spi , sing Ekblom's organ. 10. N. americolimbata ing iew of male genital capsule show 10L V Figs. 10-11. Poster Figs. 12-13. Structure of individual spines of Ekblom's organ. 12. N. americolimbata. 13. N. propinqua. ASQUITH & LATTIN: Nabicula propinqua Table 2. Description of characters used in the cladistic analysis of the subgenus Limnonabis. Character states are preceeded by their coding in the data matrix and (in parentheses) the number of times state arises in the cladogram. Female Genitalia 1. 0(1) Double sclerotized ring. 1(1) Single sclerotized rings. 2. 0(1) Sclerotized ring symmetric, located medio- dorsal. 1(1) Sclerotized ring asymmetric, located laterally. 3. 0(1) Membranous “sac” around sclerotized ring absent. 1(1) Membranous "sac symmetric, located dor- sally. 2(1) Membranous sac” asymmetric, not located dorsally. 4. 0(1) Muscular lobe” antero-ventral to sperm ducts absent. 1(1) Muscular “lobe” antero-ventral to sperm ducts present. Male Aedeagus 5. 0(1) Aedeagus with fewer than three sclerites. 1(1) Aedeagus with 3 sclerites. 2(1) Aedeagus with 4 sclerites. 3(1) Aedeagus with more than 4 sclerites. 6. O(1) Length of sclerite I less than four times its width at middle. 1(1) Length of sclerite I at least four times its width. 7. 0(2) Sclerite I without flanges present. 1(1) Sclerite I with flanges. 2(1) Sclerite I with flanges only slightly deve- loped. 8. 0(1) Sclerite II absent. 1(1) Length of sclerite II less than three times its width. 2(1) Length of sclerite II three times its width. paramere (2 characters), aedeagus (6 characters), male genital capsule (1 character), female genitalia (3 characters), body structure (4 characters) (table 2). Determining the polarity of some of the char- acters was difficult, because the homology of some of the structures in other genera could not be deter- mined and no previous cladistic analysis within the Nabidae was available for comparison. Because of these problems, we included the following three taxa in our analysis as outgroups, Nabicula (Dolichonabis) limbata (Dahlbom); Nabicula (Nabicula) flavomarginata (Scholtz); Anaptus ma- jor (A. Costa) (table 3). These taxa were chosen because they represent the other two subgenera of Nabicula and a more distant member of the tribe Nabini. All multistate characters were coded as or- dered except characters 5 and 10 because we were uncertain of their transformation sequences. The analysis produced a single tree of minimal length (31 steps, consistency index of 83.0, fig. 9). The subgenus Limnonabis appears to be a mono- phyletic group, as indicated by component E of the cladogram. It is identified by characters 14 and 15, the diverging posterior margin of the head, and the recurved connexivum of the male respectively. 9. 0(1) Sclerite II without flanges. 1(1) Sclerite II with flanges. 10. 0(1) Sclerite III linear without flanges. 1(1) Sclerite III prong-shaped. 2(1) Sclerite III comb-shaped. 3(1) Sclerite III short, not linear. 4(1) Sclerite III linear with flanges. 11. 0(1) Sclerite IV absent. 1(1) Sclerite IV with recurved arm. 2(1) Sclerite IV comb-shaped. Male Paramere 12. 0(1) Tip of paramere not elongate. 1(2) Tip of paramere elongate. 13. 0(1) Apex of paramere straight. 1(2) Apex of paramere bent laterally. External Structure and Form 14. 0(1) Head straight or converging behind eyes. 1(1) Head diverging behind eyes. 15. 0(1) Medial half of male connexivum straight. 1(1) Medial half of male connexivum curved un- der and appressed to abdomen. 16. 0(1) Spines of Ekblom’s organ in a single linear row. 1(1) Spines of Ekblom's organ in a bunched row. 17. 0(1) Body length less than 5 times the width. 1(1) Body length greater than 5 times the width. These are two of the characters Kerzhner (1968) used to define the subgenus. We are doubtful of the integrity of the first character, the diverging poster- ior lobe of the head. We have examined species in other genera (Nabis Latreille, Lastomerus Reuter) in which this character displays almost the same development as seen in Limnonabis. At this time, we can find no genitalic characters that unite all species currently placed in Limnonabis and the only character that we feel defines the group is the re- curved connexivum of the male. The wssuriensis group does not appear to be a natural one, its members are united only by plesi- omorphic characters, such as the single row of spines of Ekblom’s organ. Nabicula ussuriensis is actually united with the /neata group by the pres- ence of an asymmetric sclerotized ring (2-1) (char- acter-character state), presence of a membranous “sac! around sclerotized ring (3), aedeagus with four sclerites (5-2), length of sclerite II three times its width (8-2) and sclerite II with flanges (9-1) (component C). The /neata group of Kerzhner (1968) is clearly monophyletic, represented by com- ponent B on the cladogram and defined by the asymmetric membranous sac (3-2), muscular lobe Il TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 3. Character matrix for Limnonabis processed by HENNIG86. l 2 5) 4 5 6 7 8 9) JO) iil 2 15 14 15 16.17 Anaptus t © @ @ SOON cane = JO) te 122 ? 1 © © © © 0 flavomarginata O © © © OA EE QE ? 2 © i 0 © © @ © limbata l © © 0 Od. 1 lame? ? CE) LI ® © © © »@ sauteri Lr @ © 8 1 1 TOO ANO Om 200 3] Io 30 demisa L @ Orr © ROME Oso ORT er Od 1 TO OS OI usseriensis 1 1 le 2 1 1 2 dm ERD 0 1 1 LAON, dl propinqua 1 1 2 1 2 an 2 1 2 1 @ O 1 1 1 lineata 1 1 2 l 2 1 0 2 Il 0 1 1 0 l Il Il Il pontica 1 le 2 1 2 OZ OR I ® @ A 1 1 1 behind the sperm ducts (4-1), and the bunched row of spines comprising the Ekblom’s organ (16-1). The latter character appears to be unique among the Nabidae. Nabicula propinqua, N. lineata, and N. pontica display a bunched row, 3-5 spines wide, on each side of the anal tube (fig. 11). All other species of nabids that we have examined possess only a single row of linearly arranged spines (fig. 10). The structure of the individual spines may also prove to be an informative character. In N. propin- qua, the spines are thin, with the distal third sharply narrowed, sinuous, with the apex curved laterally (fig. 13), while in N. (Limnonabis) americolimbata the spines are wider, with the distal halves flattened and expanded (fig. 12). Nabicula lineata and N. pontica are united by having sclerite III linear and without flanges (10-0) and sclerite I without flanges (7-0). This latter char- acter is homoplasious, because the 0 state also ap- pears in N. demisa. BIOGEOGRAPHY All members of the subgenus Limnonabis, with the exception of Nabicula propinqua, are found in the Palaearctic region. The three species arising i i SCHOOL SERIES Outline Map WORLD - Scale of Milen at Equator » iP A 1000 2000 3000 MAP NO. 1739 + + — del | SC — | COPYRIGHT * AMERICAN MAP CO., INC. WORLD MAP CO., INC. mn R 1 100, 120, 160, 10 180 160 140 120 100 Fig. 14. Distribution of the subgenus Limnonabis. Vertical lines and solid circles in North America, N. propingua. Large dots in Europe, N. lineata. Small dots in southeastern Europe, N. pontica. Slanted lines with small dots in southeast Asia, composite ranges of N. ussuriensis, N. demisa and N. sauteri. 12 lineata (Europe) pontica (Europe) Eocene) —————— propinqua (N Am) LR, ——— Cretaceous ussuriensis (NE Asia) Fig. 15. Cladogram of Limnonabis with the present dis- tribution indicated after each species name. Hypothes- ized ages of intercontinental vicariances indicated at left. from the cladogram first, N. ussuriensis, N. demisa and N. sauteri occur in far eastern Asia (fig. 14). Two members of the /ineata group, N. lineata and N. pontica occur in Europe and the Mediterranean region and N. propingua occurs only in North America. With regard to the views of Ball (1975), we generated a testable biogeographic hypothesis to explain the present distribution of the members of the monophyletic group represented by compo- nent C on the cladogram. We chose to use only this group and not all of Limnonabis, because N. sauteri and N. demisa are primitive taxa whose placement in the subgenus appears tenuous. We hypothesize that the distributions of the spe- cies in component C are the result of vicariance caused by the breakup of Laurasia beginning in the Cretaceous. With the development of the Turgai straits in the late Cretaceous, Asiamerica and Eur- america were separated into two distinct land masses (Cox 1974). This would have resulted in the ancestral population being divided into two groups, with the ancestor of N. ussuriensis in western Asi- america and the ancestors of the /ineata group in Euramerica. The ancestor of the /ineata group was distributed throughout Euramerica and with the opening of the North Atlantic eventually separating North Amer- ica from Europe in the Eocene, the ancestor of N. propinqua was isolated from that of N. /imeata and N. pontica. The exact time of separation of these two groups cannot be known because there were several distinct land bridges that closed at different times, with at least one dispersal route across the Thulean bridge which may have persisted into the Miocene 20 millions years ago (Noonan 1988). With the exception of the elongate apex of the paramere in N. lineata, N. lineata and N. pontica are very similar, suggesting that the separation of these two species is a more recent event. In summary, we invoke two vicariant events to ASQUITH & LATTIN: Nabicula propinqua explain the origin and present distribution of the taxa in component C of the cladogram (fig. 15). 1) The ancestor of N. ussuriensis was isolated in Asiamerica from component B in Euramerica by the Turgai Straits. 2) The ancestor of N. propinqua was isolated from component A in Europe by the opening of the Atlantic. If our phylogenetic hy- pothesis is correct, we believe this biogeographic scenario to be the most parsimonious one. A disper- sal hypothesis for the evolution of component C would require the ancestral form to have moved into, and then gone extinct from an area reaching from Eastern Europe to Eastern Asia. Likewise, dispersal of the ancestor of N. propinqua from Europe to North America via Beringia as suggested by Schaefer & Calabrese (1980), would require the extinction of this taxon in all of northern Asia. Another hypothesis for propinqua reaching North America is by dispersal across the North Atlantic. Because of the specialized habits, low incidence of macroptery and low vagility of this species (it has not been recorded from flight traps or aerial sam- pling), we consider the latter hypothesis unlikely. DISCUSSION Using examples from the Trichoptera, Homop- tera and Coleoptera, Allen (1983) discussed the North America - Europe - Northeast Asia distri- butions for insects. He found a common pattern among these groups in which the North American and European taxa shared a common ancestor after the origin of the Northeast Asian groups. Similar to our hypothesis for Limnonabis, Platnick (1976) discussed the vicariant patterns in the spider genus Callilepis (Gnaphosidae). His phylogenetic analysis of the genus showed that species groups were dis- tributed in areas that reflected the breakup of Laur- asia. He also suggested that the Turgai straights separated a Asiamerica group from a Euramerica one, and the Atlantic rift further separated the Euramerica group. The species in the N. /ineata group are very similar morphologically, differing primarily in the structure of the internal genitalia. It may seem un- usual that we invoke such great ages for the sepa- ration of these species in light of the slight degree of morphological divergence. Without rehashing the old and continuing debate regarding rates of evolution, we feel that our hypothesis is tenable for three reasons. First, with few exceptions (e.g. Na- bicula subcoleoptrata Kirby) the lineages within the tribe Nabini appear to be highly conservative or canalized in their morphology. Many species differ only slightly in external morphology and are distin- guished primarily by the male aedeagus and the seminal depository of the female. Second, explosive speciation and morphological evolution is often correlated with changing envi- TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 ronments or lineages radiating into new adaptive landscapes (Simpson 1944), while groups occupy- ing stable, constant and predictable environments may undergo very little change through time. The lineata group inhabits moist, marshy habitats and secondarily, estuarine marshes, environments that prevailed during the Cenozoic but also have per- sisted relatively unchanged to the present. Consid- ering the stability of morphology in the group, low vagility and their specific, unchanged habitat, large scale vicariant events resulting in superficially sim- ilar, disjunct species might be expected. Finally, it is becoming increasingly apparent that in some groups of insects, extant species are of great antiquity. In the Coleoptera for example, the Pleis- tocene climatic oscillations greatly altered distribu- tions but did not result in speciation (Coope 1970, Matthews 1977). Some species of beetles are appar- ently in excess of 10 million years old (Larsson 1978). Similar examples are available for the Heter- optera, (Calabrese 1978, 1980). ACKNOWLEDGMENTS We thank Michael D. Schwartz for critically read- ing this manuscript and for help in the data analysis and Bonnie B. Hall for the fine habitus illustrations. We thank the following for their assistance in ex- amining critical specimens: Thomas J. Henry, Uni- ted States National Museum of Natural History (USNM); Randall T. Schuh, American Museum of Natural History (AMNH); Kathryn C. McGiffen, Illinois Natural History Survey (INHS); Charles A. Triplehorn, Ohio State University; Philip J. Clausen, University of Minnesota (UM); B.J. Har- rington and S. Krauth, University of Wisconsin- Madison (UWIM); Robert W. Brooks, University of Kansas (KU); John A. Chemsak, University of California, Berkeley (UCB); Paul H. Arnaud Jr., California Academy of Sciences (CAS); James Lieb- herr and E. Richard Hoebeke, Cornell University (CU); Frank Mead, Florida State Collection of Ar- thropods (FSCA); Jeffrey C. Burne, University of Wyoming; Edward U. Balsbaugh Jr., North Dakota State University; Brett C. Ratcliffe, University of Nebraska. REFERENCES Allen, R. T., 1983. Distribution patterns among arthrop- ods of the North Temperate Deciduous Forest biome. — Annals of the Missouri Botanical Garden 70: 616- 628. Ball, I. R., 1975. Nature and formulation of biogeographi- cal hypotheses. — Systematic Zoology 24: 407-430. Barnosky, C. W., P.M. Anderson & PJ. Bartlein., 1987. The northwestern U.S. during deglaciation: vegetational history and paleoclimatic implications. — In: W. F Rud- diman and H.E. Wright (eds.). 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Chech List of the Hemiptera (Excepting the Aphididae, Aleurodidae and Coccidae) of America, North of Mexico. - New York Entomolo- gical Society, New York, 111 pp. Received: 18 Seprember 1989 Accepted: 1 March 1990 ASQUITH & LATTIN: Nabicula propinqua APPENDIX Literature Records Canada. — Alberta: Edmonton (Strickland 1953). United States. — Illinois: Cook Co.: Argo; West Pull- man (Blatchley 1926). - Iowa: Hancock Co.: 8 mi. SE Britt, July 6, 1928 (Hendrickson 1930). — Maryland: Somerset Co.: Deal Island, June 29, 1970 (Froeschner 1970). - Michigan: Berrien Co.: New Buffalo & Stevens- ville (Hussey 1922b). - New Jersey: Morris Co.: Snake Hill; 2? Co.: White's Bay, July 20, 1914 (Froeschner 1971). - North Dakota: Ramsey Co.: Devils Lake, July 22 & 25 (Hussey 1922a). - South Dakota: Day Co.: Waubay, August 21, 1924 (Harris 1943); Grant Co.: Bigstone, August 20, 1924 (Harris 1943). Specimens Examined Canada. — Alberta: High Prairie, July 17, 1961, A. R. Brooks, 1 6, 1 9 (OSU); Slave Lake, August 14, 1924, O. Bryant, 2 4, 1 9 (CAS). - Manitoba: Red Deer R., August 3, 1937, C. L. Johnston, 1 & (KU); The Pas, August 11, 1937, D. G. Denning, 1 9 (UM). - North West Territory: 5 mi. SE Ft. Providence, August 15, 1965, J. & W. Ivie, 1 2 (AMNH). - Ontario: Thessolon Lake, shore, July 21, 1965, J. & W. Ivie, 19 (AMNH). - Quebec: Quinze Lake, August 15, 1907, W.J. Palmer, 1 Q (CAS); Saskatchewan: Qu’ Appelle River, N Tuxford, July 29, 1965, J. & W. Ivie, 292 (AMNH). United States. — Illinois: Boone Co.: Belvidere, June 27, 1955, J.A. Slater (OSU); Fulton Co.: Havana, June 9, 1905 / Ill. Sands, Hart Coll. / Nabis elongatus Hart, type / Reduviolus elongatus Hart / TYPE Nabis elongatus, C. A. Hart, 1 & (INHS); Lake Co.: Fox Lake, August 23, 1944, Frison & Ross, 4 6,9 2 (INHS); Lake Villa, swamp, August 10, 1906 / Reduvolius vicarius Reuter, micr. F / Nabis propinquus Reuter, H. M. Harris / Van Duzee Cat. No. 825 Det. Harris, 1 9 (INHS); Waukegan, May 14, 1930 4 & (INHS); Waukegan, beach, August 23, 1906 / Reduvolius vicarius Reut, Micro. M / Nabis propinquus Reuter, Det. H. M. Harris / Van Duzee Cat. No. 825, Det. Harris 1 @, | nymph (INHS). - Kansas: Meade Co: September 13, 1944, R. H. Beamer, 1 & (OSU). - Maine: Sagadahoc Co.: Popham Beach, September 4, 1920, A. P. Morse, 1 9, (AMNH). - Massachusetts: Essex Co.: Beach Bluff, 22 June, 1914, H. M. Parshley, 1 nymph (CAS); 13 August, 1914, ex. Carex, 2 8,4 9; June 21, 1915, 1 nymph (CAS); August 17, 1916, ex. Carex, 3 6, 11 9 (AMNH); Middlesex Co.: Faneuil, August 2, 1904, A. P. Morse, 1 6; August 12, 1904, A. P. Morse, 1 9; October 1, 1904, A. P. Morse, 1 Q (AMNH). - Michigan: Cheboygan Co.: July 25, 1940, L. Spencer, 1 Q (OSU); July 1, 1950 J. D. Lattin, I nymph (OSU); Duncan Bay, July 27, 1957, W. J. Han- gan, 2 9 (KU); Emmet Co.: August 10, 1950, J. D. Lattin, 1 2 (OSU); Huron Co.: Sand Point, June 24, 1922, R. 9. Hussey, 1 9 (FSCA). - Minnesota: Saint Louis Co.: Ea- glesnest, July 15, 1959, W. V. Balduf, 2 &, 1 9 (UM); Traverse Co.: Lake Traverse, 7.5 mi SW Wheaton, July 27, 1974, B. Tollefson, 1 4, 1 9 (UCB). - New York: Catta- raugus Co.: South Dayton, 23 July, 1946, R. H. Beamer, 1 Q (KU); Erie Co.: Buffalo, July 16, 1901, 1 9 (CAS); Nassau Co.: Long Island, Cold Spring Harbor Biol. Lab., B. Darnall, 1 ® (CAS); Piermont, June 17, 1934, Scholt, 1 9 (AMNH). - North Dakota: Ramsey Co.: Devil's Lake, July 22, 1920, T. H. Hubbell, 1 @ (FSCA). - Ohio: Erie Co.: Cedar Point, Sandusky, August 31, 1905, Van Duzee, 1 2 (UCB); Union Co.??, Camp Perry, September 5, 1921, W. L. McAtee, 1 Q (USNM). - Oregon: Coos Co.: 15 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 South Slough Sanctuary, SW Coos Bay, September 24. 1988, A. Asquith, 1 9 (OSU); Tillamook Co.: Island Camp, near woods, July 18, 1959, K. Fender, 1 6 (OSU), 5 mi E Pacific City, meadow, September 9, 1962, J. Ca- pizzi, 2 Q (OSU); Sand Beach St. Pk., Sand Lake, Sep- tember 7, 1988, A. Asquith & J. D. Lattin, 4 4, 12 9 (OSU). - Wisconsin: Dane Co: TON, RTE, S28, stream edge, aquatic net, September 14, 1973, J. Hender, 1 Q (UWiM); Wood Co.?, Nevin Marsh, July 23, 1974, site 6, sweep net, D. Bach, 1 @ (UWiM); July 11, 1974, site 2, D. Bach, 1 6 (UWiM). 16 A. BEDOS & L. DEHARVENG Université Paul Sabatier, Toulouse, France NEW SPECIES OF SUPERODONTELLA STACH (COLLEMBOLA: ODONTELLIDAE) FROM THAILAND Bedos, A. & L. Deharveng, 1990. New species of Superodontella Stach (Collembola: Odon- tellidae) from Thailand. - Tijdschrift voor Entomologie 133: 17-26, figs. 1-28, tabs. 1-3 [ISSN 0040-7496]. Published 31 July 1990. The genus Superodontella Stach (Odontellidae) is recorded from Thailand for the first time. Four new species are described, S. cicoria sp. n., S. gouzet sp. n., S. longispina sp. n. and S. flammata sp. n. Some new morphological characters are presented. A. Bedos and L. Deharveng, UA333 du CNRS, Laboratoire de Zoologie, Ecologie des Invertébrés Terrestres, Université Paul Sabatier, 118 route de Narbonne, 31062 Toulouse Cedex, France. Key words. - Collembola, new species, Thailand. INTRODUCTION Superodontella was created by Stach in 1949 for the species Odontella ewingi Folsom, 1916. Long considered as a synonym of Odontella in the liter- ature, it was redefined as a subgenus of Odontella by Deharveng (1981a). It differs from Odontella s. str. in the absence of an apical exsertile bulb on antennal segment IV. This character has proved to be consistent with biogeographical data, as no spe- cies of Odontella s. str. has been found outside the austral region in spite of extensive recent collecting in Madagascar, tropical America and South East Asia. Here we accept Superodontella as valid at generic level with the diagnosis given by Dehar- veng (1981a), because of the large number of spe- cies now known to have this antennal character. TAXONOMIC CHARACTERS USED IN THE DESCRIPTIONS The most important diagnostic characters for Superodontella occur in the antennal and buccal areas. Several new characters are introduced here. Body length. — Adults are scarce, sometimes absent in many Superodontella species. ‘Large juve- niles’ are specimens devoid of any sexual differen- tiation, but of a size as large as or larger than adults. The biological problem underlying these observa- tions is under study. Integument granulations. — (1) Shape, size and arrangement of secondary granules, particularly on head and abd. VI (abdominal segment VI). (2) Ar- rangement and morphology of muscular insertions on head (which could be studied on other areas of the body). Antennae. — (1) Apical structures of ant. IV (antennal segment IV). (2) Number and morphol- ogy of s-setae on ant. IV. (3) Number of trumpet- setae on ant. IV (mostly ventral; low intraspecific variability in adult, but character instar-dependent). (4) Form and length of s-setae on ant. III. (5) Number of ordinary setae on ant. III. Ocular area. — (1) Morphology of PAO (postan- tennal organ) (‘amiboid’ with coalescence of lobes at base, or ‘hypogastruroid’ with well separated lobes). (2) Integument swellings between eyes. Buccal area. — (1) Length and morphology of buccal cone (range from very short and thick to very long and thin). (2) Labrum chaetotaxy, difficult to observe, is usually not mentioned in descriptions. A conspicuous pair of longitudinal sclerotized struc- tures terminating distally could be special modified setae; they are present in all studied species (fig. 12). (3) Labium chaetotaxy, following Massoud (1967) nomenclature (fig. 12). (4) Number and length of perilabial setae (following the nomencla- ture of Yosii (1971)) and postlabial setae (following in part the nomenclature of Da Gama (1988) (fig. 19). Both nomenclatures are incomplete when all Poduromorpha are considered; a general complete nomenclature is presently under study. (5) Form and length of maxilla stipa and fulcrum, including those of E.P.P. (external posterior processus) and LP.P. (internal posterior processus), highly discrim- inant interspecific characters (fig. 6). (6) Number, form and length of mandibles. Dorsal chaetotaxy. — (1) Morphology of ordi- nary setae, particularly on abd. VI. (2) Relative length of s-setae. (3) Position of s-setae on the tergites (stable in the species examined here). (4) Presence/absence of c2 seta on head. (5) Setae of central area (d- and sd-setae) are the same in all our 17 TIJDSCHRIFT VOOR ENTOMOLOGIE. VOLUME 133. 1990 / à \ IN, u a8 4 i È it r i "i Figs. 1-7. Superodontella ciconia sp. n. — 1. Abdomen VI tergite; 2. Central area.of head between dl and d5 setae: integument secondary granules and muscular insertions; 3. Right fourth antennal article, ventral side; 4. Right fourth antennal article and distal part of ant. III, dorsal side; i: i-seta; ms: s-microchaeta; or: distal organite; s1 to s4: s-setae of ant. II] organite; x: ordinary distal seta belonging probably to ant. III; 5. labial, perilabial and postlabial setae; 6. Maxilla and mandible, dorsal view; fulcrum with arm, I.P.P. and E.P.P. (cf text); md: mandible; st: stipa; 7. Dorsal chaetotaxy; setae used in descriptions are named; ms: s-microchaeta of thorax. 18 species and reduced from the basic pattern in Hy- pogastruridae: sdl, sd2 and d4 are absent (fig. 7). (6) Presence/absence of ml and a3/4 setae on th.II- HI (thoracic segments I-II), of ml and m3 on abd. IV and of al on abd. V. (7) Relative position of axial setae. The strong paurochaetosis that occurs in Odon- tellidae results in some uncertainty about the exact nomenclature of the remaining setae. For example, it was difficult to say whether the second seta of the a-row on th. II was a3 or a4; so, it was named a3/4. Appendices and ventral chaetotaxy. — (1) Tro- chanter. Number of setae (stable in the species examined here). (2) Femur. Number of setae. (3) Tibiotarsus. Number of setae (proximal + distal whorls). Important paurochaetosis from the prim- itive pattern 19, 19, 18 of Poduromorpha. (4) Te- nent hair of tibiotarsus. (5) Unguiculus. Always absent in the species studied. (6) Ventral tube. Al- ways 3 + 3 setae in the species described here, the same as found in the first instar of Poduromorpha (primitive state). (7) Ventral setae of abd. I, II and III (8) Dentes and mucro. Few noticeable variation in our species which have always a large mucro compared to dens. Ratio dens/mucro (d/m). Anal spines. — Length and morphology. Anal spines are not derived from setae, but directly from integument secondary granules. Only S. longispina has spine-like anal spines. SYSTEMATIC PART Material is deposited in the collection of Labora- toire de Zoologie, Université Paul Sabatier (Tou- louse). Superodontella ciconia sp. n. (figs. 1-7) Type material. — Holotype young female, Doi In- thanon, 2500 m a.s.l., humus, 2.1.81, sample n° THA 72, Deharveng leg; 49 paratypes from numerous samples, same station, 2000-2500 m, litter, humus and moss, 2 & 9.1.81 and 1.VIII.85, Deharveng and Gouze leg. Description. — Colour pale blue. Length up to 1.5 mm. Integument granulations (figs. 1 & 2). — Roun- dish to polygonal secondary granules all over the body, thickened on abd. VI which is globular. On head, muscular insertions convex and 5-6 secondary granules between d3 and d5 setae. Antennae (figs. 3 & 4). — Ant. IV chaetotaxy as follows: a large apical swelling of the integument; dorso-distally 1 small ovoid organite (or) and 1 very small s-microchaeta (ms); 9 rather long subcylin- drical s-setae (the lateral ones slightly longer and strongly bent), of which 7 are dorsal and 2 ventro- lateral; dorsally and ventrally 16 long slender mou- BEDOS & DEHARVENG: Superodontella setae (“soies mousses”, Deharveng 1981b), 10-15 short pointed setae (including the dorsal i-seta); ventrally 11 rather long trumpet-setae. Ant. Ill organite composed of 2 long bent s-setae (sl et s4), 2 swollen rods (s2 and s3) and an ex- tremely minute, triangular, s5 microchaeta. Ant. III, ant. II, ant. I with 15-16, 10, 6-7 ordinary setae respectively. Ocular area. — 5 + 5 subequal eyes; a few swel- lings in the center of the ocular field; postantennal organ amiboid with 4 lobes, the posterior one very reduced. Buccal area (figs. 5 & 6). — Buccal cone subcylin- drical, slender and elongate. Labium with only 3 setae: F much longer than G, E very small, f absent, without distinct apical small spines. Perilabial area with 4 + 4 setae: b3 and b4 very long, bl and b2 shorter. Postlabial m1 seta more than twice as long as bl; pl absent. Maxilla stipa bowed; I.P.P. and E.P.P. apparently articulated with fulcrum arm; LP.P. slightly longer than E.P.P.; a single mandible present on the right side, as long as the stipa, with arm bent at apex. Dorsal chaetotaxy (fig. 7). — Ordinary setae sub- equal, smooth and pointed, not long except on abd. VI; s-setae quite similar in length and morphology to ordinary setae, slightly thinner (position from th. II to abd. V: 3, 3/4, 4, 4, 4, 3). On head, c2 present. Th. II, th. IT and abd. IV with ml present, abd. V with al present. Appendices and ventral chaetotaxy. — As fol- lows: fr E 18 EI 5 10 O0 LI 5 10 6+ 10 LHI 4 9 yar Lo) abd. I MES abd. II Ve» abd. III Veo) Distal whorl of tibiotarsus with one acuminate tenent hair. Unguiculus absent. Dentes with 5 + 5 dorsal setae, the 3 internal ones stronger; mucro typical of the genus, as long as the dentes. Anal spines (fig. 1). — Rather short, with strong granular papillae. Derivatio nominis. — The species is named for its long and thin buccal cone. Discussion. — Only one Superodontella species with very long and thin mouth parts has been des- cribed, S. cornuta Yosii, 1965 from Japan. It differs from S. ciconta by the presence of spiniform setae on ant. I-II and coxae, apically swollen tenent hairs on tibiotarsi and clavate setae on abd. VI tergite. The axial chaetotaxy of tergites is the same in the two species but the complete dorsal chaetotaxy of S. cornuta has not been described. 19) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 n ssi O n D n di ada te 20 Superodontella gouzei sp. n. (figs. 8 to 14) Type material. — Holotype male, Doi Inthanon, 2500 m a.s.l., moss on rock, 1.VIII.85, sample n° CL 47, Dehar- veng leg; 30 paratypes from numerous samples, same station, 2000-2500 m, litter, humus and moss, 2 & 9.1.81 and 1.VIII.85, Deharveng and Gouze leg. Description. — Colour pale blue. Body length 0.7 to 1 mm (0.75 mm for the holotype male, 1 mm for a paratype female, 0.7 to 1 mm for large juveniles). Integument granulations (figs. 8 & 9). — Roun- dish secondary granules all over the body; they are slightly fused around setal sockets on head and on abd. VI tergite. On head, granules ordered in two longitudinal rows (axial area) and in oblique rows (frontal area); 8 secondary granules between d3 and d5; muscular insertions as fig. 8. Antennae (figs. 10 & 11). — Ant. IV chaetotaxy as follows: no swelling of the integument at the apex; dorso-distally, 1 small ovoid organite (or) and 1 very small s-microchaeta (ms); dorsally and ven- trally about 20 long mow-setae (including non dif- ferentiated s-setae), 12-13 rather short pointed or- dinary setae (including the dorsal i-seta); ventrally 9-10 fairly long trumpet-setae. Ant. III organite composed of 5 ovoid short s- setae hidden under an integument fold; sl and s4 longer than the others. Ant. III, ant. II, ant. I with only 12-13, 10, 7 ordinary setae. Ocular area. — 5 + 5 subequal eyes with a few swellings in the center of ocular field; postantennal organ amiboid with 4 lobes, the posterior one very reduced. Buccal area (figs. 12 & 13). — Buccal cone short and wide. Labrum with 1 + 1 setae and a pair of longitudinal sclerotized structures. Labium with 6 setae; G and F long, f short, E and ?A very short, hardly distinct at higher magnification, ?d rather short; 4 small spines on the reduced distal part. Perilabial area with 4 + 4 setae: b2 very long, bl and b3 subequal, b4 much smaller than the others; post- labial ml seta slightly longer than bl; pl absent. Maxillary stipa straight, oblique and bent at the tip towards fulcrum; I.P.P. and E.P.P. apparently artic- ulated with fulcrum arm; LP.P. twice longer than Figs. 8-14. Superodontella gouzei sp. n. — 8. Central area of head between dl and d5 setae: integument secondary granules and muscular insertions; 9. Abdomen VI tergite; 10. Right third and fourth antennal articles, dorsal side; i: i-seta; s5: external s-seta of ant. III organite, x: ordinary distal seta belonging probably to ant. III; 11. Right third and fourth antennal articles, ventral side; 12. Labral, labial and perilabial setae, lateral view; lr: labral seta; sc: distal sclerotized structure of labrum; sp: small distal spines; 13. Maxilla and mandible, dorsal view; 14. Dorsal chaetotaxy; starts: setae absent compared with pattern in S. ciconia. BEDOS & DEHARVENG: Superodontella Table 1. Differences between Superodontella gouzei and S. salmoni. S. gouzei S. salmoni s2 and s3 setae oval, short T-shaped on ant. III 2 anterior ocelli subequal to larger than others others labial setae ? A (4 sn D N < ia BAE SN N EEE ND = EE 2 ‘ Te A 2 24 ( N N TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 BEDOS & DEHARVENG: Superodontella Table 2. Differences between Superodontella longispina, S. distincta, S. nepalica and S. macronychia. S. longispina S. distincta S. nepalica S. macronychia sp. n. Yosii, 1954 Yosii, 1971 Prabhoo, 1976 (Thailand) (Japan) (Nepal) (India) apical swelling absent present present ? on ant. IV rows of setae 3 22 2 on abd. IV internal teeth basal about 1/3 basal “median” of unguis from basis tibiotarsus long, long, acuminate, not same as tenent hair acuminate capitate differentiated nepalica anal spines long, long, “smaller than long, large, pointed pointed in O. distincta”, subcylindrical pointed and truncated Description. — Colour grey blue, white muscular insertion, conspicuous in alcohol. Body length up to 1.5 mm. Integument granulations (figs. 15 & 16). — Roundish or hexagonal secondary granules all over the body, of small size (12 secondary granules be- tween d3 and d5 setae on head), more convex and thickened on abd. VI; granules joint around setae sockets. Muscular insertions conspicuous, especially a median longitudinal one on head as fig. 15. Antennae (figs. 17 & 18). — Ant. IV chaetotaxy as follows: no apical swelling; dorso-distally, 1 small ovoid organite (or) and 1 very small s-micro- chaeta (ms); 7-8 rather long subcylindrical s-setae (1 thinner), dorsal and ventro-lateral; dorsally and ventrally 16 long slender 7204-setae and 15-16 short pointed setae (including the dorsal i-seta); 31 short ventral trumpet-setae and 2 long ones at the apex. Ant. III organite composed of 4 T-shaped s-setae (sl to s4) under an integument fold and an ex- tremely minute, triangular, s5 microchaeta. Ant. IH, ant. II, ant. I with 18, 10, 7 ordinary setae. Ocular area. — 5 + 5 subequal eyes; about 30 secondary granules in the center of the ocular field; postantennal organ hypogastruroid with 4 large, well separated lobes. Figs. 15-21. Superodontella longispina sp. n. — 15. Cen- tral area of head between d2 and d5 setae: integument secondary granules and muscular insertions; 16. Abdomen VI tergite; 17. Right third and fourth antennal articles, dorsal side; stippled setae are on ventral side of ant. III, except sl-s4 of organite which are dorsally covered with an integument fold; i: i-seta; s5: external s-seta of ant. III organite; x: ordinary distal seta belonging probably to ant. UI; 18. Right fourth antennal article, ventral side; 19. Labial, perilabial and postlabial setae; 20. Maxilla and mandibles, dorso-lateral view; 21. Dorsal chaetotaxy; stars: setae absent compared with pattern in S. ciconza. Buccal area (figs. 19 & 20). — Buccal cone strongly protruded. Labium devoid of distal small spines, with 4 setae: F long, the others short. Peri- labial area with 5 + 5 setae: b3 long, bl, b2, b4 and a fifth seta (b5) short; postlabial ml seta more than 5 times longer than bl; pl present or absent, short. Maxilla fulcrum very strong; I.P.P. and E.P.P. appar- ently partially articulated with fulcrum arm; E.P.P. reduced and more or less fused to fulcrum arm; stipa rather thin with apex bent towards the fulcrum; 2 small mandibles present with rounded apices, the right one slightly longer. Dorsal chaetotaxy (fig. 21). — Ordinary setae subequal, smooth and pointed, not long, except on abd. VI where setae are longer, stronger and acum- inate (sometimes slightly capitated); ratio s- seta/ordinary seta decreasing from 3 to nearly 2 from thorax to abd. V (position from th. II to abd. V: 3, 3/4, 4, 4, 4, 3). On head, c2 present. Th. II, th. III and abd. IV with ml present, abd. V with al present; on th. II and th. IH, ml not aligned with al and pl; on abd. IV, al shifted from the axis and m3 absent. Appendices and ventral chaetotaxy. — As fol- lows: ir: E ali LI 5 12 Tats UO LU 5 Len mar 0) LIII 4 10 6+ 10 abd. I IVES abd. II Mens abd. III Ve:4 Setae of coxae nearly spiniform. Distal whorl of tibiotarsus with one tenent hair very long and acuminate. Unguiculus absent. Unguis with 1 basal inner tooth and 1 + 1 strong laterobasal teeth. Dentes with 5 + 5 dorsal setae, the 3 internal ones much stronger; mucro typical of the genus, shorter than the dentes (d/m = 1.6). 23 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 „ N \ L si tel Figs. 22-28. Superodontella flammata sp. n. — 22. Abdomen VI tergite; 23. Central area of head between dl and d5 setae: integument secondary granules and muscular insertions; 24. Right third and fourth antennal articles, ventral side; circles: setae present in some specimens; 25. Right third and fourth antennal articles, dorsal side; i: i-seta; s5: external s-seta of ant. III organite (ventral); x: ordinary distal seta belonging probably to ant. III; circles: setae present in some specimens; 26. Labial, perilabial and postlabial setae; 27. Maxilla and mandible, dorsal view; 28. Dorsal chaetotaxy; stars: setae absent compared with pattern in S. ciconia. 24 BEDOS & DEHARVENG: Superodontella Table 3. Differences between Superodontella flammata, S. gladiolifer and S. biwonensis. S. flammata apical digitation of ant. IV present thickened s-setae on ant. IV 0 c2 setae on head present basal labial setae 3 rather long, S. gladiolifer S. biwonensis present absent 0 9 absent present 3 short (a fourth could ? have been overlooked) absent present #3 #1,2 well developed absent l very long ml on th. II-III, abd. IV present s setae/ordinary setae on th. II-III #1,2 anal spines minute Anal spines. — Strong, with very long granular papillae (fig. 16). Derivatio nominis. — The species is named for its long anal spines. Discussion. — Three species of Superodontella, all from Asia, have strong anal spines, large and rather long buccal cone and T-shaped sensory rods on ant. III like S. /ongispina. On the basis of the original descriptions, we can give the differential characters as mentioned in table 2. A redescription of the species S. distincta, S. nepalica and S. macro- nychia would be however necessary to precise their real taxonomic status: we lack data about antennal and dorsal chaetotaxy as well as morphology of maxilla and mandible in these species. Superodontella flammata sp. n. (figs. 22-28) Type material. — Holotype juvenile, Doi Inthanon, 2500 m a.s.l., litter and humus near a bog, 2.1.81, sample n° THA 66, Deharveng leg; 4 paratypes in 4 samples, same station, 2500 m, litter and humus, 2 & 9.1.81, Dehar- veng and Gouze leg. Description. — Colour pale blue. Body length 0.55 to 0.72 mm (no adult observed). Antennae short and conical. Integument granulations (figs. 22 & 23). — Rather flat, roundish secondary granules all over the body, enlarged on abd. VI. On head, 2 median and 2 small lateral muscular insertions and 4 secon- dary granules between d3 and d5 setae. Antenna (figs. 24 & 25). — Ant. IV chaetotaxy as follows: no apical swelling but a long distal flame-shaped digitation; dorso-distally, 1 small ovoid organite (or) and 1 small s-microchaeta (ms); dorsally and ventrally 5 long slender mow-setae, 12 short pointed setae (including the dorsal i-seta); 9- 10 short ventral and 2 long apical trumpet-setae. Ant. III organite composed of 5 short and bent s- setae Ant Ii ant: IL, ant I with 4-16" 1026-7 ordinary setae. Ocular area — 5 + 5 subequal eyes; postantennal organ amiboid with 4 lobes, the posterior one very reduced. Buccal area (figs. 26 & 27). — Buccal cone rather long. Labium with 4 setae: F very long, others me- dium with G longer than E and f. Perilabial area with 4 + 4 subequal setae; postlabial ml seta slightly longer than bl; pl absent. Maxilla fulcrum long and thin; distal part of LP.P. apparently artic- ulated; E.P.P. nearly four times shorter than LPP; LPP, fulcrum arm and stipa of equal length; pres- ence of a single mandible on the right side, longer than the stipa, with enlarged arm apex. Dorsal chaetotaxy (fig. 28). — Ordinary setae subequal, smooth and pointed, not long, longer on abd. VI; s-setae slightly longer and thinner than ordinary setae (position from th. II to abd. V: 3, 3/4, 4, 4, 4, 3). On head, c2 present. Th. II, th. II and abd. IV with ml present, abd. V with al present; on th. II and th. III, a3/4 absent. Appendices and ventral chaetotaxy. — As fol- lows: IGE F T LI 5 Jul Wars) LII > 11 Var 2 LIII 4 10 CRIS abd. I TV:3 abd. II Ve:3 abd. III Mes Distal whorl of tibiotarsus with one acuminate tenent hair, slightly differentiated. Unguiculus ab- sent. Dentes with 5 + 5 dorsal setae, the 3 internal ones stronger; mucro typical of the genus, nearly as long as the dentes (d/m = 1.2). Anal spines. — Reduced to minute integument swellings (fig. 22). Derivatio nominis. — The species is named for its flame-shaped digitation at the antennal apex. 25 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Discussion. — In the presence of a long and thin digitation at the apex of antenna, this new species is near S. gladiolifer Massoud, 1965 from New Gui- nea. In the other characters, it seems to be closely related to S. biwonensis Lee, 1974 from South Ko- rea. Differences are summarized in table 3. REFERENCES Deharveng, L., 1981a. La famille des Odontellidae: phy- logénèse et taxonomie. — Travaux du Laboratoire d'E- cobiologie des Arthropodes Edaphiques, Toulouse 3 (Me ZL, Deharveng, L., 1981b. La chétotaxie dorsale de l'antenne et son intérét phylogénétique chez les Collemboles Neanuridae. - Nouvelle Revue d'Entomologie 11 (1): 3-13. Gama, M. M. da, 1988. Filogenia das espécies de Xenylla à escala mundial (Insecta, Collembola). - Evolucion Biologica 2: 139-147. Massoud, Z., 1967. Monographie des Neanuridae, Collem- boles poduromorphes à pièces buccales modifiées. — Biologie de l'Amérique Australe 3: 7-399. Stach, J., 1949. The apterygotan fauna of Poland in rela- tion to the world fauna of this group of insects: families Neogastruridae and Brachystomellidae. - Acta Mono- graphica Musei Historiae Naturalis, Kraków: 341 pp. Yosii, R., 1971. Collembola of Khumbu Himal. - Khumbu Himal 4 (1): 80-130. Received: 21 March 1989 Revised version accepted: 6 September 1989 JEAN BELLE Velp, The Netherlands PROGOMPHUS NIGELLUS AND PHYLLOCYCLA HAMATA, TWO NEW DRAGONELIES FROM BRAZIL (ODONATA: GOMPHIDAE) Belle, J., 1990. Progomphus nigellus and Phyllocycla hamata, two new dragonflies from Brazil (Odonata: Gomphidae). - Tijdschrift voor Entomologie 133: 27-30, figs. 1-10. [ISSN 0040-7496]. Published 31 July 1990. Descriptions and illustrations are given of two new species of Gomphidae (Odonata) from Brazil, viz. Progomphus nigellus and Phyllocycla hamata. Both male nolorypes were col- lected in Brazil, State of Rondonia, Fazenda Rancho Grande. Correspondence: Dr. J. Belle, Onder de Beumkes 35, 6883 HC Velp, The Netherlands. Key words. — Brazil; Progomphus, Phyllocycla, new species. INTRODUCTION To the courtesy of Dr. Rosser W. Garrison at Azusa, California, I had the pleasure of receiving for description two new gomphid species which he had collected himself during his investigations in the area southwest of Ariquemes in the State of Ron- donia, Brazil. The material in question consists of three males; the corresponding females still remain to be discovered. The new taxa are here published under the names Progomphus nigellus and Phyllocycla hamata. The holotypes are deposited in the National Museum of Natural History, Smith- sonian Institution, Washington, D.C; the paratype is in the collection of Dr. Garrison. Progomphus nigellus spec. nov. (figs. 1-6) Material. — Brazil: State of Rondonia, Fazenda Rancho Grande, 62 km SW of Ariquemes (10° 50’ S, 63° 7’ W, 187 m), 2-11 November 1989, 1 @ (holotype), R. W. Garrison. This species is the fourth member of the guyanensis group to which also belong Progomphus guyanensis Belle, 1966, Progomphus approximatus Belle, 1966 and Progomphus boli- viensis Belle, 1973. The males of this infrageneric group differ from those of the closely allied complicatus-group in having the superior anal ap- pendages armed with a sharply-pointed basal ex- terno-lateral dilatation. Progomphus nigellus is perhaps as nearly related to Progomphus boliviensis as to the other two members of the group. The male superior anal appendages are similar to those of boliviensis, the inferior anal appendage approaches more that of approximatus while the anal tubercles resemble those of guyanensis. The morphological differences and similarities in the terminalia of the abdomen will be seen by comparing the figures of the present species with the corresponding ones of the other three members earlier published by me (Belle 1966, 1975); In order not to tamper with the unique specimen I have studied the accessory genitalia 27 situ. This can easily be done because the posterior hamules are in a somewhat extruded position. The accessory genitalia then exhibit a great resemblance to those of boliviensis. Male (holotype). — Total length 38 mm; abdo- men (incl. app.) 29 mm; hind wing 22.5 mm; costal edge of pterostigma of fore wing 2.7 mm. Smaller and more delicate than the other mem- bers of the group. Coloration done in black, brown and greenish white (= pale) but the pale markings on the abdomen are yellow. Face predominantly pale, slightly tinged brown at posterior parts of labrum and postclypeus. Superior surface of frons and occipital plate pale. Vertex brown. Rear of head brown with a pair of small submedian pale spots on top and a pair of larger pale markings on temporae. Crest of occiput slightly and evenly concave, thinly fringed with pale hairs which are about as long as half the middorsal length of the occipital plate. Prothorax with pale middle lobe and black hind lobe. Thoracic colour pattern black with pale mark- ings, resembling that of boliviensts but with metep- imeral pale stripe partly developed and interrupted in middle (fig. 1). Legs dark brown but inner side of first femora pale and second and third femora becoming lighter brown toward the bases. 27 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 1-10. — 1-6, Progomphus nigellus spec. nov., male holotype: 1, diagram of thoracic colour pattern; 2, right posterior genital hamule, ventral; 3, seventh abdominal segment, left profile; 4, tenth abdominal segment and anal appendages, dorsal; 5, the same, ventral; 6, the same, left profile. — 7-10. Phyllocycla hamata spec. nov., male holotype: 7, diagram of thoracic colour pattern; 8, tenth abdominal segment and anal appendages, dorsal; 9, right posterior genital hamule, ventral; 10, apical segments of abdomen and anal appendages, left profile. Wings with a brown tinge, blackish brown vena- tion and brown pterostigma. Brace vein present. Ante- en postnodal cross-veins of first series 10:14- 16:12/11-11-11:10 in fore and hind wings, respec- tively. Second primary antenodal cross-vein the fifth. Basal subcostal cross-vein present. All supra- triangles one-celled. All subtriangles and triangles two-celled with two rows of cells following but with an extra initial cell at hind angle of triangle in hind wings. Intermedian cross-veins 7-7/5-4 in fore and hind wings, respectively. Anal field of fore wing 28 two cells wide for a distance of two cells. Hind wings with five paranal cells, three (left) and four (right) postanal cells, three rows of cells behind Cu2, and a three-celled anal triangle. Abdomen predominantly dark brown or blackish brown. Terminalia of abdomen black, including upper surface of superior anal appendages (con- trary to the other members of the group which have the apical part of the upper surface pale or partly pale). Sides of segments 1 and 2 largely yellow. Sides of segments 3 to 7 with a small, yellow basal spot. Middorsum of segment 2 with a round yellow spot that tapers to hind border of segment. Segments 3 to 7 with a middorsal yellow line over whole length of segment, the line being very fine on segments 4 to 7. There is no tubercle of any sort on venter of segment 1. Segment 7 slightly widening on apical half of segment (fig. 3). Posterior genitalhamule and anal appendages shaped as shown in the accompan- ying figures. Tip of branches of inferior anal appen- dage ending with three teeth. In his letter of 7 April 1990, Dr. Garrison in- formed me, that he had collected this male on a trail at about 4:00 p.m., just before it rained, in company with a male of Aphylla dentata Selys. In addition to these species and Phyllocycla hamata spec. nov. described below, he had also taken Zonophora ca- lippus klugi Schmidt and Phyllogomphoides ce- pheus Belle in the same environment. Phyllocycla hamata spec. nov. (figs. 7-10) Material. — Brazil: State of Rondonia, Fazenda Rancho Grande, 62 km SW of Ariquemes (10° 50 ’ S, 63° 7’ W, 187 m), 2-11 November 1989, 2 ¢ (holotype and para- type), R. W. Garrison. This species is the seventh member of the volsella group (cf. Belle 1988). Dr. Garrison recog- nized it as an undescribed species and wrote from Azusa on 19 February 1990 to me that the males are nearest to my armata but whose morphology of the appendages is different. The species is peculiar by the huge hook which arises erect on the upper surface of each male superior anal appendage. Male (holotype; abdomen broken between seg- ments 4 and 5). — Total length 43 mm; abdomen (incl. app.) 33 mm; hind wing 25.5 mm; costal edge of pterostigma of fore wing 2.9 mm. Head brown with pale (= leaden grey) markings. Labrum lighter brown along free border and with a symmetric pair of weakly developed pale spots. External surface of genae largely pale. Anteclypeus pale. Postclypeus with a pale spot on each lateral side. Superior surface of frons pale, darker on the middle line. Rear of head lighter on temporae. Crest of occiput slightly concave, fringed with brown hairs which are about as long as the middorsal length of occipital plate. Prothorax black above. Prerothorax black to dark brown with pale (grey to grey-green) stripes; its colour pattern shaped as shown in diagram (fig. 7). Femora brown, but brownish yellow on inner sides of first and second femora and on posterior side of third femora basally. Distal spines of ante- rior outer row of third femora more or less spaced and one-fifth to one-fourth as long as local diameter of femur. Tibiae, tarsi and claws black. BELLE: New dragonflies from Brazil Wings with a brown tinge, black venation and light brown pterostigma. Brace vein present. Ante- and postnodal cross-veins of first series 12:17- 15:13/11:12-12:11 in fore and hind wings, respec- tively. Second primary antenodal cross-vein the fifth. Basal subcostal cross-vein present in all wings. Supratriangle in right hind wing three-celled, in other wings two-celled. Subtriangles in fore wings two-celled, in hind wings one-celled. Triangles two- celled with two rows of cells following but with an extra initial cell at hind angle of triangle in hind wings. Intermedian cross-veins 9-8/6-6 in fore and hind wings, respectively. Anal field of fore wings two cells wide for a distance of two cells in anterior row. Hind wings with four paranal cells and three postanal cells (the fourth paranal cell is the first postanal cell), a one-celled anal loop, three rows of cells behind Cu2, and a four-celled anal triangle. Abdomen dark brown with paler markings as follows: Sides of segments 1 and 2 largely yellow. Sides of segments 3 to 7 with yellow baso-lateral spots, the basal spots of both sides of segment 7 connected on the middorsum at extreme base. There is a yellow middorsal line on segments 2 to 7. Sides of segments 8, 9 and 10 largely brown- yellow. Lateral dilatations of segments 8 and 9 nar- row and black. Lateral margins of segment 8 with four (left) and three (right) denticles at apex, those of segment 9 denticulated at extreme base. Poste- rior margin of segment 10 denticulated at level of bases of superior anal appendages. Dorso-apical rim of segment 10 about one-fourth the middorsal length of segment. Superior anal appendages black and shaped as shown in figs. 8 and 10. Rear margin of vesicle deeply cleft, bottom of cleft rounded and without a median elevation of any sort. Tip of posterior genital hamule rather long (fig. 9). The male paratype lacks the tip of the left super- ior anal appendage but for the rest the specimen is in a perfect condition. The pale markings of the pterothorax are yellowish green and the dark lateral stripes brown. The wings are slightly brown- tinged. The cubito-anal interspace of the right hind wing has two cross-veins instead of the usual single cross-vein and the pterostigma is a trifle larger than that of the male holotype. The measurements of the male paratype are: Total length 42.5 mm; abdomen (incl. app.) 32.5 mm; hind wing 24.5 mm; costal edge of pterostigma of fore wing 3 mm. Remark. — I take this opportunity of correcting a misprint in the description of Phyllocycla armata Belle, 1977. On page 7, in rule 7 from above, part of a line has been left out. We have to read: “Trig- onal interspace in fore wings starting with two rows of cells from triangle outwards, that in hind wings starting with a row of three cells against triangle followed by two rows of cells.” 29 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 REFERENCES Belle, J., 1966. Surinam dragon-flies of the genus Progomphus. - Studies on the fauna of Surinam and other Guyanas 8: 1-28; pls. 1-4. Belle, J., 1973. A revision of the New World genus Progomphus Selys, 1854. - Odonatologica 2: 191-308. Belle, J., 1977. A new species of Phyllocycla Calvert, 1948 from Brazil (Odonata: Gomphidae). - Entomologische Berichten, Amsterdam 37: 6-7. 30 Belle, J., 1988. A synopsis of the species of Phyllocycla Calvert, with descriptions of four new taxa and a key to the genera of Neotropical Gomphidae (Odonata, Gomphidae). - Tijdschrift voor Entomologie 131: 73- 102. Received: 17 May 1990 Accepted: 17 May 1990 K. A. MONK & R. K. BUTLIN* Department of Pure and Applied Zoology, University of Reading. * School of Pure and Applied Biology, University of Wales. A BIOGEOGRAPHIC ACCOUNT OF THE GRASSHOPPERS (ORTHOPTERA: ACRIDOIDEA) OF SULAWESI, INDONESIA Monk, K. A. & R. K. Butlin, 1990. A biogeographic account of the grasshoppers (Orthoptera: Acridoidea) of Sulawesi, Indonesia. — Tijdschrift voor Entomologie 133: 31-38, tabs. 1-4. [ISSN 0040-7496]. Published 31 July 1990. Nineteen acridid species were collected from the Dumoga Bone National Park and its environs in North Sulawesi, Indonesia, during 1985: two were new to science and one was known previously only from the Philippines. The total acridoid fauna of Sulawesi, including these three species, comprises 61 species of Acrididae (53 Catantopinae, four Oedipodinae, three Acridinae, one Gomphocerinae) and one species of Pyrgomorphidae. This fauna has four main characteristics: (a) a high level of specific and generic-level endemism, (b) low generic-level diversity but a high number of species per genus, (c) diverse distributions outside Sulawesi, and (d) localised distributions within the island. In the Catantopinae, 83% of species are endemic to Sulawesi, reflecting the importance of the island as an area of endemism. Species distributions within Sulawesi suggest that each region of the island (North, Central, South and South-east) may also be an area of endemism in its own right. These observations are discussed in relation to the geological history of Sulawesi. They suggest that for much of its history the island has been more isolated from other land masses than it is at present, and that it has been fragmented either into separate islands or by ecological barriers. Correspondence: Dr. R. K. Butlin, School of Pure and Applied Biology, University of Wales, P.O. Box 915, Cardiff CF1 3TL, Wales. Key words. - Biogeography, Sulawesi, Wallacea, grasshoppers, endemism. INTRODUCTION Although the Acridoidea in temperate zones and in dry equatorial areas are well-known, only wide ranging, economically-important pests, e.g. Oxya japonica japonica Thunberg and Locusta migratoria Linnaeus, have been studied in wet equatorial areas. The taxonomic work in South-east Asia by Ramme (1941) and Willemse (1951, 1956, 1957) is an ex- ception but it has not been extended. In particular, the acridoid fauna of Sulawesi has been largely ig- nored despite the great interest in other animal groups inhabiting the island. Sulawesi lies in a deep sea region, also including the Lesser Sunda Islands (Lombok to Timor) and the Moluccas, with lies between two continental margins: the Sunda (Malaysia and the Greater Sunda Islands), and the Sahul shelves (New Guinea and offshore islands). Classically, this area was seen as the meeting point of two major biotas: the Orien- tal and the Australasian. Wallace (1869) originally separated them by dividing Bali (and Borneo) from Lombok (and Sulawesi). Although Wallace’s Line may apply to some groups such as freshwater fish and mammals, the diversity of biogeographical patterns across this region for different taxonomic groups is high (see George 1987). Even with this overlap of broad transitional zones, Wallacea (a collective term for the islands of the deep sea re- gion) is generally considered to be insular and bi- otically impoverished, but with unique characteris- tics indicating long periods of isolation (Dickerson et al. 1928). Sulawesi covers 159,000 km? divided into four regions, Utara, Tengah, Selatan, and Tenggara, or North, Central, South, and South-east Sulawesi, re- spectively (see Whitten, Mustafa & Henderson 1987, for geographical details). North Sulawesi ts a narrow peninsula, 772 by 103 km. Volcanoes, 1800- 2400 m above sea level (asl), range along the penin- sula. Seismic activity was recorded during 1985 and 31 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 1986. Limestone outcrops (possibly Cretaceous) oc- cur in the western half of North Sulawesi. The northern peninsula is linked with Central Sulawesi by a narrow neck of land at the western extremity. The Sangihe and Talaud Islands lie between the northern tip of North Sulawesi and the Philippines. Central Sulawesi is also mountainous, with peaks reaching 2000-3000 m asl. A limestone mountain range running north to south through South Su- lawesi is 500-1000 m asl and bisected by a narrow plain. The lakes on the plain are only 1-2 m deep and were once part of the sea, suggesting that at least this peninsula may have been partially sub- merged at times. The single volcano in this region, Lompobatang (2871 m asl), is extinct. The penin- sula of South-east Sulawesi has a western mountain range reaching 2800 m asl and a lower-lying eastern area. The vegetation is affected by regional climatic differences. Central Sulawesi has the greatest ratio of wet to dry months, whereas more seasonal rain- fall occurs in the peninsulas (Whitmore 1984). The central region is covered in evergreen rain forest, whereas semideciduous forest is more common in the peninsulas (Walker 1982). Open habitats are mainly secondary, due to forest clearance for agri- culture (Whitten et al. 1987). Geologically Wallacea is a highly complex region originating in an interaction between the South- east Asian, Australian, and Pacific plates (Charlton 1986). The present island of Sulawesi is believed to be derived from two fragments that originated se- parately during the breakup of east Gondwanaland about 320 My ago, and remained separate during a northerly move starting 220 My ago (Audley- Charles 1987). From the late Cretaceous onwards, these fragments probably formed part of an ar- chipelago between the Asian mainland and Austra- lia-New Guinea but their positions relative to other land masses are uncertain. Holloway (1987) em- phasized two alternative theories about the rela- tionship of Sulawesi with Borneo: 1. The fragment now forming the western part of the island (North, South and part of Central Sulawesi) has always been in about its present position relative to Borneo, whereas the eastern fragment was Australasian in origin, 2. The two fragments were both isolated island arcs, now fused and thrusted towards Borneo. The present island was probably formed about 15 My ago. The area of land above sea level, its division into separate islands and its climate are additional factors which must have contributed to the present day fauna of Sulawesi but about which little is known (Morley & Flenley 1987). It is now believed that Sulawesi was never linked by land bridges to any other land mass following its formation 15 My ago (Audley-Charles 1987; Hollo- way 1987). During the Pleistocene, the sea level fell several times when affected by the Glacial expan- 32 sion of the ice caps. The largest drop, 170,000 years ago, linked Sundaland (Sumatra, Java, Bali, and Bor- neo) with the Asian mainland, with land extensions northward to the Philippines. Exposed land also extended around South Sulawesi toward the Lesser Sunda Islands, and from Central Sulawesi eastward toward New Guinea. During these periods of low sea level a drier, more seasonal climate prevailed and dry savannah may have extended through the Philippines to the Lesser Sunda Islands and into Australia, separating two large, wet rainforest areas in Sundaland and New Guinea; some seasonal plants still survive in South Sulawesi (van Steenis 1979). The climatic fluctuations and corresponding vegetational changes of the Quaternary are now known to have occurred in the late Tertiary as well (Morley & Flenley 1987). MATERIALS AND METHODS The analysis reported here is based on a combi- nation of data from the literature and collections made by the authors and others in the Dumoga- Bone National Park, North Sulawesi during 1985. This work formed part of the Royal Entomological Society's ‘Project Wallace’ expedition. Insects were collected by sweeping, beating, and hand-searching. Agricultural crops, grassland, and primary semi- deciduous forest (undergrowth, bushes, saplings, and trees up to 2 m) were examined at all times of day and night and in three separate periods cover- ing most of the year. Specimens collected from the tree canopy during the British Museum (Natural History) fogging programme (N. Stork, unpub- lished) were also examined. The principal published records of the acridid fauna of Sulawesi are: Ramme (1941), Dirsh (1954), Willemse (1951, 1956, 1957, 1968), Hollis (1968, 1971, 1975), Kevan & Chen (1969), and Ritchie (1982). Family and subfamily classification follows that adopted in the collection of the British Museum (Natural History). RESULTS Expedition collection of North Sulawesi Acridoidea Nineteen species from eighteen genera of Acri- doidea were found in the Dumoga Bone National Park and its environs during 1985 (table 1). This includes two undescribed catantopine species, from the genera Tarbaleus and Bibracte, both of which were collected from the canopy by fogging. One species, Eoscyllina luzonica, was previously known only from the Philippines and was collected only from limestone grassland in the western part of the National Park. This suggests that the data available in the literature on the species present in Sulawesi MONK & BUTLIN: Grasshoppers of Sulawesi Table 1. Species of Acridoidea occurring in Sulawesi and their distributions. Wings! Distribution? Outside Within Pyrgomorphidae Atractomorpha Across psittacina psittacina de Haan ah W SYP All Acrididae Acridinae Acrida Across willemsei Dirsh + W SWP All Calliphlaeoba Sulawesi celebensis Ramme ar B — N,C Phlaeobacris Sulawesi reticulata Willemse W — € Gomphocerinae Eoscyllina P luzonica Bolivar I SF W P N È Oedipodinae Heteropternis Across obscurella Blanchard W Across All Atolopus Across thalassinus tamulus Fabricius Ar W Across All Locusta Across migratoria Linnaeus ali W Across All Gastrimargus Across marmoratus (Thunberg) W Across All Catantopinae - Endemic genera Alectorolophus deceptor Ramme ala B _ SE, N > obscoenus Brunner von Wattenwyl U B — N = unilobatus Brunner von Wattenwyl B — S 5 applicatus Brunner von Wattenwyl B — ISTE È speciosus Brunner von Wattenwyl B — S mutator Ramme B — N È sororum Ramme B _ E = lineatus Ramme B — C Bs guttulosus Ramme B = N 5 Acrolophus cornutus Ramme B — C ES Alectorolophellus heinrichi Ramme B — S 5 Mengkokacris olivacea Ramme W? _ SE = Heinrichius nobilis Ramme W — N > Celebesia acuticerca Bolivar C ala W? — N x ferruginata Brunner von Wattenwyl We? — N Ri heinrichi Ramme W? = S à Paramesambria flavomaculata Willemse B — S Paracranae celebesia Willemse Ar B — N © Catantopinae - Non-endemic genera Tarbaleus Brunner von Wattenwyl M, NG sp. n. te B — N = Oxya Across Japonica japonica Thunberg W SP All bolaangensis Hollis W -- N “ stresemanni Ramme W — C sa Gesonula Across mundata pulchra Rehn SF W S N Chitaura M, Java, India? brachyptera Bolivar I B N flavolineata (Willemse) B — N Ea atrata Ramme B — C 4 mirabilis Carl B C 5 ochracea Ramme B _ S Sp vidua Carl B M S mengkoka Ramme B — SE samanga Carl B _ SYSEAE poecila Ramme B — N elegans Ramme B — S + Ue (US) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 1. Species of Acridoidea occurring in Sulawesi and their distributions. (continued). Oxytauchira gracilis Willemse Stenocatantops splendens Thunberg angustifrons Walker Bibracte Stàl sp. n. Valanga transiens Walker Oxyrrbepes meyeri Willemse obtusa de Haan Cranaella carnipes Ramme Tristria pisciforme Serville Austracris guttulosa guttulosa Walker Mesambria maculipes Stàl elegans Ramme trapezina Ramme rectangularis Ramme Traulia kukenthali Ramme sanguinipes Stàl Apalacris gracilis Willemse incompleta Willemse celebensis Willemse Moessonia tenebrifera Walker Notes + indicates a species collected by Project Wallace 1 - W - fully winged W? — winged but probably a poor flier B - brachypterous or wingless 2 - Distributions outside Sulawesi: M - Moluccas S — Sundaland P - Philippines A - Australasia NG - New Guinea Across - at least Sundaland to New Guinea are reasonably reliable since intensive collecting produced so few undescribed species, and those only from inaccessible habitats. Data on distribution within Sulawesi is less reliable with a significant proportion of species described from only one or a few localities. However collecting localities in the past have been widespread and so this may reflect very localised species distributions (see below). A probable exception is Central Sulawesi, parts of which remain very inaccessible. The expedition col- lection represents only a small proportion of the total acridoid fauna of Sulawesi (16 out of 59 described species) and this is also likely to be a result 34 Wings! Distribution? Outside Within Burma W = C * Across ala W Across all W Across S S, M, P + B = N = Across Ww = N,C S W = ? * W S N, C P B — N È S W S N PA W A N India? + B — all B S B = @ = B = N + S,P, M Ww? pus > * We? S ? S,P W = S W = S Ww wee ? * NG We = N 2 - Distributions within Sulawesi: all - all four regions N - North S - South SE - South-East € - Central * indicates that the species is known only from one or a few sites. of localised species distributions rather than under- collecting. The acridoid fauna of Sulawesi Expedition and literature data give a total of 62 Sulawesi acridoid species in 33 genera and five sub- families (table 1). The Catantopinae form by far the largest group with 53 species. This very diverse subfamily, which some authors consider to be an amalgam of several distinct subfamilies (eg Dirsh 1961), is one of the few acridoid groups to have adapted to forest environments. All of the non- catantopine species in Sulawesi are forest edge or open habitat species whereas about 80% of catan- topine species occupy habitats within the forest. Related to this is the occurrence of flightless spe- cies, with wings either absent or reduced, which are much more common in the Catantopinae (37/53 species) than in the other subfamilies (1/9 species). Flightlessness in grasshoppers is probably an adap- tation to poor habitat quality, diversion of resources away from wing and flight muscle production al- lowing earlier reproduction and/or greater fecun- dity (Ritchie et al. 1987), and is common in forest species in other areas (Jago 1973). Flightlessness clearly limits dispersal and so may have an impact on levels of endemism. In Sulawesi endemism is high for the Acridoidea, as it is for many other groups (Whitten et al. 1987) including Fumastacidae and Tetrigidae in the Orthoptera (Butlin et al 1989, Blackith & Blackith 1988). Over- all 74% of acridoid species are endemic to Sulawesi but the proportion is higher in the Catantopinae (83%) than in the other subfamilies (22%). There is a clear link with the ability to fly: only two out of nine non-endemic catantopines are flightless, and the one flightless non-catantopine is endemic (table 1). Table 2. Comparison of the Catantopinae of Sulawesi and Java. Java Sulawesi Number of species recorded 47 DI, Number of genera recorded 34 24 Number of species per genus 1.58 DAI Proportion of endemics — species 40% 83% - genera 15% 33% Table 3. Distributions outside Sulawesi. Non- Non- Relationships endemic endemic of Endemic Species! Genera? Genera’ Across >) 1 (10) 0 Sunda Shelf 7 4 (8) À Borneo 0 0 (0) 1 (exclusively) Philippines | 1(1) 0 (exclusively) Moluccas I 0 (0) 0 (exclusively) Australasia l 2 (3) 1 Notes 1. Includes distributions of Oxya japonaci japonaci, Ge- sonula mundata pulchra, and Austracris guttulosa gut- tulosa. All species have distributions ‘Across’ but the subspecies have informative distributions. 2. Excluding (including) genera with non-endemic spe- cies. 3. See text for details. MONK & BUTLIN: Grasshoppers of Sulawesi The special nature of the Sulawesi fauna can be appreciated better if it is compared with the fauna of Java. Java is the most comparable island in the region in terms of land area (126,500 km?) but is part of the Sunda Shelf and has a clearly Oriental fauna. This comparison can be made most easily for the Catantopinae using data from C. Willemse (1956, 1957), E Willemse (1965) and Hollis (1971, 1975). The total number of species recorded is sim- ilar (table 2) but the number of genera present in Sulawesi is lower with a significantly greater number of species per genus (X? = 6.5 P < 0.01) Endemism at the species level in Sulawesi is twice that in Java, as is generic level endemism. Biogeographical links Sharing of taxa between Sulawesi and its sur- rounding areas is summarised in table 3 for non- endemic species (and subspecies) and genera. It is clear that Sulawesi shares the greatest number of species and genera with the Sunda Shelf as a whole (Peninsula Malaysia, Borneo, Sumatra, Java and Bali). This bias is strongest at the specific level but is also present among non-endemic genera. We have found no exclusive links with Borneo at either level, despite the close proximity of the two islands. Links with the Philippines are mostly due to species or genera which also occur on the Sunda Shelf but there are two exclusive links, the gom- phocerine Eoscyllina luzonica and the catantopine genus Cranaella. No species shared between Su- lawesi and the Lesser Sunda Islands have been iden- tified. To the east there is one species, Chitaura vidua, which occurs in Sulawesi and the Moluccas and one, Austracris guttulosa, which is distributed widely in Australasia and for which Sulawesi is the western limit. At the generic level there are two further links to the east, the genera Moessonia and Tarbaleus. For endemic genera biogeographical information can only be derived from the distribution of sister genera but since the phylogeny of these groups has been studied very little these relationships are not known with any certainty. The following relation- ships, suggested by Ramme (1941) and Willemse (1951, 1956, 1957), are included in table 3. A group of four endemic genera, the Acrolophi (Acrolophus, Alectorolophus, Alectorolophellus and Mengkok- acris), has apparently evolved within Sulawesi, in- dicating a long period of isolation. The nearest relatives of this generic group are apparently Paralectorolophus and Lyrolophus which occur on Lombok and Java respectively. The acridine genera Calliphlaeoba and Phlaeobacris are both probably related to Phlaeoba which has a Sunda Shelf plus Philippines distribution. Heinrichius is believed to be most closely related to Noliba and therefore provides the only exclusive 35 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 link with Borneo. Paracranae is part of the Cranae group of genera which occurs in the Moluccas and New Guinea (Willemse 1977a, b). Two genera are problematic because their near- est relatives are non-endemic Sulawesi genera. This implies that the latter are polyphyletic since the Sulawesi representatives of the two genera of each pair are likely to be more closely related to one another than the Sulawesi species of the non-en- demic genus are to species in the same genus occur- ing elsewhere. Celebesia is believed to be most closely related to Traulia (Sulawesi and the Sunda Shelf), an Paramesambria to Mesambria (Sulawesi and India, Henry 1942). The distribution of Mesambria is particularly surprising but it is paral- leled by Chitaura which also has an Indian represen- tative (Hollis 1975). Neither of the two endemic genera (Celebesia and Paramesambria) has been included in table 3. Endemism within Sulawesi Mosaic distributions of species or subspecies within Sulawesi have been noted for several animal groups, eg macaques, carpenter bees, pond skaters (Whitten et al 1987 and see Knight & Holloway 1990). Ramme (1941) commented on a similar phenomenon in the Catantopinae and this appears quite striking when the island is divided into re- gions (table 4). A high proportion of the species present in any one region are known only from that region. This is particularly clear in the two most species rich genera in Sulawesi: Alectorolophus has nine species, seven of which are known from only one region, and Chitaura has ten species, nine of which are known from only one region. However the distributional information available is limited. Many species (table 1) are known from only one locality, but collecting localities overall have been widely distributed around the island and so this may represent genuinely very restricted distributions. Where more localities are known, five out of seven endemic and six out of eight non-endemic species are restricted to one region. Intensive searching of a restricted area during the Project Wallace expedi- tion yielded only a small proportion of the total number of catantopine species known to occur in Table 4. Regional distribution of Catantopine species within Sulawesi. Region Number of species occurring in: that region only that region and elsewhere North 19 5 Central 8 6 South 12 5 South-East 3 5 36 Sulawesi (10/53) but a much higher proportion of the species known to occur in North Sulawesi (9/21). Only one species previously reported from another region was collected. Nine of the species not collected on the expedition were previously reported only from the western part of the penin- sula of North Sulawesi, (particularly Toli-Toli) dis- tant from the Dumoga-Bone National Park, sug- gesting that distributions are also restricted within regions. The within island endemism in the genus Chitaura is particularly striking in this context. Spe- cies in this genus are brightly coloured and conspic- uous and occur on the forest edges as well as in light gaps within the forest. It is unlikely that the expe- dition would have failed to collect any species occur- ring in the Dumoga Bone area and yet only one of the ten Sulawesi species was found. This argument applies to collecting of Chitaura ın general and so this genus gives strong support to the existence of intra-island endemism, and would be a good candi- date for further work. DISCUSSION The Acrididae of Sulawesi show four main char- acteristics: (a) a high level of endemism, (b) low generic diversity but a high number of species per genus, (c) diverse distributions outside Sulawesi, and (d) localised species distributions within the island. These features have parallels in several other animal groups (Whitten et al 1987, Knight & Holloway 1989) and can be related to the geological history of the island. The Acridoidea did not begin their radiation until the mid-Tertiary (Sharov 1968), that is after the break up of Gondwanaland and the beginning of the proposed northward movements of the fragments of present day Sulawesi. Thus the current Acridoid fauna of Sulawesi must have originated by dispersal and its low generic diversity can be explained in one of two ways: either Sulawesi was more isolated for much of its history than it is at present, or it has only recently become dry land due to tectonic uplift. The former explanation is more consistent with the observation of high levels of specific and generic endemism and high numbers of species per genus on the island, all of which suggest a long period of independent evolution for the Sulawesi acridoids. The sources of colonists for Sulawesi appear to have been numerous although the largest number of shared species and genera is clearly with the Sunda Shelf region. This may be partly due to the higher diversity of Acridoidea in Asia than in Australasia but there are several species or subspecies distribu- tions which extend eastwards only as far as Sulawesi and very few equivalent Australasian distributions which extend westwards to Sulawesi. Thus at least a part of Sulawesi appears to have been closer to the Sunda Shelf than to Australasia for a significant part of its history. However its current proximity to Borneo is probably recent since there are no exclusive biogeographical links at the level of shared species or genera. This is also true in other animal groups, notably the butterflies (Vane- Wright 1990). Exclusive links with the Philippines and Moluccas are also weak and may be recent. This is consistent with a general geological view of the area as a collision zone in which the area of dry land is increasing and land masses are moving closer together. The high proportion of endemic species and genera in Sulawesi, the number of species per ge- nus, and the evidence for evolution of new genera within the island all suggest not only a long period of isolation but also the opportunity for consider- able evolutionary change and speciation. The mos- aic distributions of species, particularly in the forest dwelling, flightless catantopines, may provide a clue to the conditions which have favoured these developments. Mosaic distributions are a feature of several other animal groups (Whitten et al 1987, Knight & Holloway 1990) and the patterns in dif- ferent groups appear to be broadly coincident, de- fining areas of endemism within Sulawesi. The simplest explanation for these areas of endemism is past fragmentation of Sulawesi. This could have been in the form of an archipelago at times of higher sea level or before tectonic uplift and vol- canic activity had created the present land mass (Musser 1987), or due to reduction in the extent of forest habitats in periods of drier climate (Morley & Flenley 1987). The wingless, forest adapted ca- tantopines would be particularly susceptible to pop- ulation fragmentation by such processes. Evolu- tionary divergence on islands or in refugia would have been followed by expansion to form the pres- ent mosaic distribution. Contact zones between the forms have not been studied, or indeed accurately located, and so the consequences of renewed contact are unknown. In general several possible types of interaction exist: sufficiently divergent forms may be able to spread into sympatry progressively obs- curing the mosaic pattern, less divergent forms may interact at sharp parapatric boundaries due to eco- logical exclusion, or where reproductive isolation is incomplete a hybrid zone may form (Barton & He- witt 1985). Identification and analysis of such con- tact zones is likely to be a very productive area for future research. Strict isolation on islands or in refugia may not be necessary for evolutionary divergence (Barton 1989), especially in an island with the curious shape of Sulawesi in which gene flow between popula- tions inhabiting different peninsulas would be ex- tremely restricted even if they were connected by continuous suitable habitat. However,divergence without isolation is likely to produce different pat- terns of variation for individual characters within MONK & BUTLIN: Grasshoppers of Sulawesi species as well as for different species. Coincidences of patterns of variation are most likely to be pro- duced by contraction and expansion of populations, and this process may be necessary for the comple- tion of speciation (Hewitt 1989). The mosaic distributions of species within Su- lawesi constitute evidence either for the archipe- lagic nature of the land mass in the past or for habitat fragmentation during climatic fluctuations, or both. The time scale for these changes, and thus for the evolution of the Sulawesi endemics, is un- certain. They could have occurred mainly in the Pleistocene or over a much longer time scale of the order of the 15Myrs suggested as the time since the formation of Sulawesi as a single unit (Audley- Charles 1987). Most probably the present fauna of Sulawesi is the result of several superimposed pe- riods of divergence. Studies of genetic divergence and detailed investigations of the contact zones appear to be the way forward in understanding the evolution of the Sulawesi grasshoppers. ACKNOWLEDGEMENTS We thank Judith Marshall (BM(NH)) for much help in the identification of specimens, and Drs J. Holloway, R. Vane-Wright, H. Malicky, D. Clayton, and M. Robertson for comments on earlier drafts of the manuscript. We are grateful to Prof. R. Blackith for the use of his unpublished data. KAM thanks the Royal Society, the Godman Exploration Fund, and the Percy Sladen Memorial Fund for financial support. RB thanks the Percy Sladen Memorial Fund, Mr C. R. Butlin and the School of Biological Sciences, University of East Anglia. 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Willemse, F, 1977a. A study on the genus Cranae Stall (Orthoptera, Acridoidea, Catantopinae). — Tijdschrift voor Entomologie 120: 121-152. Willemse, F, 1977b. A study on the genus Cranaella Ramme (Orthoptera, Acridoidea, Catantopinae). — Tijdschrift voor Entomologie 120: 121-152. Received: 7 July 1989 Accepted: 19 September 1989 THOMAS PAPE Zoologisk Museum, Copenhagen TWO NEW SPECIES OF SARCOPHAGA MEIGEN FROM MADEIRA AND MAINLAND PORTUGAL (DIPTERA: SARCOPHAGIDAE) Pape, T., 1990. Two new species of Sarcophaga Meigen from Madeira and mainland Portugal (Diptera: Sarcophagidae). — Tijdschrift voor Entomologie 133: 39-42, figs. 1-9. [ISSN 0040-7496]. Published 31 July 1990. Sarcophaga (Discachaeta) amputata sp.n. (Madeira) and Sarcophaga (Pierretia) iulicida sp.n. (Portugal) are described and their phylogenetic affinities are discussed. S. zwlicida was bred from the millipede Ommatoiulus moreleti (Lucas, 1860). Dr. T. Pape, present address: Danish Bilharziasis Laboratory, Jaegersborg Allé 1D, DK-2920 Charluttenlund, Denmark. Key words. - Diptera; Sarcophagidae; New species; Madeira; Portugal. INTRODUCTION A number of specimens representing two undes- cribed species of Sarcophaga Meigen were reco- vered from various sources. As the accompanying biogeographical and biological information is rather interesting, it is felt that the following des- criptions are justified, although they do not form part of a more thorough revision. Depositories for specimens examined are indi- cated by the following acronyms: BMNH — British Museum (Natural History), Lon- don, England; NMwc - National Museum of Wales, Cardiff, England; ZMUC - Zoologisk Museum, Uni- versity of Copenhagen, Denmark. SYSTEMATIC PART Sarcophaga (Discachaeta) amputata sp.n. (figs. 1-4) Type material. — Holotype @, Madeira (Portugal): Be- tween Canical & Prainha, 5.viii.1981, M. Jones (NMWC). The holotype is in good condition, glued to the pin along the right side of thorax and with the terminalia dissected and glued to a piece of cardboard pinned with the specimen. Paratypes. Madeira: 18 19, [no further locality, but with the number 1948-301], T.W. Wollaston (BMNH, NMWC); 29, data as holotype (NMWC, ZMUC); North end of Deserta Grande, 14 19, 18.viii.1981, M. Jones (ZMUC, NMWC). Description Male. — Head. Narrowest part of frons 0.25-0.30 X head width. Parafacial plate with a uniserial row of setae along the eyemargin, the lower setae of this row being long and bristly. Post- gena with white setae in about posterior 0.50-0.75 (strict lateral view). Thorax. Proanepisternum setose in about upper half. Chaetotaxy: 3 indistinct pairs of presutural acrostichals, postsutural acrostichals absent or re- presented by the prescutellar pair only, dorsocen- trals = 3 + 3, intra-alars = 1 + 2, supra-alars = 1 + 3, post alars = 2. Scutellum with a pair of apicals, 3-4 pairs of marginals and without discals. Mid femur with 2-3 anteroventral bristles, no postero- ventral bristles. Hind tibia with a sparse row of elongated posteroventral setae. Abdomen. Tergites 3-4 with silvery grey micro- tomentum mainly restricted to anterior half and almost non-changing; interrupted by a median black stripe. Tergite 5 with microtomentum re- duced to narrow strips in antero-lateral position. Terminalia black or blackish brown. Protandrial segment with a row of marginal bristles. Cercus in profile with a distinct subapical dorsal hump. Ae- deagus with very long harpes that terminates in a distinct hook, and a long, terminally bifid juxta. Gonopod with the usual row of long setae along dorsal margin but with some additional setae on the median surface. Length: 8-9 mm (estimated from dissected spe- cimens). Female. — Very like the male apart from the usual sexual dimorphism. Abdominal pattern slightly more changing and with the median black stripe almost absent. Mid femoral organ not differ- entiated. Terminalia red. Tergite 6 broadly arched, the dorsal part reduced to a narrow strip, and mar- ginal bristles only present laterally. Length: 7.0-7.5 mm. 39 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Distribution. — Palaearctic: Madeira islands (Madeira, Deserta Grande). Etymology. — A Latin adjective, amputatus = cut off, amputated. The name refers to the structure of the aedeagus where the pair of very short arms at juxtal base will separate the species from all other species of the subgenus Discachaeta. Remarks. — The sarcophagid fauna of Madeira was reviewed by Pape (1986), who listed a total of five species, three of which were judged to be recent introductions by man. The present species is inter- esting as it is the third apparently endemic species, and much evidence is in favour of considering this the sister species of the Madeiran Sarcophaga (Discachaeta) kunonis (Pape, 1986). The latter spe- cies was described in the genus Discachaeta Ender- lein, which here is given subgeneric rank following the wide concept of Sarcophaga employed by Séguy (1941), Downes (1965) and Pape (1988). Disca- chaeta is probably a monophyletic group, this hy- pothesis being corroborated by at least the follow- ing character states that all seem to be derived with CX h Figs. 1-4. Sarcophaga amputata, male terminalia. 1, right cercus and surstylus, lateral view, setae omitted; 2, cerci, pos- terior view, setae omitted; 3, right par- amere + gonopod, lateral view; 4, ae- deagus, lateral view. Abbreviations: h = harpes, j = juxta, p = process at juxtal base. Scale = 0.1 mm. 40 regard to the groundplan of the Sarcophaginae and probably with regard to that of Sarcophaga sensu lato as well: 1) Male hind coxa with a ventro- median pad of short spiny bristles; 2) Aedeagal juxta more or less dome-shaped and arching over the lateral styli; 3) Base of juxta with a pair of arms or processes; 4) Male cercus dorsally with a flat- tened or concave area and with a more or less distinct hump subapically; 5) Female mid femoral organ, if present, in apical position. Defined in this way, Discachaeta contains six species, all distributed in the western Palaearctic. (Note that the species Discachaeta gigas Povolny, 1986 was transferred to Heteronychia Brauer & Bergenstamm (Ezpierretia Rohdendorf) in an ad- dendum of the original paper.) The hypothesis that Sarcophaga amputata and S. kunonis are sister species is corroborated by the presence in these two species of at least five char- acter states that are judged to be derived with regard to the ground plan of Discachaeta and not found in any other species of this taxon: 1) Postgena with at least some black setae in anterior part, 2) Proanepi- sternum setose, 3) Abdominal tergite 5 almost de- void of microtomentum, 4) Male gonopod with long setae on the median surface (fig. 3), and 5) Female mid femoral organ reduced, i.e. indistinct or absent. Sarcophaga amputata is easily separated from S. kwnonis in both sexes by the greater exten- sion of white postgenal setae. These cover at least one half (strict lateral view) of the postgena in S. amputata while they are almost absent in S. kuno- nis. However, as the Madeiran fauna of Sarcopha- gidae may still be incompletely known, the shape of the aedeagus still provides the best clue to reliable identifications. Sarcophaga amputata may be widespread on the main island and is the first species of Sarcophagidae recorded from the Desertas islands. Sarcophaga (Pierretia) iulicida sp.n. (figs. 5-9) Type material. — Holotype 8, Portugal: Mouräo, ex Ommatoiulus moreleti, larva 8.v.1987, pupa 14.v.1987, adult 30.v.1987, P.T. Bailey (BMNH). The holotype is in good condition, with the puparium glued to a piece of cardboard and the dissected terminalia stored in glycerine in a microvial, both items pinned with the specimen. Description Male. — Head. Narrowest part of frons 0.25 X head width. Parafacial plate with a row of setae along the eyemargin. The upper part of the row consists of uniserial setae, the lower part is irreg- ularly biserial with 3-5 long bristles anterior to the x PAPE: New species of Sarcophaga setae. Arista with longest hairs about 4 x as long as second aristomere. Gena with black setae only, postgena with white setae only. Thorax. Proanepisternum bare. Chaetotaxy: 2-3 distinct pairs of presutural acrostichals, postsutural acrostichals (including prescutellars) not differen- tiated, dorsocentrals = 3 + 3, intra-alars = 1 + 2, supra-alars = 1 + 3, postalars = 2. Scutellum with 2 pairs of lateral scutellar bristles, 1 pair of short apicals, and 1 pair of discals. Mid femur with 3 anteroventral bristles and a row of posteroventrals. A pical posteroventrals slightly stronger than apical anteroventrals. Hind tibia without elongated setae. Abdomen. Tergites 3-4 with strong median mar- ginals on T5 with a complete row of marginals. Terminalia black, protandrial segment without marginal bristles. Aedeagus with well developed harpes, each of which has a narrow winglike pro- cess that is directed laterally and opposite to its counterpart. Vesica reduced to a simple swelling or hump. Juxta moderately long and deeply cleft. Length: 7 mm (estimated from dissected holo- type). Female. — Unknown. Puparium of the usual barrel-shape and with the posterior spiracles in a deep pit as in most species of Sarcophaginae. Distribution. — Palaearctic: Portugal. Figs. 5-9. Sarcophaga tulicida, male ter- minalia. 5, right cercus and surstylus, lateral view, setae omitted; 6, cerci, pos- terior view, setae omitted; 7, aedeagus, lateral view; 8, distiphallus, dorsal (= posterior) view; 9, right paramere + gonopod, lateral view. Abbreviations: h = harpes, j = juxta. Scale = 0.1 mm. 41 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Etymology. — A noun in apposition. From the Latin wluys = multiped, millipede, and -cida = suffix denoting cutter, killer, killing. The name refers to the millipede-parasitizing habit of the species. Remarks. — The present species belongs to a probably monophyletic group whose members all possess an aedeagus very similar to that of Sarco- phaga nigriventris Meigen. The group is defined by the characteristic and probably apomorphic shape of the aedeagal juxta, which can be dissolved into the two character states: 1) Juxta deeply cleft, and 2) Juxtal prongs tapering. Both states are readily seen on figs. 7-8. No formal genus-group name has been applied explicitly to this taxon, but the concept of Pierretia Robineau-Desvoidy (sensu stricto) of Verves (1986) is very close, the only difference being that this author includes Sarcophaga granu- lata Kramer, which does not possess either of the derived character states. I prefer to restrict the name Pierretia, as a subgenus of Sarcophaga, to the group of species possessing the two above menti- oned character states, ie. Sarcophaga discif- era/Pandellé, S. sulicida, S. lunigera Böttcher, S. nigriventris, S. socrus Rondani, $. soror Rondani, S. sororcula (Rohdendorf) and S. villeneuvei Böttcher. Sarcophaga tulicida may easily be separated from other members of the subgenus by the wing-like projection of each of the harpes which is directed laterally and set at right angles to the longitudinal axis of the aedeagus. This is most easily seen in dorsal (= posterior) view (fig. 8). The biology of Sarcophaga iulicida is interesting as very few Diptera are known to parasitise milli- pedes, and among the Sarcophagidae only species of the New World genus Spirobolomyia Townsend have been repeatedly bred from live millipedes. The habits of other species of the subgenus Pierretia (as defined above) indicate a broad to very broad spec- trum of hosts or prey. Thus, the species Sarcophaga nigriventris has been bred from snails, beetles, grasshoppers and bees (see references in Pape 1987). ACKNOWLEDGEMENTS I extend my sincere thanks to Mr J.C. Deeming, Cardiff, and Mr A.C. Pont and Mr N.P. Wyatt, both London, for loan of specimens in their care. REFERENCES Downes, W. L., Jr., 1965. Family Sarcophagidae. Pp. 933-961. In: A. Stone et alii (eds): A catalog of the Diptera of America north of Mexico. — Agriculture Handbook No. 276, Washington, D.C. Pape, T., 1986. The Sarcophagidae (Diptera) of Madeira, with the description of a new species of Discachaeta Enderlein. - Bocagiana 93: 1-4. Pape, T., 1987. The Sarcophagidae (Diptera) of Fenno- scandia and Denmark. — Fauna entomologia scandi- navica 19: 1-203, 2 pls. Pape, T., 1988. A revision of the Palaearctic Sarcophagidae (Diptera) described by C. Rondani. - Stuttgarter Bei- träge zur Naturkunde (Serie A) 416: 1-22. Povolny, D., 1986. Discachaeta gigas sp.n. und Heterony- chia (Heteronychia) vachai sp.n. - zwei unbekannte Sarcophaginae-Arten aus dem Dinarischen Küstenge- biet (Diptera, Sarcophagidae). — Acta Universitatis Agriculturae 34(2): 229-238. Séguy, E. 1941. Etudes sur les mouches parasites. 2. Cal- liphoridae, calliphorines (suite), sarcophagines et rhi- nophorines de l'Europe occidentale et méridionale. — Encyclopédie entomologique (A) 21: 1-436. Verves, Yu. G. 1986. Family Sarcophagidae. Pp. 58-193. In: A. Soös (ed.): Catalogue of Palaearctic Diptera. Vol. 12, Calliphoridae - Sarcophagidae. - Akadémiai Kiadó, Budapest. Received: 29 May 1989 Accepted: 17 July 1989 THOMAS PAPE Zoologisk Museum, Copenhagen REVISIONARY NOTES ON AMERICAN SARCOPHAGINAE (DIPTERA: SARCOPHAGIDAE) Pape, T., 1990. Revisionary notes on American Sarcophaginae (Diptera: Sarcophagidae). Tijdschrift voor Entomologie 133: 43-74, 66 figs. ISSN 0040-7496. Published 31 July 1990. Selected New World genera of Sarcophaginae are redefined: Argoravinia Townsend, 1917 is treated as a senior synonym of Raviniopsis Townsend, 1918, syn. n., Comasarcophaga Hall, 1931 as a senior synonym of Archimimus Reinhard, 1952, syn. n., and Emblemasoma Aldrich, 1916 as a senior synonym of Pessoamyia Lopes, 1938, syn. n. and Colcondamyia Reinhard, 1963, syn. n. Microcerella Macquart, 1851 is considered as synonymous to the tribe Microcerellini, except that Cryptosarcophila Townsend, 1931 is excluded, and 6 new names are proposed to replace junior secondary homonyms: Microcerella aulacophyto nom. nov. for Aulacophyto auromaculata Townsend, 1919; Microcerella austrohartigia nom. nov. for Austrohartigia bicoloricauda Lopes, 1981; Microcerella boettcherimima nom. nov. for Boettcherimima hypopygialis Lopes, 1950; Microcerella boetia nom. nov. for Boettia ecua- toriana Lopes, 1982; Microcerella carchia nom. nov. for Carchia andina Lopes, 1982; and Microcerella jujuyia nom. nov. for Jujuyia alvarengai Lopes, 1980. Microcerella bermuda sp. n. (Bermuda Is) and M. adelphe sp. n. (eastern Canada & USA) are described and the phylogenetic relationships to other members of the genus are discussed. Microcerella acridiorum (Weyenbergh, 1875) is considered a senior synonym of Xenoppiella dyscineti Blanchard, 1966, syn. n. Fletcherimyia Townsend, 1917 and Spirobolomyia Townsend, 1917 are discussed in a phylogenetic context and it is argued that both taxa should be excluded from Blaesoxipha (s.l.) |Protodexiini + Impariini]. Species limits within the genera are revised and Fletcherimyia abdita sp. n. (southeastern USA) and Spirobolomyia latissima sp. n. (southeastern USA & Mexico) are described. Tripanurga Brauer & Bergenstamm, 1891 is redefined and treated as a senior synonym of Metoposarcophaga Townsend, 1917, syn. n. and Erwcophaga Reinhard, 1963, syn. n. Tripanurga guatemala sp. n. (Guatemala) is described, and a probable sister group relationship to T. villipes (Wulp, 1895) is demon- strated. Infraspecific variation within T. aurea (Townsend, 1917) is discussed and the species is divided into a Baja Californian form and a mainland form. Dr T. Pape, present address: Danish Bilharziasis Laboratory, Jaegersborg Allé 1 D, DK-2920 Charlottenlund. Key words. — Diptera; Sarcophagidae; generic redefinitions; new species; New World. CONTENTS INTRODUCTION Imrtoductionsv een NUE ee 43 The New World fauna of Sarcophagidae deviates Methods, terminology, depositories............ 44 markedly from that of the Old World with regard Generic redefinitions and descriptions of to diversity. The subfamily Miltogrammatinae has SPÉCIES Rae ne RL N or sdraiata 45 significantly fewer species, which in part is to be GenustAncoraviniane RAMA 45 expected as the fauna of the Palaearctic Region is GENUSUCO4ANEOPPAZARA IO 45 much more fully known and the extension of gener- GeUStE7 IENA OMAR OR 46 ally species-rich areas as dry savannas, shrublands Genussilerchenn na 47 and semi-deserts is much larger, e.g. on the African Genus Vierocenell a RENE ER EE 48 mainland and in Central Asia. Within the subfamily GENUSS piobolon pn 53 Sarcophaginae, on the other hand, species richness GENIUS ANZIO 55 per se is greater in the New World, and as the cknowledoemenesn O 58 sarcophagine fauna of especially the neotropics, References TNA leeren. 58 where the large majority of species occur, is still 43 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 very insufficiently known, this difference may be- come even more pronounced with time. The New World fauna of Sarcophaginae has diversified morphologically and biologically to a degree unri- valled by the Old World members of this subfam- ily. Some New World species have green eyes, several have a metallic abdomen, and variations in setosity and configuration of the male termina- lia are legio and often much more radical than those of the Old World species. The New World species Peckia gulo and Tricharaea occidua repres- ent the largest and the smallest species of Sarco- phaginae, respectively, and sarcophagine life hab- its include parasites of millipedes and insects, as well as predators of reptile eggs, hymenopteran kleptoparasites and generalized scavengers. The much greater morphological and biological diver- sity of New World species of Sarcophaginae com- pared to those of the Old World has caused pro- nounced differences of opinion regarding generic limits within this subfamily (especially so, because many of the previous authors have been more in- terested in differences than in similarities). Nu- merous monotypic genera have been proposed, with no other warranty than their ‘uniqueness’ and/or weirdness. My research on New World Sarcophaginae, espe- cially in connection with a major conspectus of the genus Blaesoxipha covering species limits and in- ter- and infrageneric phylogenetic relationships (Pape in prep.), during a one-year fellowship at the National Museum of Natural History (USNM), Washington, D.C., has led me to propose some phylogenetically more consistent generic defini- tions. It is felt that these definitions, in addition to being theoretically sound, support identification and information retrieval purposes better, and it is considered appropriate to make the information available as a single paper, with descriptions of several hitherto unrecognized species which turned up during my study, rather than split it into several generic revisions and isolated descriptions of spe- cies. Thus, it should be stressed that the present paper is far from a comprehensive treatment of all New World genera, and the fact that most genus- group taxa have been excluded does not mean that these are accepted by me. Older genus-group taxa such as Pachygraphia Brauer & Bergenstamm, 1891, Lepidodexia Brauer & Bergenstamm, 1891, and Johnsonia Coquillett, 1895 are especially in need of a critical revision and will probably have to be redefined as much broader entities in order to absorb the numerous ‘less aberrant’ and probably paraphyletic genera that often are considered ‘closely related’ to these genera. Much remains to be done before generic limits stabilize, and more tho- rough character analyses are needed before the taxa above species level can emerge as well corroborated monophyletic groups. 44 METHODS, TERMINOLOGY, DEPOSITORIES Specimens were dissected by conventional cut- ting of terminalia and heating in potassium or so- dium hydroxide for about five minutes with subse- quent rinsing in distilled water and transfer through alcohol to glycerine. Drawings of termina- lia were made from glycerine preparations, either directly by using an ocular grid or from semiper- manent slide mounts in glycerine gel using a pro- jecting compound microscope. Names of genus-group and species-group taxa mentioned in the text are given without the con- ventional citing of the original author except where this has any direct bearing on nomenclature, e.g. in the lists of synonyms. According to the Interna- tional Code on Zoological Nomenclature, authors do not form part of the name proper (ICZN: 51,a), and they often serve no other purpose than as a reference to the original source of the name. Recent revisions are usually a better authority on the iden- tity of a species or genus than the original descrip- tion. For the present paper, identities of taxa not explicitly defined are easily retrieved from the ref- erences cited in the associated discussion or from the most recent regional catalogues, i.e. Downes (1965) and Lopes (1969), but see also Shewell (087) Morphological terms follow McAlpine (1981) except that I have adopted Sabrosky’s (1983) use of 'microtomentum’ for the dense cuticular micro-pu- bescence that may give surfaces a pollinose or prui- nose appearance. As no generic cladogram of the Sarcophaginae has been produced in the present or any other paper I have made no attempt of evaluating transforma- tion series polarities for the character states used in the generic diagnoses, i.e. whether apomorphic or plesiomorphic at the level in question (which would be nothing but ‘qualified guesses’ based on my own more or less explicitly developed ideas of generic phylogeny). The character states listed in the diag- noses are, of course, only a fraction of those known, and I have restricted the lists to contain only those states that seem to be unique (and therefore possi- bly autapomorphic) as well as states judged by me to possess ‘diagnostic power’ (i.e., they will in com- bination serve to define the taxon in question). The institutions from which material examined or otherwise referred to is deposited have been abbreviated with the following acronyms: AMNH: American Museum of Natural History, New York, USA; CAS: California Academy of Sciences, San Francisco, USA; CNC: Canadian National Collection of Ar- thropods, Ottawa, Canada; FSCA: Florida State Collection of Arthropods, Gainesville, USA; MACN: Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina; MCZ: Museum of Comparative Zoology, Cam- bridge, USA; MNHN: Museum National d'Histoire Naturelle, Paris, France; NMW: Naturhistorisches Museum Wien, Austria. UNAM: Instituto de Biologia, Universidad Na- cional Autónomia de Mexico; USNM: National Museum of Natural History, Washington, D.C., USA; ZML: Zoologiska Museet, Zoologiska Institu- tionen, Lund, Sweden; ZMUC: Zoological Museum, University of Co- penhagen, Denmark. GENERIC REDEFINITIONS AND DESCRIPTIONS OF SPECIES Genus Argoravinia Argoravinia Townsend, 1917b: 190, 193, 195. Type spe- cies: Sarcophaga argentea Townsend, 1912 (= Sarco- phaga rufiventris Wiedemann, 1830), by original de- signation. Raviniopsis Townsend, 1918: 160. Type species: Ravint- opsis aurea Townsend, 1918, by original designation. Syn. n. Sarcophagina Curran, 1928: 102. Type species: Sarcopha- gina candida Curran, 1928, by original designation. Pachygraphomyia Hall, 1933: 259. Type species: Pachy- graphomyia spinosa Hall, 1933, by original designa- tion. Diagnosis. — The genus Argoravinia may be defined by the following character states: 1) Aedeagus with a very long and S-shaped me- dian stylus, see, e.g., Roback (1954: fig. 39); 2) One or more setae on the underside of the stem of r,,, elongate; 3) Male cercal prong bent backwards; 3) .Ejaculatory apodeme large. As tentatively suggested by Pape (in press), the genera Argoravinia and Raviniopsis should be treated as synonyms. They have been included in a subtribe of their own (within Sarcodexiini) by Lopes (1975a, 1982e), the monophyly of which al- though never corroborated by explicit reference to probable apomorphies has never been questioned. Lopes (1982e) lists a number of character states, but only some of these can be considered derived with regard to the groundplan of the Sarcophaginae, and this author does not propose any most probable sister group. The two very distinctive character states 1-2 mentioned above seem to be autapomor- phic and will fully define the genus. The remaining two character states may diagnose the taxon further, but these may be plesiomorphic at the level of Argoravinia (i.e. define a more inclusive group) as discussed under Tripanurga. PAPE: American Sarcophaginae Having corroborated the monophyly of Argora- vinia, it remains to be discussed why a splitting is opposed. One argument is the small size of this taxon, with a total of 7 species recognized at pres- ent. Even if monophyletic subgroups can be recog- nized (and in theory a complete resolution into dichotomies should be possible), nothing is gained by erecting formal genus-group names if one then has to create new family-group names (e.g. tribes and subtribes) to keep the related taxa together. Another, more formal, argument is the subgroups proper. Argoravinia in the narrow sense of Lopes (1976), including A. rufiventris and A. alvarengat, is most probably monophyletic, defined by the aut- apomorphic process from the posteroventral mar- gin of the epandrium that partially covers the sur- stylus, and by the much more distinctly S-shaped median stylus compared to the remaining Argora- vinta (see figs. in Lopes 1976 and 1988b). I have not found any character state shared only by these re- maining species, which were treated as Raviniops by Lopes (1988b), that may be considered apomor- phic at this level, and none have been suggested. If the monophyly of the taxon cannot be corroborated, it has to be either split or widened until fitting a pattern for which character state distributions cor- roborate a hypothesis of strict monophyly. In the present case, the obvious choice is a merging with the older Argoravinia. Genus Comasarcophaga Comasarcophaga Hall, 1931: 280. Type species: Comasar- cophaga texana Hall, 1931, by original designation. Tejasomyia Reinhard, 1945: 68. Type species: Tejasomyta nexilis Reinhard, 1945, by original designation. Archimimus Reinhard, 1952: 140. Type species: Archimz- mus camatus Reinhard, 1952, by original designation. Syn. n. Diagnosis. — The genus Comasarcophaga may be defined by the following character states: 1) Male mid femur apically with a short comb-like row of posteroventral spines; ) Male cercal prong bent backwards; ) Aedeagal juxta slightly displaced ventrally rel- ative to longitudinal axis of aedeagal tube, the latter thereby appearing 'hump-backed' (figs. 1-4, additional figs. in Lopes 1979 and Lopes & Tibana 1988); 4) Aedeagal vesica appearing more or less square in lateral view (cf. figs. mentioned above). D N The genus Comasarcophaga was described by Hall (1931) as a monotypic genus for his new spe- cies C. texana. Downes (1965) synonymized Coma- sarcophaga with Tejasomyia, thereby adding the species C. nexilis, and apparently only these two species were included in the genus by Shewell (1987: 1168), who gave "2 spp.; Texas, California”. Another species, Sarcophaga prolepsis, described 45 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 by Reinhard (1947) from Arizona, was transferred by Downes (1965) to Blaesoxipha subgenus Servai- sia, and later by Lopes (1988d) to Archimimus. Lopes & Tibana (1988: 324) defined Archimimus in part by the features: "styli of glans and median process spinous” (lateral and median styli in my terminology) and "arista plumose to tip”. However, by including prolepsis and longespinus (Lopes 1988d), which both possess simple lateral and me- dian styli with small denticles as well as a pubescent or short-plumose arista with a distinctly bare tip, conditions perfectly like those found in most other Sarcophaginae, the definition breaks down at least for these characters. Other character states listed by Lopes & Tibana (1988) as diagnostic for Archimz- mus are: 3-Á post-dorsocentral bristles, proanepis- ternum bare, male mid femur with apical comb-like posteroventral spines, aedeagal vesica strongly sclerotized, and female tergite 8 (7 in my terminol- ogy) setose. These states, however, do not secure an unambiguous definition of the taxon as this combi- nation is found in other genera, e.g. Blaesoxipha in part and Comasarcophaga. Of the four states listed in the definition above, 1 and 2 are found in several other genera. Only 3 and 4 seem to be unique (i.e., autapomorphic), but, unfortunately, they are less clear cut and more dif- ficult to describe as are many other characters of the male terminalia. Comasarcophaga in the present definition may be divided into two groups based on similarity (as perceived by me) of the male termi- nalia: one containing longispinus, nexilis, prolepsis and texana, another containing the remaining members. This agrees with Lopes (1988d: 918), who mentioned that longispinus and prolepsis was “provisionally included in Archimimus in spite of some important differences from the type-species: last female sternite large and clypeal arch of the first instar larva complete’. Differences as such, however, cannot reject phylogenetic relationships, and as I have found no better way to describe the species explicitly in terms of shared character states, lumping all Archimimus with nexilis and texana in a broader Comasarcophaga is necessary to maintain the monophyly of the taxon. Terminalia of species of Comasarcophaga not figured by Lopes (1979) and Lopes & Tibana (1988) have been illustrated in figs. 1-12. The only biological information available for Co- masarcophaga is a single breeding record of a male C. texana from the bulimulid snail Rabdotus deal- batus, and the fact that another male of this species has been collected from a dysentery fly trap (Neck & Lopes 1973). Genus Emblemasoma Emblemasoma Aldrich, 1916: 56. Type species: Emble- masoma erro Aldrich, 1916, by original designation. 46 Pessoamyia Lopes, 1938: 333. Type species: Pessoamyia prosternalis Lopes, 1938, by original designation. Syn. n. Colcondamyia Reinhard, 1963a: 82. Type species: Colcon- damyia falcifera Reinhard, 1963a, by original designa- tion. Syn. n. Golcondamyia Reinhard, 1963b: 152. Unjustified emenda- tion of Colcondamyia. Diagnosis. — The genus Emblemasoma may be defined by the following character states: 1) Prosternum broadened (Shewell 1987: fig. 28); 2) Male mid femur apically with a row of comb- like posteroventral spines. When Lopes (1938) described Pessoamyia he did not present any hypothesis as to possible relation- ships to other genera, and the main reason for describing the genus apparently was the "proster- num extraordinariamente desenvolvido” (p. 333). Later, Lopes (197 1b: 89) presented a "study of spe- cies showing large and inflated prosternum”, al- though without commenting on the similarly equipped Colcondamyia, and he uncritically ac- cepted the validity of the two genera discussed (ie. Emblemasoma and Pessoamyta). Actually, he did not even mention whether or not he considered the enlarged prosternum as indicating phylogenetic re- lationship, but he later erected the subtribe Emble- masomatina (Lopes 1975b, as tribe in 1982e and 1988c), including all the above-mentioned genera (Lopes 1982e), and he explicitly recognized the modified prosternum to be autapomorphic within the Sarcophagidae (Lopes 1988c). As shown in the list of synonyms given above, I prefer to reduce the (sub)tribe to the generic level, and at least three reasons should be considered. Firstly, the comparatively small size of the entire taxon (i.e. Emblemasoma in the present sense), does not, in my opinion, necessitate a generic sub- division to enhance the clarity of the phylogenetic relationships between the included species. Admit- tedly, a group consisting of Pessoamyia and Em- blemasoma sensu Lopes (1988c) may probably be monophyletic based on the shared possession of a highly modified male cercus, which in profile has a subapical dorsal knob and ends in a blunt hook (Lopes 1988c: figs. 17, 42, 45), but I prefer to name this assemblage of species the erro species-group rather than use a formal genus-group name. Se- condly, while the monophyly of Emblemasoma in the present sense seems reasonably well corrobo- rated (the species are also biologically similar in being parasites or predators of insects, as far as known), the monophyly of the groups recognized by Lopes (1988c) seems much less corroborated. Thus, the only derived character states defining the taxon Pessoamyia seem to pertain to the aedeagus (vesica reduced, vesica spinous, ‘lateral plates’ hya- line, see key in Lopes 1975b), and Lopes (1988c: 17) admitted that males are needed to decide whether species for which only females are known “belong to Pessoamyia, Emblemasoma or to a distinct ge- nus’. Thirdly, the enlarged prosternum provides an easy-to-see feature for the recognition of the taxon and thus of value from the point of view of the non- specialist (e.g. when using a generic key). Note that Pape (1987) included Emblemasoma auditrix in a wide concept of Blaesoxipha, and the bent male cercus and the fused abdominal sternites 6-8 of auditrix are indeed reminiscent of the condi- tion found in some species of Blaesoxipha (Soper et al. 1976). However, my revised concept of the latter genus (Pape in prep.) includes all sarcophagine species with the lateral styli non-conducting and fused ventrally through a median sclerotization (?vesica). Therefore, the three free and apparently sperm-conducting (i.e. hollow and communicating with the spermduct) styli of Emblemasoma com- bined with the lack of a row of trochanteral spines provides evidence that neither E. audıtrıx nor all of Emblemasoma should be included in Blaesoxipha (see also discussion under F/etcherimyia and Spiro- bolomyia). Evidence for either placement of audz- trix, however, is rather sparse, and I prefer the present inclusion of this species in Emblemasoma mainly because an inclusion within Blaesoxipha would require that the free styli should be consi- dered a reversal, which so far has not been shown to occur in any other species of Sarcophaginae. A detailed numeric cladistic analysis of all sarcopha- gine genera is needed to test this hypothesis. Genus Fletcherimyia Fletcherimyia Townsend, 1917b: 191. Type species: Sar- cophaga fletcheri Aldrich, 1916, by original designa- tion. Peltopyga Townsend, 1917b: 191, 194. Type species: Sar- cophaga celarata Aldrich, 1916, by original designation. Diagnosis. — The genus Fletcherimyia may be defined by the following character states: 1) Male mid femur apically with a short comb-like row of posteroventral spines; 2) Male cercal prong bent backwards; Aedeagus with lateral styli shortened; 4) Aedeagal vesica a single tongue-shaped struc- ture (figs. 13-18); 5) Aedeagal juxta with cuticular pubescence along the distal margin (figs. 13-18); 6) Female abdominal tergite 6 strongly convex; 7) Female abdominal sternites 6-7 fused; Larvae unable to develop to maturity outside pitchers of Sarracenia. Character states 5 and 6 seem to be autapomor- phic. Fletcherimyia and Peltopyga, both monotypic, were described in the same paper by Townsend PAPE: American Sarcophaginae (1917b). When Townsend (1938) later redescribed these genera, he still included only the type species in Peltopyga while he did not state how many spe- cies were included in F/etcherimyia. Downes (1965) catalogued the Nearctic species and treated Fletche- rimyia and Peltopyga as synonyms, including four similar species, all of which were known to deposit their larvae in pitchers of Sarracenia, in Fletcheri- myia as subgenus of Blaesoxipha. This apparently obligate association with pitcher plants is reflected in the distribution in that all species of Fletcherimyia have their center of distri- bution in the eastern part of the continent. Buckell & Spencer (1957) listed Sarcophaga fletcheri from Canada (Robson, B.C.), but I have been unable to confirm this and the record was apparently not accepted by Downes (1965). Species of Sarracenia are mainly eastern in distribution although the most wide-ranging species, $. purpurea, reaches far west to the District of Mackenzie and the extreme northeastern part of British Columbia (Cody & Tal- bot 1973). Sarracenia purpurea is known from one locality only in B.C., a peat bog approximately 30 km south of Fort Nelson (Krajina 1968), and the record of fletcheri from Robson seems unlikely. It should be expected that species of Fletcherimyia require a certain density of Sarracenia localities to be able to maintain stable populations. Males of Fletcherimyta can often be seen resting on the rim of the Sarracenia pitchers, and these may serve as the aggregation site. Females larviposit large first stage larvae singly into newly opened pitchers, and the larvae feed on invertebrates caught in the fluid. The larva lives submerged, but with the posterior end kept at the water surface to facilitate respiration. The mature larva leaves the pitcher and pupates at the base of the plant (For- syth & Robertson 1975, Fish 1976, Fish & Hall 1978). A few other species of Sarcophagidae breed in the pitchers of insectivorous plants: The Nearctic Sar- cophaga (Liosarcophaga) sarraceniae in Sarracenia pitchers (Aldrich 1916) and the Oriental Sarco- phaga (Pierretia) urceola in Nepenthes pitchers (Beaver 1979, Shinonaga & Beaver 1979), but these are easily shown by morphological evidence to have evolved the life habit independently from species of Fletcherimyia, and even from each other, as indi- cated by the subgeneric assignment given in paren- theses. Roback (1954) gave figures of the aedeagus of the four species known to him, but instead of straight- ening out the homologies of the acrophallic sclero- tizations, he simply introduced the morphological term 'stematis’ for the type of acrophallus found in Fletcherimyia. Lopes (1971a: 4) stated that "the opening [of the acrophallus] is very much compli- cated” and proposed that it had “a different origin” from that of Blaesoxipha. This, however, seems 47 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 little warranted as there is no evidence that the sperm exit proper is not homologous throughout the Sarcophagidae (and probably all of Diptera), and therefore may serve as an important morpho- logical landmark. If this is accepted, Fletcherimyia falls nicely within the groundplan pattern of the Sarcophaginae in that all species possess three styli which all seem to be more or less tubular and con- nected to the common sperm duct (fig. 18). With the additional species described below, the genus contains five species, but some other species should be mentioned as they have been assigned to the genus Fletcherimyia. Lopes (1946) described F. speciosa from Mexico and included Sarcophaga ces- sator as well, and Rohdendorf (1971) described Fletcherimyia zayasi from Cuba. These three spe- cies, however, all belong to Blaesoxipha, the mo- nophyly of which is corroborated mainly by the transformation of the lateral styli into non-conduct- ing structures and a sclerotized ventromedian bridge connecting these (Pape unpubl.). It should be stressed that although Lopes (1971a: 5) stated that Fletcherimyia (and some other taxa) did "not belong to Tephromyiini |= the present Blaesoxi- pha in part] simply because they were "good genera”, it is perfectly possible to argue for the inclusion of Fletcherimyia within Blaesoxipha, e.g. as a subgenus (Downes 1965), simply by widening the concept of Blaesoxipha sufficiently (e.g. by de- fining this genus on the comb-like pv bristles of the male mid femur). This does not, of course, alter the well corroborated hypothesis that species of Flet- cherimyia as presently recognized, form a mono- phyletic group. My reason for not including Flet- cherimyia within Blaesoxipha is primarily that the resulting taxon, although possibly monophyletic if also containing Comasarcophaga and Spirobolo- myia discussed elsewhere in the present paper, would be much more vaguely defined in the sense that the monophyly would be much more tentative. Considering the well corroborated monophyly of Blaesoxipha, Comasarcophaga, Fletcherimyia and Spirobolomyia respectively, as well as their rather different external morphology, I see no convincing arguments for combining them into one taxon. It could be argued that the combined taxon would be more easy to identify in conventional sorting of specimens, as dissections of the male terminalia, which often are necessary for the non-specialist and therefore prevent handling a large number of spe- cimens, would be reduced to a minimum. I still prefer, however, to give the criterion of monophyly higher priority than that of usefulness. Fletcherimyia abdita sp. n. (figs. 13, 19, 24) Type material. — Holotype 4, USA: Alabama, Theo- dore, reared from Sarracenia drummondii, 10.vi.1916, 48 FM. Jones [paratype of Sarcophaga rileyi, cf. Aldrich (1916: 243)] (USNM). Paratypes. - USA: 14, data as holotype but with the date 22.vi.1916 [paratype of Sarco- phaga rileyi) (USNM); 26, locality as holotype but with- out date (FSCA, ZMUC); Florida, Freeport, 26, 25.v1.1921, [no collector, label in Jones’ handwriting], reared from Sarracenia drummondii (FSCA); Mississippi, Biloxi, 18, 6.vi.1921, 28, 23.vi.1921, 28, 25.vi.1921, FM. Jones, reared from Sarracenia sledgei (4 in FSCA, 1 in ZMUC). Description Male. — General morphology similar to other species of Fletcherimyia (see descriptions in Al- drich 1916), but separated from these by the struc- ture of male abdominal sternite 5 and male termi- nalia (compare figs. 13-28). The following description is restricted to these diagnostic features. Abdomen: Posterior margin of sternite 5 with accessory lobes like those of F. celarata but slightly raised from the plane of the remaining sternite so that a small flange penetrates behind (morpholo- gically above) it (fig. 19). Sternite 5 of F. celarata has accessory lobes broadly rounded, level with remain- ing sternite (fig. 20), and dark brown along poster- ior margin in contrast to the yellowish remaining sternite. Sternite 5 of F. rileyt has tapering accessory lobes and posterior margin concolorous with re- maining sternite (fig. 23). Terminalia: Cercus, as seen in posterior view, with an outline intermediate between that of F. fletcheri and F rileyi. The aedeagus is similar to that of E fletcheri, but the juxta is broader and more square and the vesica is larger with a spinelike process at about middle (fig. 13). Female. — Unknown. Etymology. — A Latin adjective, from abditus = hidden, concealed, put away. The name refers to the fact that the holotype was found amongst paratypes of F. rileyi. Distribution. — Nearctic: USA (Alabama, Flor- ida, Mississippi). Biology. — Larvae scavengers in pitchers of Sar- racenia drummondu and S. sledget. Genus Microcerella Microcerella Macquart, 1851: 209 (236). Type species: Microcerella rufomaculata Macquart, 1851, by original designation. Doringia Weyenbergh, 1875: 85. Type species: Nemorea acridiorum Weyenbergh, 1875, by original designation. Syn. n. Euparaphyto Townsend, 1912: 359. Type species: Expara- phyto alpina Townsend, 1912, by original designation. Syn. n. Xenoppia Townsend, 1915: 20. Type species: Xenoppta hypopygialis Townsend, 1915, by original designation. Camptopyga Aldrich, 1916: 41. Type species: Camptopyga aristata Aldrich, 1916 [= Xenoppia hypopygialis Townsend, 1915], by original designation. Hypopelta Aldrich, 1916: 49. Type species: Hypopelta scrofa Aldrich, 1916, by original designation. Aulacophyto Townsend, 1919: 158. Type species: Aulaco- phyto auromaculata Townsend, 1919 [secondary junior homonym of Euparaphyto auromaculata Townsend, 1919: 157] [= Microcerella aulacophyto nom. nov.|, by original designation. Syn. n. Gymnopsoa Townsend, 1919: 161. Type species: Gym- nopsoa texana Townsend, 1919, by original designa- tion. Catheteronychia Townsend, 1927: 230. Type species: Cat- heteronychia chaetosa Townsend, 1927, by original designation. Syn. n. Xanthobrachycoma Townsend, 1927: 232. Type species: Xanthobrachycoma analis Townsend, 1927, by original designation. Syn. n. Townsendmyia Prado & Fonseca, 1932: 167. Type species: Stephanostoma argenteum Prado & Fonseca, 1932 |= Sarcophaga (Bellieria) halli Engel, 1931], by original designation. Syn. n. Alaccoprosopa Townsend, 1934: 202. Type species: Alac- coprosopa apicalis Townsend, 1934, by original desig- nation. Syn. n. Austrohartigia Townsend, 1937a: 115. Type species: Aus- trohartigia magellanica Townsend, 1937a [= Sarco- phaga spinigena Rondani, 1846], by original designa- tion. Syn. n. Microcerellamyia Hall, 1937: 354. Type species: Microcer- ellamyia retusa Hall, 1937, by original designation. Syn. n. Itiophyto Hall, 1937: 355. Type species: Itiophyto engeli Hall, 1937, by original designation. Syn. n. Mimophytomyia Hall, 1937: 357. Type species: Mzmo- phytomyia chilensis Hall, 1937 |= Alaccoprosopa api- calis Townsend, 1934], by original designation. Phaesarcodexia Hall, 1937: 362. Type species: Phaesarco- dexia aldrichi Hall, 1937 (also as Brachicoma aldricht) [= Sarcophaga spinigena Rondani, 1864], by original designation. Eumicrocerella Hall, 1938: 253. Type species: Eumicrocer- ella duca Hall, 1938, by original designation. Syn. n. Scopaediscus Blanchard, 1939: 802. Type species: Scopae- discus muebni Blanchard, 1939, by original designa- tion. Syn. n. Boettcherimima Lopes, 1950: 706. Type species: Boettcherimima hypopygialis Lopes, 1950 [secondary junior homonym of Xenoppia hypopygialis Town- send, 1915] [= Microcerella boettcherimima nom. nov.], by original designation. Syn. n. Parabypopelta Blanchard, 1955: 30. Type species: Parahy- popelta salavini Blanchard, 1955, by original designa- tion. Syn. n. Chapiniola Dodge, 1965: 253. Type species: Chapiniola impressa Dodge, 1965, by original designation. Syn. n. Steatopyga Dodge, 1965: 254. Type species: Steatopyga mirabilis Dodge, 1965, by original designation. Syn. n. Boetia Dodge, 1965: 255. Type species: Boetia curiosa Dodge, 1965, by original designation. Syn. n. Xenoppiella Blanchard, 1966: 182. Type species: Xeno- piella dyscineti Blanchard, 1966 |= Nemorea acridio- rum Weyenbergh, 1875, syn. n.], by monotypy. Syn. n. Azuayia Dodge, 1967: 681. Type species: Azuayıa tripar- tita Dodge, 1967, by original designation. Syn. n. Borgmeterisca Lopes, 1972: 349. Type species: Borgmeier- PAPE: American Sarcophaginae isca pilicoxa Lopes, 1972, by original designation. Syn. n. Townsendisca Lopes, 1974: 193. Type species: Townsen- disca matucanensis Lopes, 1974, by original designa- tion. Syn. n. Xenoppina Lopes, 1975d: 573. Type species: Xenoppina andina Lopes 1975d, by original designation. Syn. n. Aulacophytoides Lopes, 1978a: 765. Type species: Az/aco- phytoides alvarengai Lopes, 1978a, by original desig- nation. Syn. n. Jujuyia Lopes, 1980: 238. Type species: Jujuyia alvarengat Lopes, 1980 [secondary junior homonym of Az/aco- phytoides alvarengai Lopes, 1978a] [= Microcerella jujuyia nom. nov.], by original designation. Syn. n. Mallochisca Lopes, 1982b: 364. Type species: Mimophy- tomyia mallochi Hall, 1937, by original designation. Syn. n. Penaisca Lopes, 1982b: 365. Type species: Penaisca qui- maliensis Lopes, 1982b; by original designation. Syn. n. Cuzcomyia Lopes, 1982b: 367. Type species: Cuzcomyta rufipes Lopes, 1982b, by original designation. Syn. n. Carchia Lopes, 1982b: 368. Type species: Carchia andina Lopes, 1982b [secondary junior homonym of Xenop- pina andina Lopes, 1975d; [ Microcerella carchia nom. nov.], by original designation. Syn. n. Azuaya: Lopes (1982b: 363); erroneous subsequent spell- ing of Azuayıa. Boettia: Lopes (1982b: 363, 1982d: 607); erroneous sub- sequent spelling of Boetza. Parabybopelta: Lopes (1969a: 8, 81); erroneous subse- quent spelling of Parahypopelta. Phaeosarcodexia: Lopes (1969a: 8, 82; 1981: 327); errone- ous subsequent spelling of Phaesarcodexia. Townsendimyia: Lopes (1969a: 9, 1974: 195, 1975a: 271, 1982b: 363); erroneous subsequent spelling of Town- sendmyia. Diagnosis. — The genus Microcerella may be defined by the following character states: 1) Eyes green (live or fresh material); 2) Male protandrial segment black; 3) Hypandrial arm swollen at level of gonopod; 4) Postgena with at least some black setae close to genal suture. The generic synonymizations presented above are straightforward in the sense that the identities of the type species of most genus-group names are well known and the monophyly of what is here considered as Microcerella has never been questi- oned. One exception is the taxon Xenoptella dys- cineti, which was mentioned as being a potential member of the group by Lopes (1982b) but left with no further comment, as the holotype (which is the only specimen known) was not examined. This uncertainty resulted from the insufficient original description by Blanchard (1966), who gave a not very informative description of the male terminalia and did not provide any figures, only giving some chaetotaxic characters and stating that the termina- lia were "muy abultado” (p. 183). I have examined the male holotype (MACN), which was mounted on cardboard and completely hidden in dense 49 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 mould. After partly cleaning of the specimen and remounting of the terminalia, it turned out to be a typical member of Microcerella (s.l.) and a junior synonym of M. acridiorum, syn. n. | Note that I have not revised the type material pertaining to the lat- ter name, and I have followed Lopes (1969b), who reviewed its taxonomic history.] To corroborate this, and to facilitate recognition and identification, figures of the terminalia of the holotype of X. dys- cineti are provided (figs. 29-32, compare Lopes 1969b: figs. 55-58). The holotype, which is perfectly intact except for dissected (but well preserved) ter- minalia, is heavily moulded and glued to a piece of cardboard on its right side. Terminalia and an empty puparium are mounted on separate pieces of cardboard pinned with the specimen. It is labeled "S/ Dyscinetus gagates, adulto.”, "Tandil 6-XI-1958 Bazan, R..”, "Xenopiella dyscineti ‚gen. et sp. nov. det. E. E. Blanchard”, and "Xenopiella dyscineti E. E. Blanchard 1966 Holotypus [red label]”. Lopes (1982b) noted that the monotypic genus Sarcophagulopsis (type species: S. trigonophymt) possibly belonged to the Microcerellini. It is, how- ever, a Blaesoxipha sensu lato and will be revised by Pape (in prep.). Somewhat ironically, the identity of the type spe- cies of the genus Microcerella proper, M. rufomac- ulata, has never been thoroughly revised. Séguy (1925: 184) and Townsend (1931: 72, 1937b: 202) examined and redescribed the female holotype in MNHN, leaving no doubt of the assignment to Microcerella in the present sense, but no attempt has ever been made to associate the female with male specimens, for which reason the identity is highly uncertain. As other species are known from the male sex only, it is very probable that M. rufo- maculata is a senior synonym of one of these [an obvious candidate being M. sarcophagina, the male holotype of which, however, may be lost, as men- tioned below |. To accommodate all species of the taxa synony- mized within Mzcrocerella above, a number of new names have been proposed. These names, which all should be treated as nouns in apposition, are formed by using the name of their former genus as species epithet. Two other secondary homonyms are not covered in the list of generic synonyms. They are given below, following the procedure given above: Microcerella boetia nom. nov. for Boettia [sic!] ecuatoriana Lopes, 1982d: 613 [secondary junior homonym of Euparaphyto ecuatoriana Lopes, 1982c: 495]. Microcerella austrohartigia nom. nov. for Austro- hartigia bicoloricauda Lopes, 1981: 334 [secondary junior homonym of Mesothyrsia bicoloricauda En- derlein 019287151]: 50 The genus Mrcrocerella is defined in the most narrow sense by Lopes (1969a) in the Neotropical catalogue, as he includes only one species, M. sar- cophagina, besides the type species M. rufomacu- lata. The taxon M. sarcophagina, like M. rufomac- ulata, has remained enigmatic since its description by Thomson (1869), and the male holotype, which is the only specimen known, seems to be lost. [P. I. Persson (NMRS) has informed me that he has been unable to recover the holotype, which was sent as a loan to the late H. R. Dodge in the 1970’s, and I did not succeed in finding it among the remnants of the Dodge-collection in FSCA. | Without any clue as to the structure of the terminalia, especially the aedeagus, the identity can only be a qualified guess whenever a detailed generic revision with extensive information of species present in the type locality concerned becomes available. Lopes (1982b) provides a key to the genera of Microcerellini, and his Microcerella keys out mainly on the length of the first flagellomere relative to that of the pedicel, i.e. whether the former is “twice the length” (Microcerella) or only “a little longer” (Microcerellamyia) than the latter, and his meas- urements are apparently based partly on the ori- ginal figures of Macquart (1851, pl. 22: fig. 1). It seems to me, therefore, that although Microcerella rufomaculata fits well within the tribe Microcerel- lini, and thus within Mzcrocerella in the present sense, the narrow concept of Microcerella sensu Lopes needs revision and is tentative at best. Lopes (1982b) discussed the limits of Microcer- ellini and mentioned Austrobartigia, Doringia, and Aulacophyto with ten, four, and five species respec- tively (che latter now with an additional three des- cribed by Tibana & Lopes (1988)), and he noted that “the majority of the remaining [27] genera are monotypic’ (p. 359). Note that Lopes (1982b) di- vided the Microcerellini into the subtribes Micro- cerellina and Hypopeltina. A single two-state char- acter separates these two groups, namely the possession of a pubescent versus a plumose arista (key entry only), but obviously both states cannot be assumed a priori to be apomorphic at this level as no third state from which they could be derived independently exists within the Sarcophaginae. No further explanation is given and the classification appears to be artificial. I have not been able to split the tribe into a few well defined and probably mo- nophyletic groups, and instead of accepting the wealth of monotypic or very small genera that do not convey much information (and actually often are a burden for identification and information re- trieval), I have lumped all species into one genus. Downes (1965) and Shewell (1987), who are deal- ing exclusively with the much sparser Nearctic fauna, also apply a broader concept when they in- clude four species within Mzcrocerella, one of which has been recorded from the neotropical region (M. hypopygialis, see Lopes 1980, 1982a). Microcerella in this broadened sense is very well defined and equals the tribe Microcerellini as presented in Lopes (1982b, 1982e), except that I exclude the monotypic genus Cryptosarcophila as discussed below. Lopes (1969a: 7, 1982b: 361) mentioned that all species seen by him in fresh condition have green eyes, truly a unique condition within the Sar- cophagidae. The green colour disappears after death although the eyes may retain a feeble green- ish tinge (at least for some years). I have not seen live or fresh specimens of any of the Nearctic spe- cies, and although at least M. texana seems to be green-eyed (as judged from recently captured spe- cimens in the USNM), information is needed from living or freshly killed material. Therefore, al- though I know of no exceptions within Mzcrocer- ella, or of any (homoplasic) occurrences of green eyes in other sarcophagine genera, I cannot fully assess the strength of this character state for defin- ing the genus. The majority of species of Mzcrocer- ella have a bare, pubescent or short plumose arista and males of almost all species have a dark brown or black protandrial segment (first genital segment of authors, syntergosternite 7-8 of McAlpine 1981), distinctly contrasting to a bright red or orangish epandrium. No species have their male terminalia entirely red, and the black/black condition found in M. muebni and in the probably monophyletic clade LM. hypopygialis + M. texana + M. valgata] is most probably derived from the black/red condition. The aedeagus is highly characteristic by the very rigid connection between basi- and distiphallus that probably allows little or no articulation. A part from a few species that have a highly modified aedeagus, the ventral surface of the distiphallus is equipped with a flat, sclerotized scale or plate. All species seen by me have the hypandrial arms more or less swollen at the level of the gonopods. The female terminalia show fused sternites 6-8 much like the condition seen in Spirobolomyia, Comasarcophaga, and many Blaesoxipha, and tergite 6 is entire and more or less extended beyond the posterior rim of tergite 5. Cryptosarcophila, with C. chaetosa as type spe- cies and only species included, was transferred to the Microcerellini by Lopes (1982b), but I prefer not to include this taxon in Microcerella. The rea- son for this is simply that I have not been able to recognize any convincing shared and assumed syn- apomorphic character state defining a clade con- taining only Cryptosarcophila and Microcerella. Al- though I have not examined any specimens of C. chaetosa myself, Lopes (1975e: 46) redescribed the species from the male holotype and noted that it has the "genital segments red”, which most probably is plesiomorphic relative to the red/black or black/ black condition in all Microcerella. Moreover, the arista is plumose and the aedeagus has no scalelike PAPE: American Sarcophaginae ventromedian plate or process. Cryptosarcophila chaetosa resembles many Microcerella in having the white postgenal setae restricted to the hypos- tomal bridge below the neck, which probably is a derived condition relative to the groundplan of the Sarcophaginae. Many other genera, however, seem to share this character state, e.g. Johnsonia, Neo- phyto and perhaps Notochaeta, and it does not provide evidence for inclusion particularly in Mz- crocerella. Little information is available on the biology of Microcerella, but species have been bred from land snails, beetles and grasshoppers, and in the labor- atory on horse blood serum (Blanchard 1966; Lopes 1969a, 1973). Microcerella is most diverse in the neotropical region and especially at high altitudes and in the southern temperate zones. It was therefore surpris- ing to find two specimens of an undescribed species of Microcerella from Bermuda, and even more so as the fauna of Sarcophagidae of these islands other- wise seems to consist entirely of a few widespread and possibly introduced species. Actually, the Dip- tera fauna of the Bermuda Islands is rather meager and with only few endemics. Many older Bermudan records of Sarcophagidae are misidentifications and only Helicobia morionella, H. rapax, Oxysarcodexia ventricosa and Ravinia lherminieri, all listed from Bermuda by Williams (1958, 1959), and all very common, widespread North American species, seem reliable (I have seen recently collected mater- ial of all but the latter species in USNM). Research on the Nearctic Microcerella, evoked by the discovery of the Bermudan species, revealed that what has been considered a single taxon, M. scrofa, actually covers two sibling species. Previous illustrations of terminalia in Aldrich (1916: fig. 13), Roback (1954: fig. 39) and Lopes (1982c: figs. 43- 47) seem to be of M. scrofa proper, while the species figured as Hypopelta scrofa in Hallock (1940: figs. 1-2) probably is its sibling M. adelphe (the surstylus figured by Hallock is perfectly scrofa-like, but this may be due to the rather simple outline provided by this author and the fact that the thornlike surstylar process is difficult to see in strict lateral view). Both new species of Microcerella are described below. Microcerella bermuda sp. n. (figs. 33-36) Type material. — Holotype @, "Bermudas | printed] vii 8 [handwritten, black ink] J.8 [handwritten, pencil] Davis [printed] (MCZ). Paratype, 18, "Bermuda Islands July 4 1905, T. K” [only “4” handwritten] (ZMUC). Both types bear a label reading "Sarcophaga n. sp. det. ALDRICH”. Description. Male. — Head: Narrowest part of frons 0.24X head width. Outer and inner vertical bristle well DIL TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 developed. Two weak ocellars present, the bristles much weaker than the pair of postocellars. Frontal bristles (7)-8; row continuing to just above level of apex of scape and distinctly curving laterally when seen from above. Fronto-orbital plate otherwise bare. Parafacial plate with a row of 4 setae in ventral part close to eye margin. Postcranium with poster- iormost postgenal setae and setae below occipital foramen white; other setae black. Scape distinctly raised above lunule, pedicel somewhat elongate, first flagellomere 1.5X as long as pedicel. Arista well developed, 2X as long as first flagellomere and short plumose. Palpus black. Thorax: Proanepisternum and postalar wall bare. Metasternum setose. Chaetotaxy: acr = 0 + 1, de = 4 (anteriormost 2 weak) + 3, ia = 2 (inner posthumeral + a weak presutural) + 2, sa = 2 + 3, pa = 2. Scutellum with 2-3 marginals (if 3 then the two posterior ones close together), one pair of weak discals, and one pair of apicals (slightly irregular in the holotype). Wing: Tegula black or brownish, basicosta yel- low, veins yellowish brown. Costal spine about 1.5X as long as crossvein r-m. Setal row of 14,5 just surpassing half the distance to r-m. Legs: Midfemoral apical pv bristles not spinelike (no midfemoral comb), hind tibia without any elon- gated pv or p setae. Abdomen: Ground colour black, microtomentum forming the usual silvery grey, tessellate and chan- ging pattern. Tergite 1+2-3 without median mar- ginal bristles, T4 with a weak pair, T5 with a com- plete row of marginals. ST3-4 with long setae with wavy or undulating tips. Terminalia: Protandrial segment dark reddish brown, with a few bristles on disc and a complete row of marginals. Epandrium bright red. Cercus and surstylus densely setose, cercus broad and plate- like when seen in posterior (dorsal) view, S-shaped in lateral view. ST9 (hypandrium) with a reduced disc but with arms greatly swollen at level of gonop- odal articulation. Gonopod shortened but rather broad at base, partly hidden by the swollen hypan- drial arm (lateral view), and with setae scattered along most of the distal margin. Aedeagus with a short basiphallus that is rigidly hinged to the dis- tiphallus. Distiphallus with a large transversely flattened plate carried on a narrow petiole from the ventral surface. Juxta present, well sclerotized and forming two arms slanting ventrally and basally. Lateral styli tubular, median stylus weakly sclero- tized and somewhat inflated, all styli projecting between juxtal arms. Length. — 10.0 mm. Female. — Unknown. Etymology. — A noun in apposition. Named for the type locality. 52 Distribution. — Nearctic: Bermuda Is. Note. — Johnson (1904) mentions a "Sarcophaga sp. ? A smaller form of which a number of specimens were collected by Mr. Davis, July 8”. These specimens may or may not include the holotype of the present species but obviously the note refers to the very collecting trip where it was taken (identical collector, locality and day + month) and settles with reasonable certainty the year when the holotype was collected as 1903, when Mr. C. Abott Davis visited the islands. Microcerella adelphe sp. n. (figs. 38, 40-43) Type material. — Holotype 6, USA: Maryland, Mont- gomery County, Rockville, 14.vi.1969, G. Steyskal (USNM). Paratypes, Canada: Ontario, Ottawa, 1d, 4- 7.viii.1982, L. Huggert (ZML); Quebec, Mount Tremb- lant, Lac aux Atocas, 18, 28.viii.1956, E.B. Thurman, "Bit- ing & Sweeping” (USNM).- USA: Georgia, 18, [no date], C. V. Riley [terminalia lost] (USNM); Minnesota, Ea- glesnest, 18, 29.vii.1958, W. V. Balduf (USNM); New York, Tompkins Co, [....] (illegible ?Ellio), 18, 25.viii. 1956, B. Foote, lab reared ex Polygura thyroides (ZMUC); New York, Tompkins Co, Varna, 16, 17.vii.1946, A. Stone (USNM): North Carolina, Great Smokie Nat. Park, Newfnd Ridge, 18, 11.vii.1941, A. L. Melander (USNM); Virginia, Great Falls, 16, 21.vi.1931, A. C. Melander, 19, "x.23,20”, J. M. Aldrich (both in USNM); West Virginia, Cranberry Gla., 1, 2.vi.1955, H. V. Weems (USNM). Description Male. — Very similar to M. scrofa. I have found only few diagnostic characters, all pertaining to the terminalia, and the description is accordingly res- tricted to these structures. Moreover, to facilitate a separation from M. scrofa, the description is given as one entry of a recognition couplet. For details of general morphology, the description of M. scrofa in Aldrich (1916) should be consulted. — Surstylus distally evenly rounded. Distiphallus broadest proximal to middle (lateral view) and with the appendages at the ventral surface rather slender (figs 5/159 144) ae ee ee WE RSA LAO Re En Microcerella scrofa — Surstylus distally with a concave incision and a thornlike process. Distiphallus broadest at or distal to middle (lateral view) and with the appendages at the ventral surface more com- PACS D STORES RARI One female has been included in the type series of M. adelphe although I cannot separate it mor- phologically from that of M. scrofa. Geographical evidence, however, suggests that M. adelphe may have a more easterly extended distribution relative to M. scrofa (see below). A female from Ohio (Am- herst) is accordingly judged to belong to M. scrofa. Etymology. — A noun in apposition. From the Greek adelphe = sister, the name refers to the probable sister group relationship with M. scrofa. Distribution of M. adelphe. — Nearctic: Canada (Ontario, Quebec), USA (Georgia, Maryland, Min- nesota, New York, Virginia, West Virginia). Biology. — The only information available is the breeding record from the snail Polygura thyroides (from label of one of the paratypes from New York State), and M. adelphe is probably predatory on snails, as many of its congeners. Microcerella adelphe may actually be the “undescribed species [which] parasitizes snails” mentioned by Downes (1965: 951). Nothing is known of the biology of M. scrofa. Distribution of M. scrofa. — Nearctic: Canada (Ontario [nr. Stratford]), USA (Alabama [Bir- mingham], Illinois [Algonquin], Indiana [Lafa- yette], Kansas [Topeka], Louisiana [Opelousas], Ohio [ Amherst], Texas [College Station], and Wis- consin [state record only]). The specimens on which this distribution is based are deposited in USNM (US records) and ZML (Canadian record). The apparent sympatry of M. adelphe and M. scrofa in at least part of their range (one male of each species from Canada: Ontario) is strong evi- dence that the rather slight differences reflect a reproductive barrier. The closest relative of M. adelphe is most prob- ably M. scrofa, as already mentioned. Both species possess a highly modified distiphallus with lateral sclerotized plates that extend posteriorly, thereby forming a deep, narrow groove along the posterior surface. Moreover, the gonopods are very character- istic by being greatly extended anteriorly, strongly curved upwards (or dorsally), and almost joining in the median plane (fig. 42). The group consisting of the sibling species adelphe + scrofa is probably the sister group to Microcerella bermuda. The character corroborating this hypothesis is the shape of the gonopod. In all three species, the gonopod has transformed from the elongate and more or less narrow structure found in all other Microcerella into a short or low, but very broad plate. The scat- tered setae that cover most of the distal margin may be used as a morphological landmark indicating the homologue of the posterior (or dorsal) margin of the plesiomorphic gonopod. The sister group of adelphe + scrofa + bermuda will probably be found among the Neotropical members of the group, as the three remaining Nearctic species of Microce- rella may have the Argentinean species M. muehni PAPE: American Sarcophaginae as their closest relative, these four species sharing the derived black/black condition of terminalia. Genus Spirobolomyia Spirobolomyta Townsend, 1917a: 43. Type species: Sarco- phaga singularis Aldrich, 1916, by original designa- tion. Diagnosis. — The genus Spirobolomyia may be defined by the following character states. 1) Male mid femur apically with a row of comb- like posteroventral spines; 2) Male abdominal sternite 5 with bristles along the posterior margin; 3) Male abdominal sternite 5 with a pair of bristly pads on the disc; 4) Male cercal prong bent backwards; 5) Male cercus with a sinuous lateral margin (pos- terior view); 6) Male cercus with short spinelike setae on the dorsal surface; 7) Basal parameral sclerite elongated (“additional forcipes” of Lopes 1975f); 8) Aedeagus with a beaklike projection arching over the heavily sclerotized juxta (fig. 48); 9) Aedeagal vesica beaklike and sclerotized; 10) Female abdominal sternites 6-8 fused; 11) Female abdominal tergite 6 with the median (= dorsal) part of the posterior margin devoid of setae and produced in a tongue-like projec- tion (Lopes 1975f: figs. 8, 17, 26). Townsend seldom explained the etymological derivation of his numerous generic names, but when he erected the genus Spirobolomyia with a type species that had been bred from a millipede, it seems probable that he, although he gave no men- tion of species other than the type species, included Sarcophaga flavipalpis as well, as this very similar species had been "reared ... from a myriapod” (Al- drich 1916: 258). Accordingly, he later stated that Spirobolomyia "Ranges in two species” (Townsend 1938: 68). Downes (1965) widened the concept by including Sarcophaga basalis and S. ohioensis in Spirobolomyia (as subgenus of Blaesoxipha), the biology of which is still unknown. It might be guessed that they are all parasitoids of the millipede genus Narceus. Note that I am here following Kee- ton (1960), who revised the Spirobolidae and argued that Spirobolus (consistently misspelled Sparobolus by Aldrich 1916: 186, 258) should be used for the group of Old World (China) species only, while Narceus should be applied to its New World sister group. As the name Narceus is the oldest genus-group name within the tribe Spirobo- lini, Narceus is the valid name in the present con- text whether or not Keeton’s infratribal classifica- tion is accepted. Species of Spirobolomyia are mainly distributed DO TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 in the eastern and southeastern parts of the Nearc- tic Region but extend along the east coast into the northern part of the neotropical region. The range extends from Canada: southern Quebec, (14 of S. flavipalpis in ZMUC) south to Mexico: Tamaulipas (18 of S. latissima in FSCA, see under type material below) and Chiapas (86 29 of S. ohioensis in CNC). The western limit of distribution seems to run through Wisconsin, Iowa, West Virginia, and Ala- bama (Downes 1965), and the Nearctic distribution is closely matched by the distribution of Narceus given in Keeton (1960). The life habit, although only known for two spe- cies and without any evidence that no other food source can be utilized, is interesting insofar as very few Diptera parasitize or prey upon millipedes. Within the Sarcophagidae, I know of only two other records: Specimens of Blaesoxipha beameri in USNM (44, USA: California, Riverside) bear a label reading "Ex Tylobolus sp. near hebes” prob- ably referring to a species of the spirobolid genus Hiltonius. Although this may refer to parasitism, predation or even scavenging, the food source ne- vertheless is a millipede. No other information is available on the biology of B. beameri. A single male specimen of Sarcophaga iulicida from Portugal (BMNH), with an affiliation in the subgenus Pzer- retia sensu stricto of Verves (1986), was bred from a specimen of Ommatoiulus moreleti (Pape 1990). Note that the parasitism of millipedes has not been used as a character state corroborating the mono- phyly of Spirobolomyia. Parasitism of a particular taxon is difficult to handle in a character analysis as parasites do not parasitize taxa as such, and al- though it seems reasonable that the peculiar habit of Spirobolomyia is evidence of phylogenetic rela- tionship I prefer to avoid an explicit scoring. Spirobolomyia was, like Fletcherimyia, included in Blaesoxipha as a subgenus by Downes (1965). However, all Spirobolomyia possess tubular styli that are free of each other and apparently func- tional, i.e. sperm conducting, and they do therefore not form part of the probably monophyletic group of species with only the median stylus tubular and with platelike lateral styli that are fused in the median plane. The latter group is the genus Blae- soxipha in the sense of the present author. The male midfemoral comb, the presence of short spines on the dorsal surface of the male cercal prong, the recurved male cercal prong, and the fused female abdominal sternites 6-8, however, are character states Spirobolomyia shares with Blae- soxipha. At least the latter four states seem derived with regard to the groundplan of the Sarcophagi- nae, and including Spzrobolomyia within Blaesoxi- pha (i.e. widening the concept of the latter) may be theoretically sound on the present state of know- ledge, and my reason for not doing this is, as given for Comasarcophaga and Fletcherimyia, that the 54 monophyly of the resulting taxon seems, at least to me, to be much more tentative than the monophyly of the groups included. A comprehensive phyloge- netic analysis of all sarcophagine genera is needed to settle this issue. Lopes (1975f: 156) simply dis- carded a relationship between Spirobolomyia and Blaesoxipha by stating that “there is no close rela- tionship between the two genera”, but he later (1988a: 130) elaborated on this postulate by refer- ring explicitly to the “long styli of glans” (= lateral styli of the present paper) being present in species of Spirobolomyia but absent in his Impariini (= Blaesoxipha (in part) of the present paper). The possession of functional lateral styli will corrobo- rate that Spirobolomyia does not form part of the clade here considered under the name Blaesoxipha (unless other evidence favour a reversal), but as the character state obviously is plesiomorphic at the level of the Sarcophaginae, it is totally uninforma- tive regarding the phylogenetic position of Spzro- bolomyia relative to Blaesoxipha. Species limits were discussed in detail by Lopes (1975f), but the number of specimens at his dispo- sal was rather limited and a more extensive mate- rial has revealed another, previously undescribed species. As only characters found in the male termi- nalia distinguish this species from S. ohioensis, and as species of Spirobolomyia in general are separable only by reference to their terminalia and sternite 5, I have restricted the description to include these structures only. Information on general appearance of Spirobolomyia can be found in Aldrich (1916), Hall (1927), Townsend (1938) and Lopes (1975f). Spirobolomyia latissima sp. n. (figs. 45-48, 50) Type material. — Holotype d, USA: Texas, Menard, 1929, E. O. Cushing (USNM). Paratypes, USA: Texas: 168, data as holotype (14 in USNM, 2 in ZMUC); 18 v.1930, otherwise as holotype (USNM); NW Blanco Co., Davis Ranch, 16, 22.iv.1959, J. F McAlpine (CNC); Rea- gan Wells, 18, 27.vi.1927, R. W. Laake (USNM); Sonora, 1d, 10. [month not interpreted].1920, O. G. Babcock (USNM); Hidalgo Co., 78, 29.iii.-9.x.1946, USPHS dys- entery fly trap (USNM); Hidalgo Co., 3&: 4.ii.1932, 14: 3.viii. 1934, H.J. Reinhard (3 in CNC, 1 in ZMUC); Brews- ter Co., Big Bend National Park, Basin, 14, 14.vi.1950, R. F. Smith (AMNH). - Mexico: Tamaulipas, 9 km W An- tiguo-Morelos, 18, 21.vii.1954, Univ. Kansas Exp. (ESCA). Description Male. — Abdomen: Sternite 5 deeply incised and with strong, spinelike bristles along the basal (or median) half of the posterior margin of each lobe. A pair of raised pads with strong bristles on the posterior, flattened surface are present on the disc. Terminalia: Cercus very broad in posterior view, with a sinuous outer (or lateral) margin and a greatly swollen flaplike extension basally on the cercal prong. A few short spines are present on the dorsal surface and along the margin of the hump- like lateral convexity. Cercal setae are arranged in two groups: a smaller one close to the median line of articulation between the cerci, and a more exten- sive one distal to this. Aedeagus with the dorsal (or posterior) surface raised into a narrow crest. Vesica well sclerotized and of about the same size as the beaklike structure arching over the juxta. Female. — Probably very like the female of S. ohioensis. Among nine females from Texas, Hi- dalgo Co., 14.iii.-23.x.1946 (USNM), all caught in dysentery fly traps, may be specimens of S. latis- sima. | have not been able, however, to find any differences between these and a female specimen of ohioensis pinned with a presumed conspecific male (USNM), and as the distribution of these two spe- cies seems to be largely sympatric, I have not in- cluded any females in the type series. Etymology. — A Latin adjective, from latissimus = the broadest. The name refers to the male cercus, which is very broad in posterior view. Distribution. — Nearctic: USA (Texas). Neo- tropical: Mexico (Tamaulipas). Females of Spirobolomyia are very similar and I cannot at present identify female specimens to spe- cies with any certainty. This implies a problem, as the holotype of Sarcophaga pallipes is a female. Aldrich (1930) examined this type and established its relationship, but he was not able to make a species level identification: "I can not separate [it] from females of simgularis and several others” (p. 19). Townsend (1938) and Roback (1954) consi- dered S. pallipes a senior synonym of S singularis, while Downes (1965) listed pallipes as “Probably Blaesoxipha (Spirobolomyia) sp”. Lopes (1975c) likewise examined the type of pallipes and syn- onymized it with Sarcophaga basalis, but his deci- sion obviously was tentative as he "was unable to [find] good characters to separate the female[s] of the Spirobolomyia with red legs” (p. 548). Appar- ently, his decision was based largely on the equal size of the primary types, which really does not provide especially convincing evidence as to their conspecificity. When Lopes (1975f) later produced a key to species of Spirobolomyia, he did not include the females and he did not examine any female specimens of S. ohioensis. For the present paper, I have accepted the synonymy of S. basalis with S. pallipes, awaiting more information on diagnostic female character states. Roback (1954) illustrated the aedeagus and ab- dominal sternite 5 of S. singularis (as pallipes), S. flavipalpis (as flavipes, error) and S. basalis. Lopes (1975f) gave a detailed description of these species PAPE: American Sarcophaginae as well as of S. ohioensis and provided illustrations of various parts of both male and female terminalia. Both authors should be consulted for additional information. The key of Lopes (1975f), however, is not easy to use and the posterior view of the male cerci is provided for two species only (that of S. basalis even at a slightly skewed angle). The poste- rior outline of the male cerci provides a very easy means of identification and has the advantage that this attribute often is visible even in specimens with only partly spread terminalia. For the present paper, I have provided illustrations of the posterior cercal outline for all species (figs. 49-53), and for completeness a couplet is provided to separate S. latissima from the very similar S. ohioensis: — Aedeagus with long vesica and a crest running along the dorsal surface (fig. 48). Cercus very broad in posterior view (figs. 46, 50). Setae of cercal base separated into two groups by an area with no or very few setae: A small group close to the suture between the cerci and a larger group along the outer (or lateral) margin (fig. AO sand eds bldg net se latissima — Aedeagus with shorter vesica (Lopes 1975f: fig. 35), dorsal surface distinctly keeled but not raised into a crest. Cercus narrower in posterior view (fig. 49). Setose part of cercal base without any interruption, setae forming a continuous cover from suture between the cerci and out along the outer (or lateral) margin.......... Genus Tripanurga Tripanurga Brauer & Bergenstamm, 1891: 367. Type spe- cies: Sarcophaga albicans Wiedemann, 1830, by desig- nation of Townsend (1916: 9). Thelylepticocnema Townsend, 1917a: 43. Type species: Sarcophaga incurva Aldrich, 1916, by original designa- tion. Glaucosarcophaga Townsend, 1917a: 45. Type species: Glaucosarcophaga knabi Townsend, 1917a [= Sarco- phaga albicans Wiedemann, 1830], by original desig- nation. Metoposarcophaga Townsend, 1917a: 46. Type species: Sarcophaga pachyprocta Parker, 1916 [junior primary homonym of pachyprocta Hagen, 1881; = a species of the Sarcophaga importuna Walker complex], by orig- inal designation. Syn. n. Zygastropyga Townsend, 1917b: 191. Type species: Zy- gastropyga aurea Townsend, 1917b, by original desig- nation. Cacotrophus Reinhard, 1947: 99. Type species: Cacotro- phus beameri Reinhard, 1947, by original designation. Erucophaga Reinhard, 1963a: 75. Type species: Eruco- phaga triloris Reinhard, 1963a, by original designa- tion. Syn. n. Diagnosis. — The genus Tripanurga may be de- fined by the following character states: 1) Aedeagal basiphallus with an epiphallus-like process; 2) Aedeagal basiphallus shaped as an elongated 55 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 and narrow tube, strongly contrasting to the compact distiphallus (T. aurea showing a se- condary thickening); 3) Aedeagal distiphallus compact and more or less globular; 4) Aedeagal distiphallus with a fringe of filiform processes at the anterior margin (Lopes 1978b: figs. 6-8; the present figs. 57, 61); 5) Parameral bristle slightly flattened; 6) Ejaculatory apodeme large; 7) Male cercal prong bent backwards. The groundplan of the probably monophyletic group Paramacronychiinae + Sarcophaginae is characterized by an almost completely reduced epi- phallus, but in all species of Tripanurga the prox- imal part of the basiphallus is extended into a dis- tinct process that is broad at the base, tapers gradually and slants backwards or posteriorly (Ro- back 1954: figs. 316-335; Lopes 1978b: fig. 6; She- well 1987: fig. 58c; the present figs. 57, 61). Shewell (1987) applied the term epiphallus to this structure, but I prefer to avoid it as the basiphallic process in Tripanurga cannot be considered homologous to the epiphallus of the Miltogrammatinae and Calli- phoridae but rather to its base. Character states 3-5 are here considered autapo- morphic, and probably state 2 as well as the condi- tion in T. aurea, which possesses a rather strong basiphallic tube (fig. 61), is judged to be secondary. The shape of the distiphallus may indicate a trans- formation with a gradual increase in the size of the dorsal humps: importuna-type — villipes-type > sulculata-type — aurea-type (compare Roback 1954: plates 23-24, and the present figs. 57, 61). The unique (i.e. autapomorphic) shape of the au- rea-type distiphallus makes a reversed polarity highly improbable. If this is accepted, T. aurea can- not be considered the sister group of all other Tri- panurga and the broad basiphallus found in this species has to be considered a secondary specializa- tion (i.e. apomorphic). The genus Tripanurga was described by Brauer & Bergenstamm (1891) to contain "1. albicans Wd. (Sarcophaga). 2. dimidiata Wd. 3. bicolor S.M.C. Bras” (p. 367). The latter name is apparently a nomen nudum while the second probably is Wiede- mann’s (1830) Sarcophaga dimidiata, which cur- rently is regarded as a species of Notochaeta, e.g. by Lopes (1969a). Two years later Brauer & Bergen- stamm (1893) included (or mentioned) albicans only, and Townsend (1916) designated this as the type species, probably without having seen the type or any conspecific specimens. Townsend spent six months of 1928 visiting European and American museums and examining a large number of types, and he later published the results in various papers. For Sarcophaga albicans, he gave the type deposit- ories as: “Male Ht in Wien and male Pt in Lima” 56 (Townsend 1931: 75), which would have been an effective lectotype designation if only a single syn- typic specimen had been present in Vienna (NMW). Townsend probably incorporated one of the original type specimens in his own collection, and when he later moved from Lima to Washing- ton, D.C., bringing his collection along, the para- type (which with the present lectotype designation automatically becomes a paralectotype) was depos- ited in the USNM (Townsend 1937b: 221). Aldrich went to Europe in 1929, i.e. after Townsend, but he published his results a year earlier, although with- out designating any of the specimens examined as lectotypes. He did, however, give an extensive des- cription of two males of Sarcophaga albicans in NMW, "Undoubted types” (Aldrich 1930: 8). As both these males are still present in NMW, and as Townsend’s visit in Vienna antedated that of Al- drich, Townsend must have incorporated one of three original syntypic males in his own collection and not one of the two seen by Aldrich as stated by Lopes (1978b). One of these two remaining speci- mens was then chosen by Townsend to be the hol- otype. This is to be considered as a valid lectotype designation only if the specimen in case can be properly recognized, but as Townsend neither la- belled one of the males in question (he never la- belled his ‘designated’ lectotypes) nor cited any in- formation that could refer to one particular specimen, I have simply regarded Townsend’s de- signation as invalid and chosen one of the two available specimens, which I have labelled as lecto- type. This male is in good condition except that it lacks right first flagellomere + arista, left foretar- sus, right midleg, left hind leg, right hind tarsus, and has the terminalia dissected and glued to the lowermost label. It is labelled Brasilien” (printed) and “albicans Wd J 48 Coll. Winthem” (last two words printed), and its identity as Tripanurga albi- cans is herewith verified. The two paralectotypes in USNM and NMW respectively have been labelled as such. Silvestri (1903) described the species Tripanurga termitophila, probably on a single female which he recovered and apparently bred from a termite nest. The holotype has never been revised [I have not tried to locate this specimen, which is from Santa Ana, Misiones, in Tucumàn Province, Argentina, and not just “America Meridionalis” as stated by Lopes (1969a: 47)] and Tripanurga has in practice remained monotypic since Brauer & Bergenstamm (1893) excluded (or omitted) Sarcophaga dimidiata. Some authors, however, have stressed the strong similarity to Metoposarcophaga and related taxa. Aldrich (1930) mentioned that Sarcophaga albicans resembled the type species of both Metoposarco- phaga and Zygastropyga, and Roback (1954: 81) mentioned that Tripanurga "will probably fall within the genus Metoposarcophaga’ as T. albicans seems to be “very close to the subgenus Zygastro- pyga”. Neither author, however, made an explicit synonymization. Downes (1965) employed a broad concept of Metoposarcophaga, although he did not include Erucophaga, but he probably did not con- sider Tripanurga at all as T. albicans has not yet been found north of Mexico. Lopes (1969a) listed Sabinata, Tripanurga and Zygastropyga as distinct genera, while Shewell (1987) separated both Thely- lepticocnema and Erucophaga from Metoposarco- phaga. I think that using Tripanurga in the more inclusive sense presented above should be preferred for the following two reasons: Monotypic genera as Tripanurga (with T. albicans only) and Erucophaga are avoided, and the monophyly of the resulting taxon is better corroborated. Accepting a mono- typic Tripanurga will leave Metoposarcophaga, e.g. sensu Downes (1965), without any assumed apo- morphies not shared by T. albicans, and further splitting with the creation of additional monotypic genera has to be accepted. The genus Erucophaga provides a somewhat different case as the straight or slightly anteriorly curved male cercus of tri/orzs is unique within Tripanurga (in the broad defini- tion) and may be plesiomorphic relative to the remaining species where the cercus is distinctly curved backwards. This would be an argument for treating this species as the sister group to all other Tripanurga, and it may, therefore, be an argument for maintaining the genus Erzcophaga. I still think, however, that this is unwarranted simply because the little new information presented (= the possi- ble monophyly of Tripanurga excl. triloris) is made up for by a similar loss (= the monophyly of Tri- panurga incl. triloris). Moreover, as no well corrob- orated sister group of Tripanurga incl. triloris has been proposed, the monophyly of Tripanurga excl. of triloris will be highly tentative, and one ends with nothing but another genus-group name to handle. Actually, if Argoravinia turns out to be the best candidate for the sister group, as discussed below, the cercal condition in treloris must be con- sidered as derived from the bent cercus of its con- geners. In conclusion, the monophyly of Tripanurga in the present, broadened sense is corroborated by some very convincing autapomorphic character states of the male copulatory apparatus, which furthermore makes the taxon easily recognizable, at least in the male sex. Some of the character states listed in the diagno- sis may be apomorphic at a higher (i.e. more inclu- sive) level. The large ejaculatory apodeme, for ex- ample (Hallock 1940: fig. 13; Lopes 1978b: fig. 6; Shewell 1987: fig. 58c), is likewise found in the genus Argoravinia (although not quite as large) and it may be taken as evidence for considering these two genera as sister groups. Hall (1933: 255) even stated that Argoravinia “appears greatly like the PAPE: American Sarcophaginae genus Metoposarcophaga”. Moreover, a rather broad male front (although with proclinate orbital bristles in albicans and triloris only) is likewise shared with Argoravinia, and in all species of Tri- panurga except T. triloris, the male cercus is bent backwards or posteriorly (figs. 54, 58), somewhat like the condition found in Argoravinia. The almost straight male cercus in T. triloris may then repres- ent a derived state. Tripanurga guatemala sp. n. (figs. 54-57) Type material. — Holotype 8, Guatemala: Antigua, El Salto, 1.v.[no year], J. M. Aldrich (USNM). Paratypes, Guatemala: Antigua, 14, 14.v.1923, E.G. Smyth (ZMUC); Guatemala City, 18, vii.1923, E. G. Smyth (USNM). Description Male. — Head: Narrowest part of frons 0.27- 0.29X head width. Outer and inner vertical bristle well developed, fronto-orbital plate densely setose and without proclinate orbitals. Uppermost frontal bristle strong and reclinate. Upper half of parafacial plate with some scattered setulae, lower part with a row of setulae close to eye margin. All postcranial setae, including those on hypostomal bridge, black. Head entirely silvery microtomentose. Antennal pedicel blackish with brownish base and apical mar- gin, first flagellomere blackish grey. Palpus black. Wing: Tegula black, basicosta yellow. Vein r, setose from base almost to the knob at level of subcostal bend/costal break, r,,; setose from base almost to crossvein r-m. Legs: All femora with long setae and without av or pv bristles. All tibiae, especially mid- and hind- with dense villosity. Abdomen: Tergite 1+2-3 without median margi- nal bristles, T4 with a median pair, and T5 with a complete row of marginals. Terminalia: Colour red. Cercal prong slender, prongs closely adjoining in the distal half leaving a small window at base. Surstylus broad, with a fringe of long flattened setae. Length. — 11.0-13.0 mm. Female. — Unknown. Etymology. — A noun in apposition. Named for the type locality. Remarks. — T. guatemala is most probably the sister species of T. villipes, the monophyly of this group being well corroborated by at least three distinctive character states that may be derived rel- ative to the groundplan of Tripanurga: 1) Postcranial setae all black; 2) Alllegs equipped with dense, long pilosity and no anteroventral or posteroventral bristles; 3) Strongly convex ventral margin of the sursty- DI TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 lus with a row of closely set, strong, and elon- gated setae. Tripanurga aurea (figs. 58-66) Distribution. — Nearctic: USA (Arizona, Cali- fornta), Mexico (Baja California, Sonora). Neotrop- ical: Mexico (Guerrero, Jalisco, Morelos), Nicara- gua. The reason for discussing the present species is that it seems to be split into two populations, a Baja Californian population and a mainland population, with some striking morphological differences be- tween the two. Indeed, these differences were thought to indicate a specific separation until I recovered specimens from northern Baja California showing intermediate character states and reticu- late character state combinations. Two characters showing especially pronounced differences, the out- line of the surstylus and the shape of the male abdominal sternite 5, are described in more detail below. The typical Baja Californian form (figs. 58-63) has a slight posteroventral concavity of the sursty- lar margin (best seen in lateral view as in fig. 58), and the lobes of the male abdominal sternite 5 are strongly swollen apically and transversely flattened posteriorly (fig. 62). Specimens of the mainland form have the surstylus deeply incised from the posteroventral margin (figs. 64, 65), and the lobes of abdominal sternite 5 are flat and evenly rounded (fig. 66), i.e. they represent unmodified lobate. ex- tensions of the sternal disc. The aedeagus is com- pletely identical in both populations (fig. 61), and no differences have been found in the female sex. It is interesting that the Baja Californian form does not seem to be just a peripheral isolate that has diverged from its plesiomorphic ancestral popula- tion on the mainland. Both populations (i.e. gene pools) have probably diverged from their common (hypothetical) ancestor by the acquisition of at least one autapomorphic character state, the Baja Cali- fornian form having developed the obviously apo- morphic swollen lobes of abdominal sternite 5 and the mainland form the incised surstylus. The two forms are morphologically homogeneous through- out their range, but in the area where they occur simultaneously (northern Baja California), inter- mediary specimens and specimens with transposed character state combinations, i.e. incised surstylus and swollen lobes of ST5, may be caught at the same locality. Therefore, reproductive isolation does not seem to have been established, and rather than proposing a formal subspecific name for the Baja Californian population, I prefer to use the term “form”, awaiting further studies of the popula- tion(s) showing intermediary conditions. 58 Material examined. Type material. — USA: Holotype 8, Arizona, Tempe, [no date], Webster (USNM). Other material. — Mexico: Sonora, Alamos, 4d, 20- 27.11.1963, P. H. Arnaud, Jr. (USNM); Jalisco, Labarca, 14 19, 3.x.1934 [no collector] (UNAM); Guerrero, Arcelia, 16, 21.x.1947, W. G. Downs (UNAM); Morelos, 5 km SW Amacuzac, 36, 8.vi.1979, J. Butze (2 in UNAM, 1 in ZMUC); Baja California Norte, Arr. Santo Domingo, 5.7 mi E Hamilton Ranch, dam site, 428 529, 23.iv.1963, H. B. Leech & P. H. Arnaud, Jr. (FSCA); Golfo California, Isla Angel de la Guarda, Puerto Refugio, 24, 6.11.1986, F. Arias (UNAM, ZMUC); Golfo California, Isla Espiritu Santo, 18 19, 11.1.1987, L. Cervantes (UNAM). - Nicaragua: Chinandega, 18, [no date], Baker (USNM). ACKNOWLEDGEMENTS I am most grateful to N. E. Woodley (USNM) for facilitating my access to the collections during a one-year fellowship at the Smithsonian Institution and for supplying important information on Ber- mudan Sarcophagidae. I thank W. L. Downes, Jr., Lansing, Michigan, for sharing much unpublished information on Nearctic Sarcophagidae, and K. Rognes, Hafrsfjord, Norway, for a sober and very professional critique. Thanks also to W. N. Mathis, F.C. Thompson, and N. E. Woodley (all USNM) for heuristic discussions of various aspects of generic definitions. I acknowledge the time spent by D. Grimaldi (AMNH), G. E. Shewell and B. E. Cooper (CNC), C. Vogt (MCZ) and H. V. Weems, Jr. (FSCA) who kindly introduced me to the collections of their respective institutions during short visits and made prompt loan arrangements. Other loans were kindly supplied from L. Fernandez (UNAM), A. O. Bachmann (MACN), and R. Danielsson (ZML). Mary E. Petersen (ZMUC) made valuable linguistic corrections. Grants from the Smithsonian Institution, the Danish Natural Science Research Council (grant no. 11-6152), and the CanaColl Foundation are gratefully acknowledged. REFERENCES Aldrich, J. M., 1916. Sarcophaga and allies. — La Fayette, Thomas Say Foundation; 302 pp., 16 pls. Aldrich, J. M., 1930. Notes on the types of American two- winged flies of the genus Sarcophaga and a few related forms, described by the early authors. - Proceedings of the United States National Museum 78(12): 1-39, 3 pls. Beaver, R. A., 1979. Biological studies of the fauna of pitcher plants Nepenthes in west Malaysia. - Annales de la Société Entomologique de France 15(1): 3-18. Blanchard, E. E., 1939. Los sarcofagidos argentinos. Con- tribuciön a su conocimiento. - Physis 17: 791-856. Blanchard, E. E., 1955. Nuevos sarcofagidos acridiófagos argentinos (Diptera). - Revista de la Sociedad entomo- lógica argentina 18(3-4): 27-34. 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Stephanostoma- tini to Moriniini. — Itaquaquecetuba, Sao Paulo. 309 pp. Verves, Yu. G., 1986. Family Sarcophagidae. - Pp. 58-193 in: Soós, A. & L. Papp, eds. Catalogue of Palaearctic Diptera. Vol. 12, Calliphoridae-Sarcophagidae. 265 pp. Weyenbergh, H., 1875. Animales utiles. Nemorea acridi- orum Weyenb. — Anales Agricultura de la Republica Argentina 3(9): 85-86. Wiedemann, C. R. W., 1830. Aussereuropäische zwei- flügelige Insekten. Vol. 2. - Hamm. 684 pp., 5 pls. Williams, R. W., 1958. A study of the summer filth fly population of the Bermuda Islands. - Journal of Para- sitology 44(3): 339-342. Williams, R. W., 1959. The reported biting and filth fre- quenting arthropods of the Bermuda Islands exclusive of the Ixodoidae and Araneida. - Proceedings of the Entomological Society of Washington 61(5): 234-238. Received: 15 November 1989 Revised version accepted: 12 April 1990 61 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 1-4. Comasarcophaga spp. Aedeagus plus right paramere + gonopod, lateral view. 1, C. longespinus. 2, C. prolepsis. 3, C. texana. 4, C. nexilis. Scale: 0.1 mm. 62 PAPE: American Sarcophaginae E 10 N. 11 Figs. 5-12. Comasarcophaga spp. Male cerci. Figs. 5-8, right cercus, lateral view. Figs. 9-12, cerci, posterior view. 5, 9, C. longespinus. 6, 10, C. prolepsis. 7, 11, C. texana. 8, 12, C. nexilis. Scale: 0.1 mm. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 55 GT LLC mn, mi 5: nt mme — N (a x < Ò) i ff 1m i DI Pd, at m rm In Ama, MON MIT Figs. 13-18. Fletcherimyia spp. Figs. 13-17, aedeagus plus right paramere + gonopod, lateral view. Fig. 18, distiphallus, ventral view. 13, F. abdita. 14, F. celarata. 15, F. fletcheri. 16, F. jonesi. 17, 18, F. rileyi. Abbreviations: J = juxta, LS = lateral styli, MS = median stylus, V = vesica. Scale: 0.2 mm. 64 PAPE: American Sarcophaginae Figs. 19-23. Fletcherimyia spp. Male abdominal sternite 5, as seen from below between sternite 4 and margins of tergite 5. 19, F. abdita. 20, F. celarata. 21, F. fletcheri. 22, F jonesi. 23, F. rileyi. Scale: 0.4 mm. 65 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 | j Figs. 24-28. Fletcherimyia spp. Outline of male cerci + right surstylus, posterior view. Dotted line on right cercus delimit area devoid of setae. 24, FE abdita. 25, F. jonesi. 26, F celarata. 27, F. fletcheri. 28, F. rileyi. Scale: 0.2 mm. 66 PAPE: American Sarcophaginae Figs. 29-32. Microcerella acridiorum. Male terminalia. 29, Aedeagus, lateral view, 30, Right paramere + gonopod, lateral view, 31, Outline of cerci. posterior view. 32, Outline of right cercus + surstylus, lateral view. Scale: 0.2 mm. 67 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 33-36. Microcerella bermuda. Male terminalia. 33, Right cercus + surstylus, lateral view. 34, Outline of cerci, posterior view. 35, Right paramere + gonopod, lateral view, plus posterior end of right hypandrial arm (hatched). 36, Aedeagus, lateral view. Abbreviations: J = juxta, LS = lateral styli, MS = median stylus. Scale: 0.1 mm. 68 PAPE: American Sarcophaginae Figs. 37-44. Microcerella spp. Male terminalia. Figs. 37, 39, 44, M. scrofa. Figs. 38, 40-43, M. adelphe. 37, right cercus + surstylus, lateral view. 38, right cercus + surstylus, lateral view. 39, right cercus (in part) + surstylus, posterior view. 40, right cercus (in part) + surstylus, posterior view. 41, paramere. 42, right gonopod (stippled) and posterior part of hypandrial arm, lateral view. Fig. 43, aedeagus. Fig. 44, distiphallus. Scale: 0.1 mm. 69 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 , Figs. 45-48. Spirobolomyia Latissima, male terminalia. 45, Abdominal sternite 5, ventral view. 46, Cerci, posterior view, setae of left cercus omitted. 47, Right paramere + gonopod, lateral view. 48, Aedeagus. Abbreviations: B = basal parameral sclerite, G = gonopod, J = juxta, MS = median stylus, P = paramere, V = vesica. Scale: 0.2 mm. 70 PAPE: American Sarcophaginae 50 Figs. 49-53. Spirobolomyia spp., outline of male cerci, posterior view. 49, S. ohioensis. 50, S. latissima. 51, S. basalis. 52, S. singularis. 53, S. flavipalpis. Scale: 0.2 mm. 71 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 54-57. Tripanurga guatemala, male terminalia. 54, Right cercus + surstylus. 55, Outline of cerci. posterior view. 56, Right paramere + gonopod, lateral view. 57, Aedeagus. Scale: 0.1 mm. 72 PAPE: American Sarcophaginae Figs. 58-61. Tripanurga aurea, male terminalia. 58, Left cercus + surstylus, lateral view. 59, Right paramere + gonopod, lateral view. 60, Outline of cerci, posterior view. 61, Aedeagus, lateral view. Scale: 0.1 mm. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 62-66. Tripanurga aurea, Details of male terminalia. Figs. 62, 63, Baja Californian form. Figs. 64-66, mainland form. 62, Lobes of abdominal sternite 5, posterior view. 63, Right surstylus, posterior view. 64, Distal part of right surstylus, posterior view. 65, Outline of left surstylus, lateral view. 66, Lobes of abdominal sternite 5, posterior view. Scale: 0.1 mm. 74 VIRGINIJUS SRUOGA Zoologijos katedra VPI, Vilnius SEVEN NEW SPECIES OF ELACHISTIDAE (LEPIDOPTERA) FROM THE USSR Sruoga, V., 1990. Seven new species of Elachistidae (Lepidoptera) from the USSR. Tijd- schrift voor Entomologie 133: 75-84, figs. 1-26. [ISSN 0040-7496]. Published 31 July 1990. Seven new species of Elachistidae are described from Tadzhikistan and Turkmeniya (Soviet Central Asia) and Primorskiy Kray (Soviet Far East). They belong to the genera Perittia Stainton, Elachista Treitschke, Biselachista Traugott-Olsen & Nielsen and Cosmiotes Cle- mens. The male external features and genitalia are described and figured in detail. The distribution is mapped. A historical review of studies on USSR Elachistidae is provided in the introduction. V. Sruoga, Zoologijos katedra VPI, g. Studentu 39, Vilnius 34, Lithuania. Keywords. — Elachistidae, new species, USSR, Central Asia, Primorskiy Kray. INTRODUCTION Elachistidae larvae are typically leafminers of herbaceous, monocotyledonous plants, belonging to the families Poaceae, Cyperaceae and Juncaceae. Some species are leafminers of herbaceous dicoty- ledons belonging to the Boraginaceae, Lamiaceae, Asteraceae and Cistaceae (Traugott-Olsen & Nielsen 1977). Only in a few cases Elachistidae larvae mine leaves of woody Caprifoliaceae (Lonicera). Elachistidae larvae form galleries or blotches between the upper and lower epidermis layers and feed on the green parenchyma. The adult moths are small, with a wingspan of 6 to 13 mm. The forewings of the majority of species are brown- ish-grey with one fascia or two fasciae. Some Ela- chistidae are unicolorous brown, white or cream. The family is widely distributed. However, revi- sions of Elachistidae are only published for the Nearctic fauna (Braun 1948) and the North Euro- pean fauna (Traugott-Olsen & Nielsen 1977). The last decade a considerable number of taxo- nomic papers has been published with descriptions of new species from western and central Europe, northern Africa, the Canary Islands, western and central Asia (Parenti 1978, 1981, Traugott-Olsen 1985a, 1985b, 1988, Nielsen & Traugott-Oben 1978b, 1981, 1987, Whitebread 1984) and also from Japan (Kuroko 1982, Parenti 1983). The Elachisti- dae fauna of the USSR has been poorly studied. In the Keys to the insects of the European part of the USSR” (Falkovitsh 1981) 77 species of Elachistidae have been treated, including figures of their geni- talia. This key was entirely based on Traugott-Olsen & Nielsen’s book (1977) on the Fennoscandian and Danish fauna, and did not provide original data on Elachistidae from the USSR territories. As early as the mid-19th century, five species of Elachistidae were recorded from Latvia, i.e. Elachista quadripunctella (Hübner), E. luticomella Zeller, E. pollinariella Zeller, E. albifrontella (Hübner) and E. argentella (Clerck) (Lienig 1846). Nine other spe- cies were added by Nolcken (1871), te. Elachista regificella Sitcom, E. humilis Zeller, E. pulchella (Haworth), E. bisulcella (Duponchel), E. cerusella (Hübner), E. gleichenella (Fabricius), Biselachista serricornis (Stainton), B. wtonella (Frey) and B. albidella (Nylander). Nolcken (1871) also recorded 16 species for the Estonian fauna. Twenty-four spe- cies were included in the Catalogue of Russian Le- pidoptera by Ershov & Fild (1870). Teich (1889, 1893, 1899) recorded five additional species of Ela- chistidae for the Baltic fauna: Elachista eleochariella Stainton, E. pullicomella Zeller, E. biatomella Stain- ton, E. dispilella Zeller and Cosmiotes freyerella (Hiibner). The Estonian entomologist Petersen (1924) mentioned 25 species for the Estonian fauna, four of which (Elachista apicipunctella Stainton, E. megerlella Stainton, E. poae Stainton and E. pome- rana Frey) were recorded for the first time for the eastern baltic region (the present Baltic republics). Some additional data on the Baltic Elachistidae were provided by Saar (1930), Palionis (1932), Brandt (1942), Prüffer (1947) and Kuusik (1962). Elachis- tidae from other eastern European regions are dealt with by Schille (1931): he noted 32 species from eastern Poland (now parts of the Ukraine). In recent years investigations on Elachistidae of various regions in the USSR (including the Baltic republics) have been intensified. Sulcs & Sulcs (1983, 1984, 1987), Savenkov (1984, 1987) and Ivinskis et al. (1985) recorded 48 species of Elachis- tidae for the three Baltic republics, and two more 75 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 species will be added shortly (N. Savenkov pers. comm.). From other regions of the European part of the USSR the studies in the Vyborg reserve near Leningrad are worth mentioning: 17 Elachistidae were recorded here (Sukhareva & Falkovitsh 1984). Further, Sinev (1988) found two species, Elachista nielswolffi Svensson and Cosmuotes exactella Her- rich-Schaffer, in the Murmansk region in 1979- 1980, amongst a considerable number of Lepidop- tera species. Material of the Crimean Elachistidae is still under study (Yu. Budashkin pers. comm.). Hitherto, only one work dealt with the Elachis- tidae of the Asiatic part of the USSR (Falkovitsh 1986), in which one new genus (Kuma Falkovitsh) and three new species (Kumia integra, Elachista ilicrina and E. manca) were described. From this review it is clear that the Elachistidae fauna of the USSR is very imperfectly known, and further studies are urgently needed for a better understanding of this rich and interesting fauna. This paper contributes with the description of seven new species from two regions in the Asiatic part of the USSR. MATERIAL AND METHODS Methods for preparation of genitalia largely fol- low Falkovitsh & Stekolnikov (1978). The genitalia of the Cosmiotes species are figured here with the valvae in situ, because of difficulties in spreading them without damage. The genitalia were studied with a Biolam microscope and a MBS-10 stereo- microscope, using the drawing apparatus according to Gorodkov (1961) for the line figures. The material studied has been collected during expeditions to largely unexplored and little known areas of Soviet Central Asia. In addition material from Tadzhikistan and Primorskiy Kray collected by R. Puplesis in 1982 to 1986 has been studied. Locality names are spelled in accordance with the Times Atlas of the World (comprehensive edn. 1975 and later). Fig. 1. Perittia biloba, holotype: male genitalia. Scale 0.1 mm. 76 The type specimens are deposited in the collec- tion of the "Minology” working group, Department of Zoology, Pedagogical Institute, Vilnius, Lithua- nia (MG) and in the collections of the Zoological Institute of the USSR Academy of Sciences in Le- ningrad (ZIAS). TAXONOMY Perittia biloba sp. n. (figs. 1, 25) Type material. — Holotype g: USSR, Tadzhikistan, 30 km N Dushanbe, env. Varzob (Kondara), 27.vi.1986, R. Puplesis (MG). Diagnosis. — Related to P. lonicerae (Zimmer- man & Bradley), distinguished from this species by the shape of the valvae and juxta, the basal part of the aedeagus and the shape of the tegumen. External characters. — Male (female unknown). Forewing length 3.0 mm. Frons, neck tufts and scape white, some brown scales. Antenna white and brown ringed. Labial palp slightly mottled by white and brownish scales. Thorax and tegulae strongly mottled by white and brown scales. Forewing, ground colour white, strongly mottled due to brown tipped scales; basal part of costa densely covered by dark brown scales; cilia grey; cilia line indistinct. Hindwing brownish; cilia light creamy grey. Male genitalia (fig. 1). — Uncus lobes small, with short setae. Gnathos more or less rounded when ventrally viewed. Tegumen widening basally. Valva short and broad, costa strongly sclerotized, narrow, convex, slightly setose, sacculus curved to angular, cucullus distinct but narrow, setose. Juxta U-shaped, lobes long, broad, distally with acute spine-like pro- cess; digitate process very small, slightly setose. Vinculum typical for the genus, weakly developed. Aedeagus strongly sclerotized, slightly tapering to distal end, two triangular lobes at base. Biology. Unknown. Holotype caught in June. Distribution (fig. 25). — Only known from the holotype, Tadzhikistan. Elachista fuscofrontella sp. n. (figs. 2-4, 26) Type material. — Holotype @: USSR, Primorskiy Kray, 10 km S Slavyanka (Ryazanovka), 5.viii.1983, R. Puplesis (MG). Diagnosis. — Closely related to E. albifrontella (Hiibner). Differs in having a darker frons and labial palp, aedeagus curved in basal part without cornuti, saccus gradually tapering, digitate process club-shaped, widest above the middle, juxta lobes not dilate. External characters. — Male (female unknown). Forewing length 3.0 mm. Frons, neck tuft greyish brown, silvery lustre. Antenna greyish and dark New species of Elachistidae SRUOGA: = “EZ \ bd \ À Sia \ AR, ef — \ GET RMS A (al 2a \ ZED 07 07 2882 = => ZE De ven“ ef — [| TE 7 © RES SE Go oS Sf DITES Ci — I 4 EK Ma eg F ~ GAS ern “=F 1 SA EREN È SE — 7 CZ DEA ©. FE / Gr “ % 2a 4 ever = e SE an => ?c = (17 MY 2 4 € x CH € 7 9 I cei cati 2 CEE e Figs. 2-4. Elachista fuscofrontella, holotype. 2-3, male genitalia ventral view; 4, male genitalia lateral view. Scale 0.1 mm. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 5-8. Elachista multipunctata, holotype, male genitalia. 5-6, ventral view; 7, lateral view; 8, aedoeagus. Scale 0.1 mm. 78 brown ringed, slightly serrate distally. Labial palp whitish from above, dark brown from below. Thorax and tegulae greyish brown, some silvery lustre. Forewing, ground colour dark grey, strongly mottled, scale tips almost black; medially inter- rupted, silvery, curved fascia before middle above fold; tornal and costal spots whitish, apical spot, shiny, silvery, not regular in shape; cilia brownish. Hindwing brown; cilia brownish. Male genitalia (figs. 2-4). — Uncus deeply in- dented, lobes large, setose. Gnathos rounded; costa of valva strongly sclerotized, short prominent hump. Juxta, lobes not apically dilate; digitate pro- cess club-shaped, very wide above middle. Vincu- lum, not long, without medial ridge; saccus gradu- ally narrowing. Aedeagus curved in basal part, gradually tapering, no cornuti. Biology. — Unknown. Holotype caught in Au- gust. Distribution (fig. 26). — Primorskiy Kray. Elachista multipunctata sp. n. (figs. 5-8, 25) Type material. — Holotype @: USSR, Tadzhikistan, 30 km N Dushanbe, env. Varzob (Kondara), 20.viii.1986, R. Puplesis (MG). Diagnosis. — Very closely related to E. maculata Parenti. Differs in having the juxta lobes and dig- itate process widening and the aedeagus with a large cornutus. Easily distinguished from related E. pollinariella Zeller by shape of the uncus and juxta. External characters. — Male (female unknown). Forewing length 3.4 mm. Frons, neck tufts white. Antenna whitish and creamy ringed, pecten white. Labial palp white from above, brownish from below. Thorax white; tegulae white, some brown scales. Forewings, ground colour white, mottled due to brown tipped scales, basal part of costa covered by dark brown scales; cilia white, distal part grey; cilia line more or less distinct by dark brown tipped scales. Hindwing brownish white; cilia greyish white. Male genitalia (figs. 5-8). — Uncus deeply in- dented, lobes strongly curved towards valvae. Gna- thos oval. Valva widest in the middle, costa with convex lobe in the middle. Juxta, lobes wide, apically tapering; digitate process short, wide, slightly se- tose. Vinculum short; saccus strongly curved. Aedea- gus strongly sclerotized, stout, sclerotized humps past the middle and apical, one large cornutus. Biology. — Unknown. Holotype caught in Au- gust. Distribution (fig. 26). — Tadzhikistan. Elachista megagnathos sp. n. (figs. 9-12, 26) Type material. — Holotype 4: USSR, Primorskiy Kray, 20 km E Ussuriysk (Gornotayeznoe), 6.vii.1982, R. Puple- SRUOGA: New species of Elachistidae sis (MG). Paratype: 1 4, same data, 7.vii.1982 (ZIAS). Diagnosis. — Closely related to E. gleichenella (Fabricius), easily distinguished from this species by the very large gnathos, shape of the valva and the basal extensions of the aedeagus. Uncus in contrast to E. gleichenella deeply indented, digitate process narrower. External characters. — Male (female unknown). Forewing length 2.6 mm (holotype), 2.9 mm (par- atype). Head grey-brown, shining silvery. Antenna brownish to brown, distal half slightly serrate. La- bial palp descending, whitish, shining from above, brownish from below. Thorax grey-brown, shining silvery. Forewing, ground colour blackish brown with bronze lustre, silvery marks with goldish lus- tre, basal spot reaching costa, medial fascia distinct, slightly widened in anal end, distal part of forewing with second, V-shaped fascia, cilia greyish brown. Hindwing brown; cilia greyish brown. Male genitalia (figs. 9-12). — Uncus deeply in- dented, lobes with small setae. Gnathos prominent, very large. Valva widest in middle, costa convex, cucullus setose. Juxta, apical part of lobes narrow- ing, triangular; digitate process typical for majority of the genus, broad, short with fine setae. Vinculum not long, more or less triangular. Aedeagus strongly sclerotized, characteristically bent just below distal end, basally extended into two lateral arms. Biology. — Unknown. Specimens were caught in July. Distribution (fig. 26). — Primorskiy Kray. Biselachista spinigera sp. n. (figs. 13-16, 25) Type material. — Holotype g: USSR, Turkmeniya, western Kopet Dag, 30 km E Kara-Kala (Juvan-Kala), 18.viii.1988, V. Sruoga (MG). Diagnosis. — Closest related to B. serricornis (Stainton). Distinguished by the long and distinct distal spine of the sacculus and the shape of the valvae and gnathos. External characters. — Male (female unknown). Forewing length 3.7 mm. Frons and neck tufts white. Antenna brownish-creamy, distal part of dis- tal half slightly serrate, pecten white. Labial palp white, basal part brownish from below. Thorax white; tegulae white, some brownish scales. Fore- wing, ground colour white, three brown spots, one behind the middle, near costal margin, other two, irregular in shape, in tornal and apical position; cilia white; cilia line absent. Hindwing brownish; cilia brownish. Male genitalia (figs. 13-16). — Uncus deeply indented, lobes slightly setose. Gnathos, lobes elon- gate. Tegumen basally widening. Valva, sacculus with long and distinct distal spine, costa strongly convex in the middle, cucullus widely rounded. 79 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 7 EN SS N SS ven, mere CA av < Figs. 17-24. Male genitalia. 17-19. Cosmiotes kopetdagica, holotype. 17, ventral view; 18, lateral view; 19, juxta, anellus and digitate process. 20-24. Cosmiotes pallens, holotype. 20, ventral view; 21, valva, lateral view; 22, uncus, gnathos and tegumen; 23, juxta, anellus and digitate process; 24, aedoeagus. Scale 0.1 mm. 82 Fig. 25. Distribution map: Perittia biloba (dot), Elachista multipunctata (dot), Biselachista spinigera (triangle), Cosmiotes kopetdagica (rectangle), Cosmiotes pallens (rectangle). Juxta, lobes broad, tapering dorso-lateral, small group of setae on apical margin; digitate process narrow at base, widest above the middle, setose. Vinculum triangular. Aedeagus more or less straight. Biology. — Unknown. Holotype caught in Au- gust. Distribution (fig. 25). — Turkmeniya. Cosmiotes kopetdagica sp. n. (figs. 17-19, 25) Type material. — Holotype g: USSR, Turkmeniya, western Kopet Dag, 40 km E Kara-Kala, 28.v.1988, V. Sruoga (MG). Diagnosis. — Closely related to C. amselt Parenti. Distinguished from this species by the shape of the juxta lobes and the very large gnathos. External characters. — Male (female unknown). Forewing length 3.6 mm. Frons and neck tufts greyish white, whitish lustre. Antenna whitish grey and brown ringed, distally slightly serrate. Labial palp white with some lustre from above, brownish from below. Thorax and tegulae brownish. Fore- wing, ground colour brownish grey, some silvery lustre, strongly mottled with brown scales, distal part mottled by dark brown scales, distinct white spots at tornus and costa, indistinct small whitish spots near costa and approximately in the middle of the forewing; cilia greyish; cilia line indistinct. Hindwing greyish brown; cilia grey. Male genitalia (figs. 17-19). — Gnathos very large, oval, broadened. Valva, distal spine of saccu- lus large, distinct, acute. Juxta, lobes short, triangu- lar in ventral view, small setae; digitate process long, distal part broad and setose. Vinculum trian- gular; saccus long, gradually tapering. Aedeagus long and slender, curved at base, long spine-like cornuti. Biology. — Unknown. Holotype caught in May. Distribution (fig. 25). — Turkmeniya. SRUOGA: New species of Elachistidae Cosmiotes pallens sp. n. (figs. 20-24, 25) Type material. — Holotype 4: USSR, Turkmeniya, western Kopet Dag, 40 km E Kara-Kala, 28.v.1988, V. Sruoga (MG). Diagnosis. — Most similar to C. freyerella (Hiibner). Easily distinguished from this and other related species by more or less pale forewings, broad gnathos and broad distal spine of sacculus, large anellus and presence of cornuti in the aedea- gus. External characters. — Male (female unknown). Forewing length 2.5 mm. Frons and neck tuft whit- ish creamy. Antenna creamy brownish, distal part finely serrate, scape brown, pecten creamy. Labial palp creamy white. Thorax whitish to creamy white, some yellowish-brown scales; tegulae brown. Forewing, ground colour whitish to creamy white, some yellowish-brown scales, brown scales in two irregular spots in middle and abundantly present in apical part of forewing, basal part of forewing paler, dominated by whitish-creamy scales; cilia greyish- creamy, some scattered brown scales; cilia line in- distinct. Hindwing of holotype lost. Male genitalia (figs. 20-24). — Gnathos broad. Valva, distal spine of sacculus very distinct and broad, not acute. Juxta, lobes short with fine setae, apical margin sharp; digitate process long, distally broad, setose; anellus very large. Vinculum with long, narrow saccus. Aedeagus long, slender, with acute cornuti. Biology. — Unknown. Holotype caught in May. Distribution (fig. 25). — Turkmeniya. Fig. 26. Distribution map: Elachista fuscofrontella (trian- gle), Elachista megagnathos (dot). ACKNOWLEDGEMENTS I wish to thank very much Dr. N. V. Savenkov (Riga) for valuable information on Elachistidae of Latvia. I am indebted to Dr. Rimantas Puplesis (Minology working group, Vilnius) for his advice 85 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 and help as well as for making available the mater- ial collected by him. Also I wish to express my gratitude to R. Noreika (Minology working group) and particularly to S. Jolanta, for their continuous and kind help with my work. References Brandt, FH., 1942. Lepidopterologische Sammelerinner- ungen aus dem gebiet der Ammat (Livland) mit einem Verzeichnis der dort gesammelten Arten. - Korres- pondenzblatt des Naturforscher-Vereins zu Riga 64: 244. Braun, A.F, 1948. Elachistidae of North America (Mi- crolepidoptera). - Memoirs of the American Entomo- logical Society 13: 1-110, 26 pls. Ershov, N. & A. Fild, 1870. Katolog cesuekrylyh Rossiys- koy imperii. - Trudy russkogo entomologiceskogo ob- shestva 4: 118. Falkovitsh, M.I., 1981. Semeistvo Elachistidae zlakovye moli-miniory (in Russian). - Opredelitel nasekomyh Evropeiskoi casti SSSR 4(2): 479-523. Falkovitsh, M.I., 1986. Cesuekrylye (Lepidoptera) ostan- covyh gor Kuldzuktau i podgornoj ravnini (jugo-za- padnij Kizilkum) (in Russian). In Fauna cesuekrylyh (Lepidoptera) SSSR. - Trudy vsesojuznego entomolo- giceskogo obvestva 67: 131-186. Falkovitsh, M.I. & A.A. Stekolnikov, 1978. Otriad Lepi- doptera-Cesuekrylye. Vvedenie (in Russian). - Opredelitel nasekomyh Evropeiskoi casti SSSR 4(1): 599: Gorodkov, K.B., 1961. Prosteyshiy microproyector dlya risovaniya nasekomyh (in Russian). - Entomologces- koe obozrenie 40: 936-939. Ivinskis, P., Pakalniskis, S. & Puplesis, R., 1985. - Augalus minuojantys vabzdZiai: 107-112. Mokslas, Vilnius. Kuroko, H., 1982. Elachistidae. - In Inoue, H., Sugi, S., Kuroko, H., Moriuti, S. & A. Kawabe. Moths of Japan 1: 259-260. Kadansha, Tokyo. Kuusik, A., 1962. Märkmeid Eesti Pisiliblikatest. - Fau- nastilisi Märkmeid I, 3. Loodu-Suurijate Selts Eesti NSV Teaduste Akademia Juures: 228-235. Lienig, F, 1846. Lepidopterologische Fauna von Livland und Kurland. - Isis 3/4: 299. Nielsen, E.S. & E. Traugott-Olsen, 1978a. Elachistidae (Lepidoptera) described by O. Staudinger, J. Mann and C. Mendes. - Entomologist's Gazette 29: 5-16. Nielsen, E.S. & E. Traugott-Olsen, 1978b. A reassessment of the genus Stephensia Stainton, 1858 (Lepidoptera, Elachistidae). - Entomologist’s Gazette 29(4): 183-200. Nielsen, E.S. & E. Traugott-Olsen, 1981. A new species and a new combination in the genus Stephensia Stain- ton, 1885 (Lepidoptera: Elachistidae). - Entomolo- gist’s Gazette 32: 245-250. Nielsen, E.S. & E. Traugott-Olsen, 1987. Four new west Palaearctic species of Elachistidae (Lepidoptera). - En- tomologist's Gazette 38: 103-116. Nolcken, J.H.W., 1871. Lepidopterologische Fauna von Estland, Livland und Kurland. - Arbeiten des Natur- forscher-Vereins zu Riga 4:1-392. Palionis, A., 1932. Idélis Lietuvos drugiy faunai pazinti. — Vytauto Didziojo Universiteto Matematikos-Gamtos fakulteto darbai 1931-1932 6: 181. Parenti, U., 1978. Nuove specie paleartiche del genere Elachista Treitschke (Lepidoptera, Elachistidae). - Bollettino del Museo di Zoologia dell’ Universita di Torino 4: 15-26. 84 Parenti, U., 1981. Nuove specie di Elachistidi paleartici (Lepidoptera: Elachistidae). - Bollettino del Museo di Zoologia dell’ Universita di Torino 4: 49-64. Parenti, U., 1983. Elachistidi del Giappone (Lepidoptera, Elachistidae). — Bollettino del Museo Regionale di Scienze Naturali Torino 1(1): 1-20. Petersen, W., 1924. Lepidopteren-Fauna von Estland (Esti) 2: 511. - Reval, Talinn. Priiffer, J., 1947. Studia nad motylami Wilenszczyzny. — Towarzystwo Naukove w Toruniu. Wydzial Matema- tyczno-Przyrodniczy: 278, 296. Saar, H., 1930. Insektenbeobachtungen in Livland, II. — Korrespondenzblatt des Naturforscher-Vereins zu Riga 60: 102-104. Savenkov, N., 1984. Nekotoriye danniye po faune i eko- logiyi cesuekrylykh Latvii (in Russian). - Latvijas En- tomologs 27: 60. Savenkov, N., 1987. Noviye danniye po faune i ekologiyi cesuekrylykh Latvii (in Russian). — Latvijas Entomol- ogs 30: 72. Schille, F, 1931. Fauna motyli Polski. - Prace monogra- ficzne Komisji fizjograficznej 2: 258-265. Sinev, S.Ju., 1988. K faune cesuekrylykh (Lepidoptera) Pecengskogo raiyona Murmanskoy oblasti (in Rus- sian). — Svyazi entomofaun severnoj Europy i Sibiri: 168-172. Sukhareva, I.L. & M.I. Falkovitsh, 1984. Cesuekryliye (Le- pidoptera) bolshogo Beriozovogo ostrova (in Rus- sian). — Materialy po faune Vyborgskogo zakaznika We VSSD. Sulcs, A. & I. Sulcs, 1983. Neue und wenig bekannte Arten der Lepidopteren-Fauna Letlands. 9. Mitteilung. - No- tulae Entomologicae 63:39. Sulcs, A. & I. Sulcs, 1984. Neue und wenig bekannte Arten der Lepidopteren-Fauna Letlands. 10. Mitteilung. — Notulae Entomologicae 64: 153. Sulcs, A. & I. Sulcs, 1987. Neue und wenig bekannte Arten der Lepidopteren-Fauna Letlands. 11. Mitteilung. — Notulae Entomologicae 67: 143. Teich, K., 1889. Baltische Lepidopteren-Fauna. Microlepi- doptera. — Arbeiten des Naturforscher-Vereins zu Riga N. F. 6: 125-126. Teich, K., 1893. Nachtrag zur baltischen Lepidopteren- Fauna. - Arbeiten des Naturforscher-Vereins zu Riga N. FE 6: 1-33. Teich, K., 1899. Vervollständigtes Verzeichnis der Schmet- terlinge der Baltischen Provinzen. - Korrespondenzb- latt des Naturforscher-Vereins zu Riga 42: 9-76. Traugott-Olsen, E., 1985a. Five new Elachista species from Sierra Nevada. - SHILAP Revista de lepidopte- rologica 13(49): 73-76. Traugott-Olsen, E., 1985b. Three new Elachista — species and supplement to the description of the five n. sp. from Sierra Nevada. - SHILAP Revista de lepidopte- rologica 13(49): 169-174. Traugott-Olsen, E., 1988. The Elachista triseriatella Stain- ton complex, with descriptions of eight new species (Lepidoptera: Elachistidae). - Entomologist’s Gazette DORIS Traugott-Olsen, E. & E.S. Nielsen, 1977. The Elachistidae (Lepidoptera) of Fennoscandia and Denmark. - Fauna entomologica scandinavica 6: 1-299. Whitebread, S.E., 1984. A new species of Perittia Stainton from Switzerland (Elachistidae). - Nota lepidoptero- logica 7: 271-281. Received: 5 April 1989. Revised version accepted: 1 February 1990. BR. THEOWALD & P. OOSTERBROEK Instituut voor Taxonomische Zoölogie UvA, Amsterdam ZUR ZOOGEOGRAPHIE DER WESTPALAARKTISCHEN TIPULIDEN. IX. DIE TIPULIDEN DES VORDEREN ORIENTS. Theowald, Br. & P. Oosterbroek, 1990. Zur Zoogeographie der Westpaläarktischen Tipu- liden. IX. Die Tipuliden des Vorderen Orients. — Tijdschrift voor Entomologie 133: 85- 95, figs. 1-3, tabs. 1-4 + 1 appendix [ISSN 0040-7496] Published 31 July 1990. A review is presented of the 198 species and subspecies of Tipulidae known from the Near East. The Tipulidae fauna of the Near East distinctly belongs to the Westpalaearctic but has an isolated position as is clear from the high amount of endemic species (67%). The Near East can be divided in two regions of different faunal composition, the Southeast with predominantly Mediterranean, and the Northwest with a majority of European and Eurasiatic (sub)species. About 50% of the (sub)species belongs to the subgenus Lunatipula. It is concluded that the differentiation within the larger species-groups of Lunatipula dates back to at least the Lower Miocene. Both authors: Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Universiteit van Amsterdam, Plantage Middenlaan 64, 1018 DH Amsterdam, Nederland. Key words. — Diptera; Tipulidae; Near East; Zoogeography. EINLEITUNG In dieser Arbeit sind die Tipuliden des Vorderen Orients zusammengefafst. Zum Vorderen Orient sind hier gezählt: die asiatische Türkei (Anatolien), die Sovjet Republiken südlich des Kaukasus (Trans- kaukasien: Georgien, Azerbaidjan, Armenien), Iran, Irak, Syrien, Libanon, Jordanien, Israel und Nordägypten (Fig. 1). Untersuchungen der Tipuli- denfauna dieser Gebiete haben erst spät angefan- gen. Bis 1960 waren insgesamt etwa 50 Arten und Unterarten bekannt, heute sind es 198. Damit ist aber die Tipulidenfauna des Vorderen Orients si- cher nicht vollständig bekannt. Manche neue Aus- beute enthält wieder von dort unbekannte Arten und von schon bekannten Arten neue Fundorte. Für diese Arbeit wurden Daten aus der Literatur und aus der Sammlung des Zoologischen Museums, Amsterdam, zusammengetragen. Der Grofteil der dort befindlichen Tipuliden des Vorderen Orients wurde gesammelt von A. & U. Aspöck, H. v. d. Brink, J. Lucas, H., T. & B. v. Oorschot, H. & R. Rausch, F. Ressl und H. Wiering. Wir sind ihnen und allen anderen Sammlern für das Material, das sie uns überließen, sehr dankbar. Besonders verplichtet sind wir unserem Freund Günther Theischinger (Engadine, Australien) für Ergänzungen und für das Überprüfen des deut- schen Textes. Für "Arten und Unterarten” ist im Text der Kürze wegen die Bezeichnung “Arten” verwendet. VEGETATION DES VORDEREN ORIENTS Ursprünglich war der Vordere Orient (Rikli 1946; Atlas of the Arab World and the Middle East 1960) größtenteils trockene Grassteppe und Wüste, Biotope, die kaum für Tipuliden geeignet sind. Es gab aber auch Wald und Wiesen: hauptsächlich ent- lang der Küsten, in den Gebirgen von Transkauka- sien, Iran und Ostanatolien und entlang der größeren Flüsse (z.B. Nil, Euphrat, Tigris) (Fig. 2). In den höheren Lagen der Gebirge gab es große Flächen von sommergrünem Laubwald und Misch- wald, in den niederen Lagen mehr immergrünen mediterranen Wald. Der Vordere Orient ist eines der ältesten Kultur- gebiete der Erde. Vor etwa 6000 Jahren wurde dort bereits mit Landwirtschaft angefangen, und diese hat sich seither über den größten Teil des Gebietes ausgebreitet. Vor 3500 Jahren gab es schon viele Städte. Eine so lange Zeit von Landwirtschaft und andere Beeinflussung der Vegetation durch menschliche Aktivitäten, wie zum Beispiel Holz- nutzung für Bau und Brennstoff, hatte einen star- ken Einfluß auf Flora und Fauna. Zuerst wurden die feuchten Gebiete entlang der Flüsse für Landwirt- 85 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Balkan - halbinsel È Levantinisches Becken Schwarzes Meer SudruBland \ Kaspisches À Turk- % Rers. Golf À Meer \ menien Fig. 1. Der Vordere Orient mit Hauptgebieten, Staten, tiirkischen Provinzen und Inseln. Von tiirkischen Provinzen markiert # sind keine Tipuliden bekannt. Hauptgebiet 1: Westanatolien Le Lesbos 7 Bursa 4 # Istanbul 8 Kocaeli 5 Canakkale 9 Sakarya 6 # Balikesir 10 Bilecik Hauptgebiet 2: Levantinisches Ktistengebiet Kh Khios Cy Cyprus Ik Ikaria 14 Izmir Sa Samos 15 Aydin Ko Kos 16 Mugla Ro Rodhos 19 Antalya Hauptgebiet 3: Zentralanatolien 22 # _ Eskisehir 28a Konya p. p. 27 Ankara 34 Kayseri Hauptgebiet 4: Ostanatolien 32 # Gaziantep 50 Mardin 46 Sivas 51# Diyarbakir 47 # Malatya 52# Elazig 48 Adiyaman 53# Tunceli 49 Urfa 54 # _ Erzincan Hauptgebiet 5: Nordanatolien 23 Bolu 39 # Corum 24 Zonguldak 40 Amasya 25 Kastamonu 41 Sinop 26 # Cankiri 42 # Samsun Hauptgebiet 6: Georgien Hauptgebiet 7: Azerbaidjan Hauptgebiet 8: Armenien Hauptgebiet 9: Iran 86 Kütahya Usak Manisa Denizli Icel Hatay Adana Maras Syrien Nigde Kirsehir Erzurum Bingöl Kars Agri Mus Tokat Ordu Giresun Gümüsane Burdur Isparta Afyon Konya p. p. Libanon Israel Agypten Nevsehir Yozgat Bitlis Van Siirt Hakkari Trabzon Rize Artvin schaft genutzt, und die ursprüngliche Vegetation und Fauna ist dort fast restlos verschwunden. In den feuchteren Gebieten der Gebirge wurden Laub- und Mischwälder großtenteils abgeholzt, und der Boden wurde über kürzere oder längere Zeit landwirt- schaftlich genutzt. Im ganzen Gebiet sind heute nur noch voneinander isolierte Laub- und Mischwald- reste übrig, besonders in den weniger zugänglichen Teilen der Gebirge. Viele der in älteren Zeiten für Landwirtschaft genutzten Gebiete sind heute ent- weder mit trockener mediterraner Hartlaubvegeta- tion bewachsen, oder zu Steppen und Wüsten ge- worden. Die Vegetation des Vorderen Orients schließt im Nordwesten und Norden nahe an jene der Balkan- halbinsel und Südrußlands an. Von der Balkanhalb- insel ist sie getrennt durch Bosporus und Dardanel- len, die aber stellenweise weniger als 5 km breit sind. Von Südrußland ist sie getrennt durch das Schwarze Meer und den Kaukasus. Dieses Gebirge ist bis über 5000 m hoch, fällt aber an den Küsten des Schwarzen Meeres und des Kaspischen Meeres bis ins Flachland ab. Im Nordosten ist der Vordere Orient vom sibirischen Gebiet getrennt durch das Kaspische Meer und die Steppen und Wüsten von Turkmenien. Die Gebirge von Nordostiran gehen aber allmählich in die von Afghanistan und Zen- tralasien über. Für die mehr oder weniger feuchtig- keitsliebenden Tipuliden ist somit im Norden des Vorderen Orients Austausch mit der Balkanhalbin- THEOWALD & OOSTERBROEK: Tipuliden sel, mit Südrußland und mit Zentralasien wohl schwierig, jedoch nicht auszuschließen. Im Süden ist der Vordere Orient für die feuchtigkeitslieben- den Tipuliden aber vollständig abgeschlossen vom orientalischen und vom afrotropischen Gebiet durch die Steppen und Wüsten von Südafghanistan, Südiran, Irak, Syrien, Jordanien und Arabien. Fau- nenaustausch mit diesen Regionen ist also nicht zu erwarten. ARTENLISTE (APPENDIX 1) In der Artenliste sind die 198 Tipuliden des Vor- deren Orients alphabetisch angeführt mit ihrer Verbreitung innerhalb und außerhalb des Gebietes. Aufgrund von Klima und Vegetation wurde der Vordere Orient in 9 Gebiete aufgeteilt, wobei aus praktischen Gründen soweit möglich die Grenzen von Staaten und Provinzen benutzt worden sind (Fig. 1). Die 9 Hauptgebiete sind: 1 Westanatolien, 2 das levantinische Küstengebiet, 3 Zentralanato- lien, 4 Ostanatolien, 5 Nordanatolien, 6 Georgien, 7 Azerbaidjan, 8 Armenien, 9 Iran. Im Text zu Figur 1 sind die 9 Gebiete mit zugehörigen Staaten, tür- kischen Provinzen und Inseln verzeichnet. Die große türkische Provinz Konya ist aufgeteilt in den feuchteren gebirgigen Westen und Süden (28) und die mehr trockene Hochebene (28a). Die Inseln wurden zum nächstgelegenen Festland gerechnet. Nicht mitgezählt wurden Jordanien und Irak, von Balkan - Schwarzes Meer halbinsel Levantinisches Becken SudruBland Turk - menien Kaspisches Meer = Pers. Golf Fig. 2. Die ursprüngliche Vegetation des Vorderen Orients, schwarz: feuchte Gebiete mit Wald und Wiesen, weiß: trockene Gebiete mit Steppen und Wüsten. 87 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 welchen Staaten keine Tipuliden bekannt sind. Von Syrien sind Tipuliden nur bekannt vom Küstenge- biet, von Iran fast nur von den Gebirgen südlich des Kaspischen Meeres. Von Ägypten ist nur eine Art bekannt, der Kulturschädling T. (Tipula) orientalis. In der Artenliste ist die Verbreitung über die 9 Hauptgebiete verzeichnet, und in Klammern, in welcher der türkischen Provinzen oder auf welchen Inseln. Ebenso sind die Verbreitungstypen im pa- läarktischen Raum aufgenommen. Es bedeutet: A Hauptverbreitung in Asien; E Hauptverbreitung in Europa; EA Hauptverbreitung in Eurasien; M Nur verbreitet im mediterranen Raum der Westpalaarktis. Die Endemiten sind mit e bezeichnet und dahin- ter steht jeweils einer der obengenannten Buchsta- ben. Die großgeschriebenen Buchstaben weisen auf die Verwandtschaft hin. So bedeutet zum Beispiel eA: die Art ist endemisch im Vorderen Orient, und ihre nächsten Verwandten haben asiatische Ver- breitung. ZOOGEOGRAPHISCHE BEMERKUNGEN Stellung des Vorderen orients im palaarktischen Raum In Tabelle 1 sind die Tipuliden des Vorderen Orients zusammengefaßt und geordnet nach ihrem Vorkommen in den 9 Hauptgebieten und nach ih- rer Verbreitung im paläarktischen Raum. Der Vordere Orient beherbergt 8 Arten mit asia- tischer Verbreitung oder Verwandtschaft (A und eA), 45 mit europäischer (E und eE) und 17 mit eurasischer (EA und eEA) Verbreitung oder Ver- wandtschaft. Die eurasischen Arten gehören zur westpalaarktischen Fauna, haben die Kaltzeiten am Ende des Pleistozins im Balkan Refugium ver- bracht und sich seither über Europa und über klei- nere oder größere Teile Asiens ausgebreitet (Theo- wald & Oosterbroek 1983, 1985). Tabelle 1. Die Tipuliden des Vorderen Orients nach ih- rem Vorkommen in den 9 Hauptgebieten. Hauptgebieter MERS 475 CR SN OT Verbreitungstyp A O7 21 REDA 2 3 e tot or Oel OO) 3 E GM 3 7 PWG 9 DANS eE NND PS2 1 EA 4 2 6 © 10, 7 5. 2 eEA 1 On it AL 2 3 5 M WUO 2e Foe Oor À SA eM 16,59 maw. Se 19S SAS 107, Total DI Bik WD 3b) Dik Gy) 410530578198 Insgesamt 128 Arten gehören zur mediterranen Fauna der Westpaläarktis (M und eM). Mit 190 zur westpaläarktischen Fauna gehörigen Arten, gegen- über nur 8 ostpaläarktischen, gehört der Vordere Orient deutlich zur Westpaläarktis, ganz im Gegen- satz zum borealen Europa, wo 64% der Tipuliden Arten von ostpaläarktischer Herkunft sind (Theo- wald & Oosterbroek 1985). Innerhalb der Westpaläarktis hat der Vordere Orient jedoch eine isolierte Lage, was sich im hohen Anteil der Endemiten (67%) und im hohen Anteil der mediterranen Arten (64%) zeigt. In jedem ein- zelnen der südlichen Gebiete Europas (Balkanhalb- insel, italienisches Festland, iberisches Festland) gibt es weniger als 40% Endemiten und weniger als 35% mediterrane Arten. Faunenzusammensetzung der 9 Hauptgebiete Es gibt große Unterschiede in der Zahl der Ar- ten, die von jedem einzelnen der 9 Hauptgebiete bekannt sind (Tabelle 1). Zentralanatolien (Hauptgebiet 3) zählt nur 15 Arten. Dieses Gebiet ist eine trockene Hochebene mit Steppen und Wüsten, zum Teil bewässert und landwirtschaflich genutzt. Was Tipuliden betrifft, ist es wahrscheinlich der ärmste Teil Anatoliens. Von Iran (Hauptgebiet 9) kennen wir 28 Arten. Von diesem Land sind aber relativ wenige Ausbeu- ten bekannt, und wahrscheinlich kommen dort mehr Arten vor und sind besonders noch ostpa- läarktische Arten zu erwarten. Es gibt keine deutlichen Hinweise, daß in einigen der weiteren Gebiete auffallend mehr, in anderen auffallend weniger gesammelt worden ist, obwohl Unterschiede in der Zahl der bekannten Arten deutlich sind. Wahrscheinlich hängen diese Unter- schiede mit dem Aufbau der Landschaft zusammen. Tabelle 2 gibt die prozentuelle Faunenzusam- mensetzung der Hauptgebiete. In dieser Tabelle sind die europäischen und eurasischen Tipuliden zusammengezählt. Aufgrund der Faunenzusam- mensetzung läßt sich der Vordere Orient in ein südwestliches Gebiet (Hauptgebiete 1 und 2) und ein nordöstliches (Hauptgebiete 4 bis 9) aufteilen. Im südwestlichen Gebiet gibt es vorwiegend medi- terrane Arten. Obwohl die europäischen und eura- sischen Arten nur 31% der Gesamtfauna darstel- len, sind sie im Nordosten in der Mehrheit. Prozentuell sind die Unterschiede mehr als deut- lich: im Südwesten 69-78% mediterrane und weni- ger als 30% europäische Arten, im Nordosten 48- 67% europäische und weniger als 27-43% mediter- rane Arten. Hauptgebiet 3, mit nur 15 Arten schließt sich mit 60% europäischer Arten dem Nordosten an. In Tabelle 3 sind die Tipuliden des Vorderen Orients eingeteilt in Arten mit südwestlicher Ver- breitung, Arten mit nordöstlicher Verbreitung und Arten, die im Südwesten und im Nordosten vor- Tabelle 2. Prozentuelle Faunenzusammensetzung der Tipuliden des Vorderen Orients in den 9 Hauptgebieten. Hauptgebiete 1 2 3 4 5 6 7 8 9 Total asiatische Arten De DO Ged Be WS I Tevet europäische und eurasische Arten 29 20 60 S1 58 58 48 67 54 31 mediterrane Arten 69 78 40 43 39 38 40 27 39 65 kommen. Zu Arten mit südwestlicher Verbreitung werden jene gezählt, die wenigstens in Westanato- lien und/oder dem levantinischen Kiistengebiet vorkommen. Einige von diesen reichen mehr oder weniger weit in den Westen von Nordanatolien oder in den Südwesten von Ostanatolien, sie fehlen aber in Transkaukasien und Iran. Zu Arten mit nordöstlicher Verbreitung sind jene gezählt, die in Transkaukasien und Iran Vorkommen. Zum Teil kommen sie auch im Osten von Nordanatolien und in größeren Teilen Ostanatoliens vor. Sie fehlen aber in Westanatolien und im levantinischen Kiistengebiet. Es gibt im Vorderen Orient von 198 Arten nur 19, die weitverbreitet sind. Sie sind ziemlich gleichmäßig über das ganze Gebiet verbreitet. Es fällt aber auf, daß 11 der 15 Arten von Zentralana- tolien zu dieser Gruppe gehören. In den anderen Gebieten sind sie deutlich in der Minderheit. Unter ihnen sind mehrere bedeutende Kulturschädlinge. Die übrigen 179 Arten sind ziemlich gleichmäßig auf Südwesten (93) und Nordosten (86) verteilt. Zwischen beiden Gebieten gibt es also nicht nur einen Unterschied im prozentuellen Anteil von eu- ropäischen und mediterranen Arten (Tabelle 2), jedes der beiden Gebiete zählt überdies mehr als 80% charakteristische Arten, die im anderen Ge- biet nicht vorkommen. Die beiden Gebiete haben weniger als 20% ihrer Arten gemeinsam. Der Vor- dere Orient läßt sich somit in zwei deutlich vonein- ander verschiedene Faunengebiete einteilen: Süd- westen und Nordosten. Ursache dafür ist wohl die Tatsache, daß beide Gebiete schon längere Zeit durch die Steppen und Wüsten von Zentralanato- lien voneinander isoliert sind. Austausch von Arten war nur möglich entlang der Küste des Schwarzen Meeres über Nordanatolien und vielleicht auch noch zwischen dem levantinischen Küstengebiet und Ostanatolien im Süden der asiatischen Türkei. Die europäischen und eurasischen Arten im Vor- deren Orient Die Mehrzahl der Arten mit europäischer oder eurasischer Verbreitung (E, EA) haben die Kaltzei- ten am Ende des Pleistozäns in einer Reihe von Kleinrefugien auf der Balkanhalbinsel verbracht THEOWALD & OOSTERBROEK: Tipuliden und sich seither über Europa oder Eurasien ausge- breitet (Theowald & Oosterbroek 1983, 1985). Von den insgesamt 40 nichtendemischen europäischen Arten im Vorderen Orient zeigen 31 eine ziemlich geschlossene Verbreitung von der Balkanhalbinsel über die Ukraine und Südrußland bis nach Georgien und/oder Azerbaidjan, und von dort reichen sie zum Teil weiter bis nach Iran oder in den Osten von Nord- und Ostanatolien, 7 reichen sogar bis nach Westanatolien und 4 bis in das levantinische Küs- tengebiet. Nur 4 der 40 nichtendemischen europäi- schen Arten sind von den Gebieten nördlich des Schwarzen Meeres und vom östlichen Faunenge- biet des Vorderen Orients unbekannt. Sie sind nur bekannt vom Westen des Vorderen Orients. Die Verbreitung von 5 Arten is unklar. Ähnliches finden wir unter den 22 Endemiten mit europäischer oder eurasischer Verwandtschaft (eE, eEA). Die meisten haben eine nächstver- wandte Art in den Gebieten nördlich des Schwarzen Meeres und auf der Balkanhalbinsel. Insgesamt 16 dieser Endemiten sind im östlichen Faunengebiet verbreitet, 2 davon bis Westanatolien. Rein westli- che Verbreitung haben nur 6 Arten. Die meisten europäischen und eurasischen Arten (47 aus 62) zeigen also mehr oder weniger deutlich eine Verbreitung von der Balkanhalbinsel über die Gebiete nördlich des Schwarzen Meeres bis in das östliche Faunengebiet des Vorderen Orients. Nur 15 Arten sind entweder nur vom westlichen Fau- nengebiet bekannt oder Verbreitung und/oder Ver- wandtschaft sind unklar. Die Gebiete nördlich des Schwarzen Meeres waren jedenfalls im Saalien (Rif) trockene Kältesteppen, und wenig weiter im Norden befand sich die Eisdecke des Ural. Der Osten des Vorderen Orients war damals also deut- lich von der Balkanhalbinsel isoliert. Es ist deshalb anzunehmen, daß die europäischen Tipuliden sich erst rezent, am Ende des Pleistozäns oder im Ho- lozän, von der Balkanhalbinsel über die Ukraine und Südrußland bis nach dem Osten des Vorderen Orients ausgebreitet haben. Ausbreitung vom Sü- den der Balkanhalbinsel über Bosporus und Darda- nellen bis nach Westanatolien hat anscheinend be- deutend weniger stattgefunden. Die mediterranen Arten im Vorderen Orient Von den mediterranen Arten des Vorderen Tabelle 3. Verteilung der südwest-, nordost- und weit- verbreiteten Arten über die 9 Hauptgebiete. kauptgebieten | DIESE CRA SOS Total Verbreitung südwest BO) Fl AO BO Or 0093 nordost OO OMS ECS 2 285162202286 I CRELO UL TLS NISSAN 815157357 517765740, 302287198 weitverbreitet Total 89 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Tabelle 4. Artengruppen von Lunatipula nach den in Figur 3 erwähnten Gruppen und der Zahl der endemischen Arten und Unterarten im mediterranen Gebiete (West Mediterr. inkl. Makronesien und Nordwest Afrika; Italien inkl. Korsika, Sardinien und Sizilien; Balkan südlich Österreich und Ungarn; Vorderer Orient wie für diese Arbeit umschrieben). Die Gezamtzahl der paläarktischen Lunatipula Arten und Unterarten ist 340. Mediterrane Gebiete West Italien | Balkan Vordere Paläarktische Mediterr. Orient Arten Falcatae falcata-Gruppe 9 3 14 helvola-Gruppe 1 3 5 bullata-Gruppe 4 5 10 10 — — Fasciculatae A livida-Gruppe 11 4 10 17 74 ill 4 10 17 (= 40%) Fasciculatae B verrucosa-Gruppe 5 6 caudispina-Gruppe 3 8 11 fascingulata-Gruppe 3 2 8 1 19 3 5 16 6 (= 20%) Spinosae A fascipennis-Gruppe 2 1 8 fulminis-Gruppe 1 3 truncata-Gruppe 18 22 — 2 20 Spinosae B1 caucasica-Gruppe 1 1 5 7 macroselene-Gruppe 1 1 4 19 30 peliostigma-Gruppe 2 2 12 19 1 4 7 35 (= 74%) Spinosae B2 pustulata-Gruppe 5 7 clio-Gruppe 5 1 6 phaidra-Grupe 4 l 6 acuminata-Gruppe 3 13 33 53 lunata-Gruppe 1 8 14 9 — 22 43 (= 58%) Orients sind 21 (M) unverändert auch von Gebieten außerhalb des Vorderen Orients bekannt und sind weiter 107 Endemiten (eM). Von den 21 Nichten- demen sind 11 verbreitet vom Westen des Vorderen Orients bis auf die Balkanhalbinsel (einige sogar bis in das westmediterrane Gebiet), gegenüber nur 4, die vom Osten des Vorderen Orients bis nach Siid- rußland oder weiter verbreitet sind. Das bedeutet, daß es mehr Austausch von mediterranen Arten gegeben hat im Westen als im Osten. Eine Art (T. (L.) cressa) kommt auf Rodhos und Kreta vor. Sie gehört in die phaidra-Gruppe mit kretischer Ver- breitung (siehe unten). Von 5 weiteren Arten ist die Verbreitung unklar. Sie sind entweder über das ganze ostmediterrane Gebiet, auch nördlich des Schwarzen Meeres, verbreitet, oder sie zeigen in ihrer Verbreitung große Disjunktionen. Von den 107 Endemiten haben 60 eine südwes- tliche, und 39 eine nordöstliche Verbreitung. Ob- 90 wohl die Arten also größenteils rein südwestlich oder nordöstlich sind, gibt es unter den Arten- gruppen, zu welchen sie gehören, kaum Hinweise auf eine derartige geografische Zweiteilung. Die größeren Artengruppen (/ivida-, macroselene-, pe- liostigma-, acuminata- und lunata-Gruppe) haben etwa gleichviele Arten im Westen und im Osten des Vorderen Orients. Wahrscheinlich waren diese Gruppen schon über den Vorderen Orient verbre- itet, bevor dieses Gebiet durch Austrocknung des zentralanatolischen Gebietes in zwei Faunenge- biete geteilt wurde. Die Untergattung Tipula (Lunatipula) im Vor- deren Orient Im Vorderen Orient gehören 124 der 198 Tipu- liden (63%) zur Untergattung Lunatipula. Die pa- läarktischen Arten dieser Untergattung wurden, hauptsächlich durch Mannheims (1963-1968) und Savtshenko (1964), eingeteilt in 27 Artengruppen, von denen 19 auch im Vorderen Orient heimisch sind. Die Verwandtschaftsbeziehungen zwischen den Artengruppen wurden durch die beiden Auto- ren aber nicht klargestellt. Eine Einteilung der paläarktischen Lunatipula- Arten in einige größere Hauptgruppen wurde schon von Riedel (1913) versucht, der ohne weitere Begründung die Untergattung aufteilte in Falcatae, Fasciculatae und Spinosae. Es ist möglich diese Ein- teilung teilweise durch Synapomorphien zu bele- gen, miteine weiteren Aufspaltung der Fasciculatae und Spinosae (Fig. 3). l. Adminiculum mit deutlichen membranösen Strukturen seitlich des Penisführers, die ihn größ- tenteils oder ganz verdecken. Plesiomorph sind derartige Strukturen nicht anwesend und liegt der Penisführer unbedeckt. 2. Seitlich des Penisführers finden sich nebst die differenzierte größere oder kleinere blattförmige Strukturen überdies kleinere oder größere Dornen. 3. Seitlich des Penisführers finden sich stark sklerotisierte Dornen und nur Reste von membra- nösen blattförmigen Strukturen. 4. Am Hinterrand des 8. Sternits der Männchen sind zwei Anhänge die meist kurze Dornen tragen. 5. Die Anhänge des 8. Sternits tragen einen lan- gen geschwungenen Dorn. 6. Zwischen den Anhängen des 8. Sternits befin- det sich ein Haarplättchen. FASCICULATAE A FASCICULATAE B SPINOSAE BI SPINOSAE B2 FALCATAE SPINOSAE A Fig. 3. Kladogramm der palaarktischen Hauptgruppen von Lunatipula. THEOWALD & OOSTERBROEK: Tipuliden Tabelle 4 gibt die Einteilung der 19 Artengrup- pen des Vorderen Orients unter den 6 Hauptgrup- pen von Figur 3, und die Zahl ihrer mediterranen Endemiten. Drei Arten der Balkanhalbinsel, die Mannheims in die falcata-Gruppe stellt, werden hier, conform mit Savtshenko, zu der fascingulata- Gruppe gezählt, wohin sie aufgrund des Baues des Adminiculum gehören. Aus Tabelle 4 geht hervor, daß den hochdifferen- zierten Spinosae Bl und B2 im Vorderen Orient die größte Bedeutung zukommt. Von den zugehörigen 121 mediterranen Endemiten dieser Gruppen ha- ben 78 (64%) dort ihre Verbreitung. Es ist anzu- nehmen, daß die Spinosae Bl und B2 sich im Vor- deren Orient differenziert haben, und daß sich später Arten von dort weiter ausgebreitet haben. Von den übrigen Hauptgruppen haben Fascicu- latae B und Spinosae A vorwiegend balkanische Verbreitung, sind die Fasciculatae A ziemlich gleichmäßig über die ganze Paläarktis verbreitet (besonders die /zvida-Gruppe) und findet sich die älteste Gruppe der Falcatae nur im westmediterra- nen Gebiet. Die Differenzierung der Hauptgruppen im ost- mediterranen Gebiet hat wahrscheinlich schon im Unteren Miozän oder früher angefangen. Eine so- genannte kretische Verbreitung über die südaegäi- schen Inseln zeigt sich in der bemacula-Subgruppe (livida-Gruppe, Fasciculatae A), der verrucosa- Gruppe (Fasciculatae B), der cretis-Subgruppe (macroselene-Gruppe, Spinosae Bl) und der phaidra-Gruppe (Spinosae B2) (Theowald & Oos- terbroek 1990). Eine derartige Verbreitung geht wahrscheinlich zurück auf eine Zeit, in der es von Südgriechenland über Kreta und Rhodos bis nach Westanatolien eine Landbrücke gab. Diese Land- brücke ist in der zweiten Hälfte des Miozäns zu Inseln auseinander gefallen (Parent 1988). Dies bedeutet, daß die meisten Hauptgruppen von Lunatipula, auch die hochdifferenzierten Spinosae B1 und B2, schon damals existierten. ZUSAMMENFASSUNG Unter den insgesamt 198 Tipuliden Arten im Vorderen Orient sind 8 ostpaläarktischen gegen- über 190 westpaläarktischen. Der Vordere Orient gehört somit klar zur Westpaläarktis. Charakteris- tisch für das Gebiet ist die große Zahl von Endemi- ten und mediterranen Arten. Der Vordere Orient läßt sich in zwei Faunenge- biete aufteilen: ein südwestliches und ein nordöst- liches. Beide haben nur wenige Arten gemeinsam. Sie sind voneinander getrennt durch die zentralana- tolischen Steppen und Wüsten. Fast ein Viertel der Arten des Vorderen Orients ist auch über die mehr nördlich gelegenen und küh- leren Gebiete Europas verbreitet. Sie finden sich vorwiegend im östlichen Faunengebiet des Vorde- Oi TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 ren Orients und es ist anzunehmen, daß die meisten europäischen und eurasischen Arten sich im Spät- pleistozän oder im Holozän von der Balkanhalbin- sel über Südrußland dorthin ausgebreitet haben. Fast die Hälfte der Arten des Vorderen Orients gehören zu hochdifferenzierten Artengruppen von Lunatipula, einer Untergattung von Tipula, die sich dort seit etwa dem Unteren Miozän entwickelt hat. LITERATURVERZEICHNIS Mannheims, B., 1963. 15. Tipulidae. — In: E. Lindner (Ed.): Die Fliegen der palaearktischen Region, 3 (5) 1, Lief. 238: 137-176. Mannheims, B., 1965. Idem, Lief. 256: 177-212. Mannheims, B., 1966. Idem, Lief. 267: 213-256. Mannheims, B., 1967. idem, Lief. 270: 257-288. Mannheims, B., 1968. Idem, Lief. 275: 289-320. Parent, G. H., 1988. Esquisse biogéographique de la Crete. — Linneana Belgica 9: 344-387. Riedel, M. P., 1913. Die paläarktischen Arten der Dipte- ren-Gattung Tipula L. — Abhandlungen des Vereins für naturwissenschaftliche Erforschung des Nieder- rheins, Krefeld 1: 1-122. Rikli, M., 1943. Das Pflanzenkleid der Mittelmeerländer. Erster Band. — H. Huber, Bern: 1-436. Rikli, M., 1946. Idem. Zweiter Band: 437-1094. Rikli, M., 1948. Idem. Dritter Band: 1095-1418. Savtshenko, E. N., 1964. Tipulidae. — Fauna SSSR (New Series), Diptera 2 (4) 89: 1-503. Theowald, Br. & P. Oosterbroek, 1980. Zur Zoogeogra- phie der westpaläarktischen Tipuliden. I. Die Tipuli- den von Nordafrika. — Beaufortia 30: 179-192. Theowald, Br. & P. Oosterbroek, 1981. Idem. II. Die Ti- puliden der iberischen Halbinsel. — Beaufortia 31: 1- 20. Theowald, Br. & P. Oosterbroek, 1982. idem. IV. The Tipulidae of Corsica and Sardinia with a note on Do- lichopeza fuscipes Bergroth. — Mitteilungen der schweizerischen entomologischen Gesellschaft 55: 317-332. Theowald, Br., C. Dufour & P. Oosterbroek, 1983. Idem. III. Die Tipuliden der europäischen Tiefebenen. — Bonner zoologische Beitrige 34: 371-394. Theowald, Br. & P. Oosterbroek, 1984. Idem. V. Die ita- lienischen Tipuliden. — Fragmenta entomologica Roma 17: 245-291. Theowald, Br. & P. Oosterbroek. 1985. Idem. VI. Die Tipuliden der montanen, alpinen und borealen Ge- biete. — Bonner zoologische Beiträge 36: 185-220. Theowald, Br. & P. Oosterbroek, 1986. Idem. VII. Die Tipuliden der Balkanhalbinsel. — Tijdschrift voor Entomologie 129: 1-13. Theowald, Br. & P. Oosterbroek, 1987. Idem. VIII. The Tipulidae of Israel and adjacent regions. — Israel Jour- nal of Entomology 20: 71-83. Theowald, Br. & P. Oosterbroek, 1990. Idem. X. The Tipulidae of the Eastmediterranean islands. — Bijdra- gen tot de Dierkunde (in press). IR APPENDIX 1: Artenliste Ctenophora 1. elegans Meigen, 1818: 2 (Sa)-E 2. guttata Meigen, 1818: 1 (28), 4 (67), 6-EA 3. magnifica Loew, 1869: 6, 9-eEA 4. ornata Meigen, 1818: 2 (29)-E Dictenidia 5. bimaculata (Linnaeus, 1761): 5 (58), 6, 9-EA Dolichopeza 6. fuscipes Bergroth, 1889: 2 (Sa)-M 7. graeca Mannheims, 1954: 4 (59), 5 (58)-M 8. hirsuticauda Savtshenko, 1968: 6, 7-eM 9. schabriari Theowald, 1978: 9-eM Nephrotoma cornicina-Gruppe 10. aculeata (Loew, 1871): 4 (64), 5 (40, 56), 6, 7-EA 11. appendiculata appendiculata (Pierre, 1919): 1 (21, 28), 3 (27), 4 (63, 64), 5 (25, 57), 6,7,8, 9-E 12. beckeri (Mannheims, 1951): 2 (Cy, 29, 30, Li, Is), 4 (50)-eM 13. cornicina (Linnaeus, 1758): 1 (20, 21, 28), 2 (Sa, Cy, Li, 15), G45 35) MOD SIMCA) 51(25 5418) CAS NOLEA 14. eugeniae (Savtshenko, 1957): 6-eEA 15. gwestfalica (Westhoff, 1880): 2 (14)-E 16. minuscula (Mannheims, 1951): 2 (Cy, 30, Sy, Li, Is)-eE 17. nasuta Oosterbroek, 1975: 1 (7, 28), 4 (64)-eM 18. quadrifaria quadrifaria (Meigen, 1804): 7-E 19. quadrifaria farsidica (Savtshenko, 1957): 9-eE 20. schaueffelei (Mannheims, 1964): 7, 9-eM 21. tenuipes (Riedel, 1910): 6-EA 22. theowaldi Oosterbroek, 1978: 1 (28), 2 (Ko, Ro, 14, 15, 19, 29)-eE crocata-Gruppe 23. analis (Schummel, 1833): 1 (21, 28), 3 (45, 37), 4 (59, 62, 63, 64), 5 (57), 6, 7, 8, 9-EA 24. croceiventris lindneri (Mannheims, 1951): 2 (Ko, 33), 3 (27; 34, 35), 4 (46, 61, 62, 63, 64, Op), Of), D (C2, D1), FSE 25. nox (Riedel, 1910): 1 (28), 3 (35), 4 (48, 50, 59, 60, 61, 64, 65, 66, 67), 7, 8, 9-eEA 26. pratensis (Linnaeus, 1758): 5 (56), 6, 8-E 27. scalaris scalaris (Meigen, 1818): 1 (7, 21), 2 (Ik, Ro, Cy, 14, 19, 30, Sy, Li, Is), 3 (34), 4 (50, 594035.64,63), SACS SC) OA SS I-ER 28 scalaris terminalis (Wiedemann, 1830): 5 (58), 6, 7, 8, 9-A Nigrotipula 29. nigra (Linnaeus, 1758): 7-EA Tipula T. (Acutipula) 30. cypriensis Vermoolen, 1983: 2 (Cy)-eM 31. fulvipennis fulvipennis De Geer, 1776: 4 (59), 5 (58), 6, 7, 9-EA 32. fulvipennis nigroantennata 1961: 5 (45, 58), 6, 7, 9-eEA 33. isparta Vermoolen, 1983: 1 (20), 2 (Cy, 29)-eM 34. irrequieta Alexander, 1935: 7, 8,9-A 35. libanica Vermoolen, 1983: 2 (Li, Is)-eE 36. macra Savtshenko, 1961: 7, 9-eM 37. transcaucasica 1961: 8, 9-M 38. transcaucasica latifurca Vermolen, 1983: 1 CAS) 2) dksSankKomlOns le Te) A (C2; 164; 65, 67), 5 (56, 57)-M Savtshenko, transcaucasica Savtshenko, T. (Beringotipula) 39. unca Wiedemann, 1817: 6, 7, 8-EA T. (Dendrotipula) 40. flavolineata Meigen, 1804: 6, 7, 8-E T. (Emodotipula) 41. saginata Bergroth, 1891: 4 (59), 8, 9-E T. (Lunatipula) acuminata-Gruppe 42. angela Mannheims, 1963: 2 (Li, Is)-eM 43. angelica Theowald, 1957: 2 (Sy, Li, Is)-eM 44. artemis artemis Theischinger, 1977: 2 (Kh, Ik, Sa)-M 45. artemis asiaeminoris Theischinger, 1982: 2 (14, 15)-eM 46. auriculata Mannheims, 1963: 2 (31)-eM 47. berytia Mannheims, 1963: 2 (Li, Is)-eM 48. borysthenica Savtshenko, 1952: 1 (28)-M 49. brinki Theischinger, 1987: 2 (19)-eM 50. canakkalensis Theischinger, 1987: 1 (5)-eM 51. curvata Theischinger, 1977: 2 (31)-eM 52. cypris Mannheims, 1963: 2 (Ro, Cy)-eM 55. decolor Mannheims, 1963: 1 (7, 9)-M 54. dedecor Loew, 1873: 2 (Sa, Ko)-eM 55. emmahelene Theischinger, 1980: 2 (29)-eM 56. franzressli Theischinger, 1982: 2 (16)-eM 57. geja Savtshenko, 1968: 6-eM 58. horsti Theischinger, 1982: 1 (18), 2 (19)-eM 59. huberti Theischinger, 1982: 2 (19)-eM 60. hyrcana Savtshenko, 1973: 7-eM 61. istriana Erhan & Theowald, 1961: 1 (Le, 9, 10, 18, 21, 28)-M 62. kaplani Theowald & Oosterbroek, 1987: 2 (Is)-eM 63. kreissli Theischinger, 1987: 5 (25)-eM 64. leeuweni Theischinger, 1982: 2 (Sy, Is)-eM 65. macropyga Savtshenko, 1952: 8, 9-eM 66. montifer montifer Theischinger, 1977: 2 (31)-eM THEOWALD & OOSTERBROEK: Tipuliden 67. montifer tasucuensis Theischinger, 1987: 2 (29)-eM 68. musensis Theischinger, 1987: 4 (63)-eM 69. osmana Mannheims, 1963: 1 (7), 5 (40), 6, 7-eM 70. parasimurg Savtshenko, 1968: 7-eM 71. peteri Theischinger, 1979: 1 (7)-eM 72. simurg Savtshenko, 1964: 7-eM 73. subacuminata Mannheims, 1963: 2 (29, 31, 33), 4 (60, 64)-eM 74. suleika Mannheims, 1963: 7, 9-eM 75. theowaldi Savtshenko, 1964: 6-eM 76. turca Mannheims, 1963: 1 (7)-eM 77. valerii Savtshenko, 1968: 6-eM 78. vermooleni Theischinger, 1987: 2 (Ko)-eM adusta-Gruppe 79. adusta lucistriata Mannheims & Savtshenko, 1967: 2 (31), 4 (61)-A caucasica-Gruppe 80. caucasica Riedel, 1920: 6-eM 81. kinzelbachi Theischinger, 1982: 2 (Sy)-eM 82. quadridentata paupera Savtshenko, 1964: 6-eM 83. talyshensis Savtshenko, 1964: 7-eM clio-Gruppe 84. chelifera Savtshenko, 1964: 4 (59), 5 (58), 6-eM fascingulata-Gruppe 85. eugeniana Simova-Tosic, 1972: 6-M 86. praecox Loew, 1873: 6-M 87. rbynchos Theischinger, 1977: 2 (14)-eM fascipennis-Gruppe 88. fascipennis Meigen, 1818: 6-E helvola-Gruppe 89. helvola Loew, 1873: 1 (Le, 7, 28), 2 (Sa, 16, 19, Is), 5 (56)-E laetabilis-Gruppe 90. humilis Staeger, 1840: 6-E livida-Gruppe 91. adzharolivida Savtshenko, 1968: 6-eM 92. bimacula bimacula Theowald, 1980: 1 (Le), ?9-M 93. cinerella Pierre, 1919: 1 (20, 28), 2 (19)-M 94. deserticola Savtshenko, 1968: 7-eM 95. freidbergi Theowald & Oosterbroek, 1987: 2 (Is)-eM 96. kumerloevi Mannheims, 1968: 2 (31)-eM 97. kybele kybele Mannheims, 1968: 1 (28), 2 (29, Li, Is), 3 (35), 4 (63, 64, 66, 67)-eM 98. kybele russula Theischinger, 1977: 1 (18, 20, 28), 2 (19)-eM 93 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 99. livida morosa Savtshenko, 1964: 6-eE 100. maija Savtshenko, 1973: 6-eM 101. mendli Martinovsky, 1976: 1 (Le, 7, 17, 21), 2 (Kh, Ko, Sa, 16)-eM 102. nigdeensis Bischof, 1905: 3 (28a, 35)-eM 103. pseudowolfi Theischinger, 1979: 1 (28), 2 (Sa, 16)-eM 104. rbodolivida Theowald, 1972: 2 (Ko, Ro, 16, 19)-eM 105. saltatrix Savtshenko, 1964: 6-eM 106. sternalis Theischinger, 1977: 2 (Is), 4 (64)-eM 107. trapeza Theischinger, 1982: 1 (21)-eM 108. turcolivida Mannheims, 1968: 2 (31)-eM 109. vulpecula Theischinger, 1979: 4 (65), 5 (56)-eM lunata-Gruppe 110. furcula Mannheims, 1964: 1 (5, 7, 21, 28), 2 (Kh, Ko, Ro, 14, 15, 16)-M 111. Aybrida hybrida Savtshenko, 1952: 7-eM 112. hybrida altivolans Savtshenko, 1971: 6-eM 113. lunata Linnaeus, 1758: 6, 7, 8-E 114. paravelox Theischinger, 1987: 2 (33)-eM 115. pseudolunata pseudolunata Theischinger, 1980: 1 (21, 28)-eM 116. pseudolunata spinalonga Theischinger, 1982: 6-eM 117. soost soost Mannheims, 1954: 1 (Le, 5, 7, 18, 2028) Sa, Ro 1415416419293), 3 (34, 35), 6-M 118. soosz izmirensis Theischinger, 1987: 2 (14)- eM 119. subvelox Savtshenko, 1968: 5 (25), 6-eM 120. wmicornis Theischinger, 1977: 5 (25, 43)-eM macroselene-Gruppe 121. acudens Theischinger, 1977: 2 (33)-eM 122. adapazariensis Theischinger, 1987: 1 (8)-eM 123. aurita Riedel, 1920: 6, 7, 8-eM 124. carens Theischinger, 1987: 1 (8)-eM 125. cedrophila Mannheims, 1963: 1 (10), 2 (Ro, Li, Is)-eM 126. christophi Theischinger, 1982: 2 (Sa, 14)-eM 127. dumetorum Savtshenko, 1964: 6-eM 128. holzschuhi Theischinger, 1977: 9-eM 129. zliensis Mannheims, 1965: 2 (31)-eM 130. zmbecilla Loew, 1869: 2 (Ro)-eM 131. kerkis Theischinger, 1977: 1 (21)-eM 132. lithophila Savtshenko, 1968: 7-eM 133. /ucasi Theischinger, 1987: 4 (67)-eM 134. oorschotorum Theischinger, 1987: 1 (21)-eM 135. palifera Mannheims, 1965: 2 (19, 33), 3 (35)- eM 136. rutila Savtshenko, 1952: 2 (33), 8-eM 137. selenis Loew, 1873: 2 (Ro)-eM 138. subbispina Savtshenko, 1952: 9-eM 139. tibonella Theischinger, 1977: 1 (Le, 11), 2 (14, 19)-M 94 140. zaitzevi Savtshenko, 1952: 6, 8-eM mellea-Gruppe 141. affinis Schummel, 1833: 6-E peliostigma-Gruppe 142. antilope Theischinger, 1977: 4 (67), 9-eM 143. astigma Savtshenko, 1968: 6-eM 144. didymotes Theischinger, 1977: 4 (61, 64, 65, 67)-eM 145. micropeliostigma Mannheims, 1965: 1 (21)- eM 146. ornithogona Theischinger, 1982: 2 (16)-eM 147. peliostigma peliostigma Schummel, 1833: 1 (US) 2A (Sa xls ID) ZSNSTI NOMAD E 148. peliostigma burdurafyonensis Theischinger, 1987: 1 (18, 21)-eE 149. pseudopeltostigma Mannheims, 1965: 2 (Li, Is)-eM 150. renate Theischinger, 1982: 1 (18)-eM 151. selene Meigen, 1830: 6-E 152. semipeliostigma Mannheims, 1965: 2 (31)-eM 153. strigosa Savtshenko, 1952: 5 (58), 6-eM 154. ulrike Theischinger, 1982: 1 (28)-eM 155. wrswlae Mannheims, 1965: 2 (19)-eM phaidra-Gruppe 156. cressa Mannheims, 1965: 2 (Ro)-M 157. sciurus Theischinger, 1977: 1 (20, 28), 2 (19, 31)-eM truncata-Gruppe 158. arnoldu Savtshenko, 1957: 6-M verrucosa-Gruppe (= brunneinervis-Gruppe) 159. dracula Theischinger, 1977: 1 (21), 2 (Kh, Ko, 16)-eM 160. neutra Theischinger, 1982: 1 (28), 2 (19)-eM 161. quinquespinis Theischinger, 1980: 1 (Le, 17, 18, 28), 2 (16, 19, 29)-eM 162. teunisseni Theischinger, 1979: 2 (Ro)-eM 163. verrucosa verrucosa Pierre, 1919: 6, 7, 8-M 164. verrucosa sinedente Theischinger, 1980: 1 CUS A 23) AGO) 3) (AID), A (EO), SIE 43, 55)-eM zimini-Gruppe 165. zimini semiopaca Savtshenko, 1964: 7-A T. (Mediotipula) 166. anatoliensis Theowald, 1978: 1 (28)-eE 167. caucasiensis Theowald, 1978: 6-eE 168. stigmatella Schummel, 1833: 1 (7), 8-E T. (Pterelachisus) 169. luteobasalis luteobasalis Savtshenko, 1964: 4 (64), 5 (58), 6, 8, 9-eE 170. luteobasalis alata Theowald, 1980: 4 (64)-eE IA trichopleura Savtshenko, 1964: 6-eE T. (Savtshenkia) 172. 17,5 174. 75 176. 7/7. 178. 179: 180. 181. baltistanica Alexander, 1935: 6, 7-A interserta Riedel, 1913: 6, 7, 8-E jeekeli Mannheims & Theowald, 1959: 1 (Le), 2 (Ro)-M nivalis Savtshenko, 1961: 6-eEA obsoleta Meigen, 1818: 5 (56), 6-E ornata Theowald & Oosterbroek, 1987: 2 (Is)-eE pechlaneri Mannheims & Theowald, 1959: 9-E rufina rufina Meigen, 1818: 2 (Cy, Li, Is), 4 (67), 7, 9-E serrulifera Alexander, 1942: 5 (58), 6-E subalpium Savtshenko, 1961: 6-eE T. (Schummelia) 182. 183. ahrenst Savtshenko, 1957: 6-eE varicornis Schummel, 1833: 8-E T. (Tipula) 184. 185. 186. italica errans Theowald, 1984: 1 (Le), 5 (56), 7, 8, 9-M oleracea Linnaeus, 1758: 2 (Ro)-E orientalis Lackschewitz, 1930: 1 (Le, 5, 18, 20, DIN SNN NRO el 4 15129 Sv Is, Ag), 3 (34), 4 (46, 48, 59, 63, 64, 65), 5 (41, 44, 58), 6, 7, 8, 9-M 187. THEOWALD & OOSTERBROEK: Tipuliden subcunctans Alexander, 1921: 8-EA T. (Vestiplex) 188. 189. 190. LOE 192: kosswigi Mannheims, 1953: 1 (7)-eA pallidicosta pullata Savtshenko, 1960: 1 (28), 4 (61), 5 (56), 6, 7, 8-eE relicta Dia & Theowald, 1982: 2 (Li, Is)-eA semivittata semivittata Savtshenko, 1960: 5 (45, 58), 6, 7-E semivittata dissimilis Savtshenko, 1964: 6-eE T. (Yamatotipula) 193. 194. 195. 196. 197. 198. caesia Schummel, 1833: 1 (17, 21, 28), 3 (27), 4 (59, 63, 64), 6-E caucasimontana Savtshenko, 1955: 4 (67), 7-eA lateralis lateralis Meigen, 1818: 1 (7, 17, 18, ZO W228) 82 (Sato Ko Ro 942993933) Sy, Li, Is), 3 (27), 4 (46, 49, 50, 59, 60, 64, 67), 5 (24, 56), 6, 8-E lateralis tranensis Theowald, 1978: 4 (64), 9- eE pruinosa Wiedemann, 1817: 6-EA submontium Theowald & Oosterbroek, 1981: INNEN IES TA ICE 8-E Received: 23 March 1990 Revised version accepted: 16 May 1990 95 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 BOOK REVIEW Johansson, R., E. S., Nielsen, E. J. van Nieukerken and B. Gustafsson, 1989. The Nepticulidae and Opostegidae (Lepidoptera) of North West Europe. - Fauna Ento- mologica Scandinavica 23: 1-739, text-figs. 1-111, ta- bles 1-3, figs. 1-1122. [ISBN 90 04 08698 6 (2 parts), E. J. Brill, Leiden, The Netherlands. Price 220 Dutch Guilders/US $ 110.-]. For the first time an up to date treatment is given for all the 121 species of Nepticulidae and four species of Opostegidae occurring in Fennoscandia, Denmark, the British Isles, The Netherlands, Bel- gium, France north of Paris, West and East Ger- many, Poland and the Baltic states. This two vo- lume work figures in full colour all species and gives additional colour illustrations in cases of sexual dimorphism or polymorphism. The colour illustra- tions are of very high quality and make the book even more a must for specialists, lepidopterists in general and book-lovers alike. Line drawings of male and female genitalia and leaf-mines for all species are given. Also the larvae of most species are figured. Identification of the species is facilitated by var- ious keys: key to the tribes and families of Nepti- culoidea; key to the mines and larvae, arranged per host plant genus; keys to the males, based on exter- nal characters; keys to the males, based on genitalia; keys to the females, based on external characters and keys to the females, based on genitalia. Unfor- tunately the key for the females of Nepticulini (the first 76 species) based on the genitalia would be too much a copy of the full descriptions and has been omitted. But upon checking the line drawings of the female genitalia a definite identification is easily reached. Each species is fully redescribed and biology and distribution given. The chapters on immature stages, biology, collecting and rearing, will undoubt- edly stimulate many people to keep an eye out for the mines of the Nepticulidae. The phylogeny, classification and nomenclature are briefly, but adequately, dealt with in a separate chapter. One minor point to be mentioned in the section which deals with the Nepticulini is the lack- ing of the flight periods for the adults and the months in which the larvae can be found. This in contrary to the other part of the book where this information is given. This book has a very high scientific standard, but is easily accessible for the non-specialist at the same time. [R.T.A. Schouten] 96 J. vAN TOL National Museum of Natural History, Leiden KEY TO THE MALESIAN SPECIES OF LEPTOGOMPHUS SELYS, WITH THE DESCRIPTION OF A NEW SPECIES FROM SABAH (ODONATA, GOMPHIDAE). Descriptions and records of Malesian Odonata, 1. Tol, J. van, 1990. Key to the Malesian species of Leptogomphus Selys, with the description of a new species from Sabah (Odonata, Gomphidae). Descriptions and records of Malesian Odonata, 1. — Tijdschrift voor Entomologie 133: 97-105, figs. 1-37, tabs 1-2 [ISSN 0040- 7496]. Published 31 July 1990. A new species of Leptogomphus, L. pasia, based on one female from S. Sabah (Northern Borneo), is described. A congeneric and presumably conspecific male from E. Sabah is also described. A key to the eleven species of this genus recorded from Malaysia, Indonesia, the Philippine Islands, and the adjacent areas in Burma and Thailand, is included. Many characters are illustrated for the first time. J. van Tol, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Key words. — Leptogomphus; key; Malesia; new species. INTRODUCTION During the 1987 expedition of the Rijksmuseum van Natuurlijke Historie (Leiden) to Sabah, north- ern Borneo, a wealth of interesting Odonata was collected. In this paper a new species of Lepto- gomphus Selys is described, based on a single fe- male. It was collected in the vicinity of Long Pasia, a village still surrounded by vast areas of virgin rainforest. This area, situated close to the borders of Sarawak and Kalimantan at c. 1000 m above sea level, is poorly investigated for the aquatic fauna up to now. The Odonata fauna seems to be most closely related to those of Mount Dulit and other higher peaks in Sarawak. This may be exemplified by re- cords of Coeliccia cyaneothorax Kimmins, formerly only known from Mt. Dulit, and Burmagomphus insularis Laidlaw, of which only the holotype from Sarawak was known. A complete list of the species encountered will be published in due time. SYSTEMATIC PART Leptogomphus pasia spec. nov. (Figs. 1-6, 8, 16, 32-33) Type material. — Holotype female: 'RMNH Leiden/N. Borneo, S. SABAH. Beaufort, 105 km S of: Long Pasia area. Sungai Pasia near confluence with Sungai Maga. Rockpools and stream. Lower montane evergreen tropical rainforest. 1250 m. 4°26’N 115°40’E. 3 April 1987. Leg. C. van Ach- terberg’, in Museum Leiden (RMNH). Other material. — Sabah, 60 km W of Lahad Datu: Danum Valley Field Centre, 4°58’N 117°48’E. 2 May 1987, 1 male (J. van Tol) in RMNH. Description Female. — Closely related to Leptogomphus lansbergei and L. semperi, but immediately recog- nizable by its extremely long vulvar lamina (vulvar scale) (authors of species-group names in Lepto- gomphus are given in table 1). Head. — Depressed; frons, clypeus and labrum protruding. Labium with prementum dark brown to brownish black, lateral lobes greenish yellow, broadly bordered with brownish black, especially distally; movable hooks brownish black, central part of mentum pale brown, laterally bordered with creamish white. Mandibles reddish brown-black, but visible part beside the labrum mostly greenish yellow. Labrum black with two relatively small sub- triangular pale spots close to the clypeus. Clypeus black, with a paired oval depression on postclypeus, together approximately as wide as clypeus between the emarginate anterior side. Frons distinctly swollen with a broad greenish yellow transversal stripe, covering nearly half of the surface, pale stripe interrupted in the middle by a dark triangular 97 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 1 Species included in the key of the Leptogomphus species of Malesia, with original references Species Author; date: pages L. coomansi Laidlaw 1936: 267-269 L. gestroi Selys 1891: 476-477 L. inclitus Selys 1878: 444 (sep 39) L. lansbergei Selys 1878: 446 (sep 41) L. lansbergei assimilis Kriiger 1899: 308 L. mariae Lieftinck 1948: 249-251 L. palawanus Asahina 1968: 364-365 L. pendleburyi Laidlaw 1934: 555-556 L risi Laidlaw 1933: 95-96 L. semperi Selys 1878: 443 (sep 38) L. williamsoni Laidlaw 1912: 94-95 Original genus Leptogomphus Leptogomphus Gomphus (Leptogomphus) Gomphus (Leptogomphus ?) Leptogomphus Leptogomphus Leptogomphus Leptogomphus Leptogomphus Gomphus (Leptogomphus) Leptogomphus Type depository RMNH / H male MCSN / H male BMNH / L female IRSN + SMTD / S female ZMPA ? / S male + female RMNH / H female CASC / H male BMNH / H male BMNH / H male IRSN / H male BMNH / H male Abbreviations: BMNH = British Museum (Natural History), London; CASC = California Academy of Sciences, San Francisco; IRSN = Institut Royal des Sciences Naturelles de Belgique, Bruxelles; MCSN = Museo Civico di Storia Naturale, Genoa; RMNH = National Museum of Natural History, Leiden; SMTD = Staatliches Museum für Tier- kunde, Dresden; ZMPA = Zoological Museum Polish Academy of Sciences, Warszawa. - H = holotype, L = lecto- type, S = syntype. Note: The type of L. assimilis should be selected from material in the Dohrn collection, which was deposited in Stet- tin (now Szczecin, Poland); some parts of the collection of this museum have been transferred to ZMPA. emargination. Vertex shiny black, but dullish black behind the tubercles posterior to the lateral ocelli; occiput also dull black, tubercles behind lateral ocelli consisting of a larger pair in the centre between the ocelli and a smaller pair latero-posteriorly of the former; occiput without tuberculi (fig. 2), which are usually present in congeneric species (e.g. L. semperi, L. coomansi, L. mariae and L. williamsont) (see also Lieftinck 1948, plate 8); occiput poste- riorly with a distinct triangular emargination. Thorax (for nomenclature of thoracic markings, cf. Williamson 1907: fig. 29 (1)). - Prothorax with anterior lobe erect, black, but greenish yellow in lateral edges; middle lobe with deep median depres- sion anteriorly. Synthorax (fig. 5) brownish black with pale markings yellow; dorsal thoracic stripe connected with mesothoracic half collar (as in L. coomansı), antehumeral stripe narrow, somewhat wider at posterior side; pale markings on mesepi- meron and metepisternum more or less as in L. lansbergei, but pale area on metepimeron much smaller than in L. lansbergei (the dark stripe under the pale area approximately of equal width). Legs stoutly built; hind femora with two rows of 9-12 distinct short spines on innerside; all tibiae with two rows of 9-10 long spines on innerside. Wings: forewing 37 mm, hind wing 35 mm. Forewing with 16 Ax and 13 Px; hindwing with 12 Ax and 13 Px. Hind wing with one cubito-anal crossvein; an in- complete crossvein proximal to first primary Ax present; distal costal brace closer to proximal costal brace than to nodus; triangle, supratriangle and subtriangle all without crossveins. Pterostigma 3.5 mm. Abdomen. — Length 39 mm. Dark brown, nearly black, except for a yellow middorsal stripe, which is rather wide on segments 1-2, but much narrower to 98 nearly indiscernable on segments 3-8; lateral side of segment 1 dark yellow, lateral side of segment 2 and antero-lateral side of segment 3 also dark yellow. Auriculae hardly developed, yellow. Vulvar lamina diagnostic (figs. 3-4), extremely elongate, the tip reaching posterior margin of segment 10, very slender, the top acutely v-shaped. Male. - As female (figs. 6, 16), but pale spots on labrum larger than in female, subquadrangular (fig. 8); hind femora with many short spines on inner- side instead of the rows of medium-sized spines as in the female; wings with anal triangle three-celled, crossveins meeting in one point; secondary genita- lia and anal appendages very similar to L. lansber- gei, but superior appendage of /ansbergez in dorsal view curved outwards more conspicuous than in pasia (figs. 30, 32); base of superior appendage of pasia bulgy on outer margin; consequently, laterov- entral ridge with spines along the outer margin of the superior appendage nearly straight in lansber- get and distinctly curved in pasta (figs. 31, 33). Although I consider this male conspecific with the female described above, I prefer to include only the female in the type series of this species. Differential diagnosis. - The female of this spe- cies, superficially looking as L. lansbergei, is most easily distinguished from its congeners by the pe- culiar shape of the vulvar lamina. Also the male is very similar to L. lansbergei, but can be distin- guished from most species by the thoracic mark- ings, and from lansbergei by the shape of the super- ior appendages. Etymology. - Pasia from the type locality Long Pasia. To be considered as a noun in apposition. Discussion Up to now the following species of the genus Leptogomphus were known from Borneo: L. coo- mansi, L. martae, L. pendleburyi and L. williamsoni. There is still some doubt about the occurrence of L. lansbergei assimilis on Borneo, of which a male was recorded from Bettotan, 4 August 1927 by Laidlaw (1931). This specimen is not in the collection of the British Museum (Natural History) (Brooks in litt.). Also L. semperi, known from Mindanao, should be VAN TOL: Malesian species of Leptogomphus taken into account when considering the fauna of Borneo. Earlier records by Martin (1904) and Wil- liamson (1907) were considered erroneous by Laid- law (1936). This species was also attributed to the lansbergei Formenkreis by Laidlaw (1936). L. pasia shows some affinity to L. coomansi and L. mariae, sharing the connection between the dor- sal stripe and the mesothoracic half collar (cf. Lief- tinck 1948, plate 8). The absence of spiny tubercles on the occipital plate, and the peculiar shape of the Figs. 1-7. Leptogomphus from northern Borneo. - 1, L. pasta female holotype, head in frontal view; 2, Idem, hind margin of occiput in dorsal view; 3-4, Idem, last abdominal segments in ventral and left lateral view; 5, Idem, synthorax in left lateral view; 6, L. cf. paria, male from Danum Valley, synthorax in left lateral view; 7, L. pendleburyi, male holotype, synthorax in left lateral view. 99 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 8-14. Leptogomphus. - 8, L. cf. pasia, male from Danum Valley, head in frontal view; 9, L. pendleburyi, male holotype, idem; 10, L. lansberget assimilis, male from S. Sumatra, idem; 11, L. inclitus, female lectotype, head in frontal view; 12, L. lansbergei assimilis, female from S. Sumatra, last abdominal segments in ventral view; 13, L. inclitus, female lectotype, idem; 14, L. inclitus, female lectotype, hind margin of occiput in dorsal view. 100 vulvar scale are diagnostic characters for the pres- ently described species. As already mentioned above, it is, however, most closely related to L. lansbergei from Java and Sumatra. KEY TO THE LEPTOGOMPHUS SPECIES OF MALESIA In the next key I have summarized the characters to distinguish the species of Leptogomphus occur- ring in Malesia (i.e. the Malay peninsula, Indonesia, the Philippine Islands and the island groups north- east of New Guinea). I have added several species recorded from Burma and Thailand that might oc- cur in this area. All species mentioned in the key are listed in table 1 with the author and a reference to the original publication. Leptogomphus svibleri Asahina, however, recorded from Burma and Thai- land, is not included in the key. The absence of an incomplete basal cross-vein (Asahina 1970: 116) is an indication that this species is not a Leptogomphus in the present sense. Asahina (1986: 17) has placed it in Heliogomphus. The genus Leptogomphus Selys as defined by Carle (1986) forms with Africogomphus Fraser and Heliogomphus Laidlaw the tribe Leptogomphini in the Epigomphinae. Characters to distinguish Leptogomphus and Heliogomphus include lamina on fore tibia of male (present/absent, respectively), basal subcostal crossvein (present/absent), cells of anal triangle in male (meet in one point/serially), lamina anterior (raised transverse-platelike/bilo- bate), superior appendages of male (more or less straight/lyrelike). The key is mainly based on characters in the markings of the specimens, predominantly of the head and the thorax. Colour pattern is a reliable character in this group and the interpretation of Table 2 Species ( coomansi gestroi inclitus lansbergei lansbergei lansbergei assimilis mariae palawanus pasia pendleburyi risi semperi williamsoni | +++t++++++++ | H++++++)] (1) Dorsal stripe complete (+) / incomplete (—) (2) Dorsal stripe connected (+) / not connected (—) with mesothoracic half collar (3) Yellow mark on frons separate (+) / one stripe (—) (4) Anterior side of postclypeus with yellow patches (+) / dark (—) (5) Yellow mark on vertex present (+) / absent (—) A We _ | ++|HH++++] VAN Tor: Malesian species of Leptogomphus conspecificity is based on this character for several species. Structural characters include the appen- dages of the male and the vulvar scale of the female. Also the tubercles on vertex and occiput are fre- quently used, but these characters have to be con- sidered carefully. I have seen several specimens with very apparent differences in the tubercles on each side of the head. Asahina (1986: 22) mentions female specimens of L. gestroz Selys with an occip- ital structure significantly different from the (allo) type. This phenomenon was also observed by him in Merogomphus parvus. The distribution of the species was taken from the following works: for India in the wider sense (Laidlaw 1922, Fraser 1927, 1934), for Thailand (and Burma) (Williamson 1907, Asahina 1986), for Sumatra (Krüger 1899, Ris 1927, Lieftinck 1935, 1948), for Java (Lieftinck 1934), for Borneo (Laid- law 1912, 1914, 1934, 1936, Lieftinck 1948) and for the Philippine Islands (Needham & Gyger 1937, Asahina 1968). Lieftinck (1954) is a most useful summary of knowledge for the larger part of this area. Table 2 provides a data matrix for several impor- tant characters used in the dichotomous key below. I have examined all species myself. 1. Thorax with complete antehumeral stripe run- ning close to the humeral suture (e.g. figs. 16- 17, 19-21) Antehumeral stripe on thorax absent or strongly reduced, usually to a small pale spot in the posterior corner of the thorax above the humeral suture (e.g. figs. 15, 18) 10 2. (1) Vertex with yellow marking; transversal stripe on frons not interrupted by a longitudi- nal black line inithe, middle; td. 3 (4) (5) (6) (7) (8) (9) Ar — — — ap + att + 4 im ? 24 = AL + = - ? u I ie al n a — — al — — ali = i r = + ? — _ r = SF Ar EL di Da + ons (+) — — r ? ? sr = an 5A Al aif + a = a = dE gE + + + a + ? (6) Antehumeral stripe present (+) / absent (—) / re- duced (r) (7) Valve (female) far beyond segment 9 (+) / shorter (—) (8) Tubercles on occiput in female present (+) / absent =) (9) Tubercles on abdominal segment 10 of male present (+) / absent (—) 101 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 19 20 21 Figs. 15-21. - Thoracic pattern of left half of Leptogomphus species, dorsal view. - 15, L. pendleburyi, male holotype; 16, L. cf. pasia, male from Danum Valley; 17, L. williamsoni, female E. Borneo; 18, L. coomansi, male E. Borneo; 19, L. lansbergei assimilis, male S. Sumatra; 20, L. gestroi, male Thailand, Doi Suthap; 21, L. #nclitus, female lectotype. — Vertex black, without yellow or otherwise pale — Females ............... 7 markings; transversal stripe on frons inter- rupted by a longitudinal black line in the middle (CREfios ss MO Mio RE Tor. RER BRR. * 4 . (2) Characteristic markings on thorax with very short dorsal stripe and large mesothoracic half collar (fig. 17). Occiput black. Postclypeus with latero-anterior corners pale coloured. Dis- tabution#BorneoY.. nee ae. ER EN AA ee L. williamsoni Laidlaw Markings on dorsal side of thorax less charac- teristic, with a mesothoracic half collar as in most other congeneric species and a complete dorsal stripe, which is not connected with the mesothoracic half collar (fig. 20). Occiput with a pale centre. Postclypeus brown, without pale markings. Superior appendages of male, figs. 22-23. Distribution: Thailand, Burma. ....... OS calc ee er MNT L. gestrot Selys Note. Redescription of male and female with illustra- tions in Lieftinck (1960: 238-240, fig. 6). MAO) IMAL ES sas aa EN Henk: 5 Note. The male of L. #nclitus (distribution Thailand) is unknown. See fig. 21 for the thoracic markings of the female. In all other species these markings agree well between male and female. 102 . (4) Tip of superior appendage sharp with sub- terminally a blunt ventral tubercle, thus in lat- eral view bifid in appearance (figs. 34-35). Dis- tributions Philippineiisiands aac, Mr mean ER eee vai L. semperi (Selys) Tip of superior appendage sharp, but without subterminal ventral tubercle ……. 6 (5) Dorsal stripe (nearly) connected with me- sothoracic half collar (fig. 5, 16); head pattern as in fig. 8; superior appendages of male as in figs: 32-334 Distribution: Borneo) .... ar We tl eon oer we L. cf. pasia spec. nov. Dorsal stripe and mesothoracic half collar se- parated by a narrow black line; head pattern as in fig. 10; superior appendages of male as in figs. 30-31. Distribution: Java and Sumatra. ite Soy N L. lansbergei (Selys) Note. Specimens from Java belong to subspecies lansbergei and specimens from Sumatra to subspecies assimilis. According to Ris (1927: 28-29) the superior appendages of both subspecies are different. . (4) Valvula vulvae extremely long, extending to the hind margin of segment 10 (figs. 3-4); oc- ciput without tubercles (fig. 2); head pattern as in fig. 1. Distribution: northern Borneo. ..... SIERRA GEE RSS REES L. pasta spec. nov. VAN TOL: Malesian species of Leptogomphus 25 24 27 26 29 FTR Figs. 22-37. - Leptogomphus superior appendages of male, for all species in dorsal and lateral view respectively. — 22- 23, L. gestroi from Thailand, Doi Suthap; 24-25. L. rist from Thailand, Nakhom Sithammarat province; 26-27, L. coomansi from E. Borneo; 28-29, L. pendleburyi, holotype from northern Borneo; 30-31, L. lansbergei lansbergei from W. Java; 32-33, L. cf. pasia from eastern Sabah, Danum Valley; 34-35, L. semperi from Basilan I.; 36-37, L. palawanus from C. Palawan. 103 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 — Valvulae vulvae much shorter, extending not further than the hind margin of segment 9 8. (7) Occiput with four slender spiny tubercles; dorsal stripe connected with mesothoracic half collar. Distribution: Philippine Islands. ...... DR RE tr NAR L. semperi (Selys) Note. A female in the collection of the RMNH from Mindanao keys out here. I consider this specimen doubtfully conspecific with the male from Basilan I., which was compared with the type of L. semperi by M. A. Lieftinck (for a detailed description of the male specimen, see also Laidlaw 1936: 269). — Occiput with ridges or smooth, but always with- out spiny tubercles; dorsal stripe and mesotho- racic half collar separated by a narrow black line 9. (8) Hind margin of occiput nearly straight or with two roundish tubercles in the middle (size of tubercles showing much variation). Labrum black with a paired subrectangular pale spot (cf. fig. 10). Last abdominal segments, fig. 12. Dis- tributions avarandiSumattay nun. Bee Metre A L. lansbergei (Selys) Note. According to Lieftinck (1948, p. 245 and fig. 9) specimens from Java (subspecies lansbergei tend to have more conspicuous tubercles than specimens from Sumatra (subspecies assimilis Krüger). — Hind margin of occiput against both eyes with an erect squarish tubercle (fig. 14). Labrum pale (creamish yellow ?) with a narrow dark stripe along the anterior margin and a very narrow longitudinal dark line in the middle. Last ab- dominal segments, fig. 13. Distribution: Thai- landen er L. inclitus (Selys) 10. (1) Middle of postclypeus with a yellow patch; antehumeral stripe absent or reduced to a small spot in posterior part of synthorax above hu- meral suture (fig. 18). Superior appendages of male, figs. 26-27. Distribution: Borneo. ...... ERO ON. L. coomansı Laidlaw — Postclypeus black or brownish black, without yellow markings; antehumeral stripe reduced to a larger or smaller pale spot in posterior part of synthorax above humeral suture ...... 11 11. (10) Yellow mark on frons a single transversal stripe (fig. 9); thoracic pattern, fig. 7; superior appendages of male, figs. 28-29. Distribution: Northern Borneo. .... L. pendleburyi Laidlaw Note. Only known from male holotype. — Transversal pale stripe on frons divided in the middle, or at least with distinct dark triangular emarginations in the middle ........... 12 12, MIG. a ean Ee 15 Note. The male of L. mariae is unknown. TDA EN 14 13. Superior appendages in dorsal view curved in- 104 wards, tip bluntly shaped (figs. 24-25). Distri- bution: Malaysias EP ea: L. risi Laidlaw — Superior appendages in dorsal view straight, tip bifid (figs. 36-37). Distribution: Palawan. ENE MA L. palawanus Asahina Note. Specimens from Borneo that key out here, may belong to L. martae, of which the male is undescribed. See Lieftinck (1948, p. 248) for an illustration of the thoracic pattern of the female. 14. Occiput smooth, without tubercles. Distribu- tion: Malayase o L. risi Laidlaw — Occiput with conspicuous tubercles (Borneo andiPhilippinesislands) We 15 15. Area behind lateral ocelli with a sharp tubercle, directed posteriorly. Distribution: Palawan. RS DO a Se L. palawanus Asahina — Area behind the lateral ocelli with a semicircu- lar transversal rim divided into two parts by a central\deptessioni(Borneo) ME SE Ee L. mariae Lieftinck Note. This species is only known from the female holotype. In other species with tubercles on the oc- ciput, these tubercles may be absent or broken at one or both sides of the head. ACKNOWLEDGEMENTS I am indebted to Mr. Stephen J. Brooks, British Museum (Natural History) for his help during my visits and for promptly sending type material under his care, and to Dr. Erik J. van Nieukerken for his comments upon a draft of this paper. REFERENCES Asahina, S., 1968. Records and notes on Philippine Odo- nata. — Japanese Journal of Zoology 15 (4): 349-376, figs. 1-58 + plate 1-2. Asahina, S., 1970. Burmese Odonata collected by Dr. Ar- thur Svihla with supplementary notes on Asiatic Ce- riagrion species. — Japanese Journal of Zoology 16 (2): 99-126, figs. 1-68 + plate 1. Asahina, S., 1986. A list of the Odonata recorded from Thailand, Part XIV. Gomphidae-2. - Tombo 29: 7-53, figs. 1-201. Carle, F. L., 1986. The classification, phylogeny and bio- geography of the Gomphidae (Anisoptera). I. Classi- fication. - Odonatalogica 15 (3): 275-326, tables 1-2. Fraser, F. C., 1927. Indian dragonflies. Part XXVI. - Jour- nal of the Bombay Natural History Society 31(4): 882- 889, figs. 1-3. Fraser, F. C., 1934. Odonata. Volume 2. - The Fauna of British India including Ceylon and Burma: i-xxiii + 1- 398, figs. 1-119 + plate 1-4. Krüger, L., 1899. Die Odonaten von Sumatra. IL. Theil. Familie Aeschniden. — Stettiner entomologische Zeitschrift 59 (1898): 267-331, figs. Laidlaw, F. F, 1912. List of the Odonata taken on an expedition to Mt. Batu Lawi together with descriptions of supposed new species. — Journal of the Straits Branch of the Royal Asiatic Society 63: 92-99, figs. 1- DE Laidlaw, F. F, 1914. Contributions to a study of the dra- gonfly fauna of Borneo. — Part II. The Gomphinae and Chlorogomphinae. — Proceedings of the Zoological Society of London 1914: 51-63 + plate 1. Laidlaw, F. F., 1922. A list of the dragonflies recorded from the Indian Empire with special reference to the collec- tion of the Indian Museum. Part V. The subfamily Gomphinae. (with an appendix by F. C. Fraser). — Records of the Indian Museum 24 (3): 367-426, figs. EZRA Laidlaw, F. F., 1931. A synonymic list of dragonflies of the family Gomphidae (Odonata, Anisoptera) found in the Oriental region. — Transactions of the Entomological Society of London 78 (2): 171-197. Laidlaw, F. F, 1933. Notes on Malaysian dragonflies (Odo- nata), with descriptions of new species. — Bulletin of the Raffles Museum, Singapore, Straits Settlements 7 (1932): 95-96, figs. 1-2b. Laidlaw, F. E‚ 1934. A note of the dragonfly fauna (Odo- nata) of Mount Kinabalu and of some other mountain areas of Malaysia: with a description of some new or little known species. - Journal of the Federated Malay States Museums 17 (3): 549-561, figs. 1-4. Laidlaw, F F, 1936. On some examples of the genus Leptogomphus (Odon., Gomphidae). - Treubia 15 (3): 267-270, figs. 1-2. Lieftinck, M. A., 1934. An annotated list of the Odonata of Java, with notes on their distribution, habits and life- history. - Treubia 14 (4): 377-462. Lieftinck, M. A., 1935. A synopsis of the Odonata (Dra- gonflies) of Sumatra. - Miscellanea Zoologica Suma- trana 92/93: 1-23, figs. 1-2. Lieftinck, M. A., 1948. Descriptions and records of South- east Asiatic Odonata. - Treubia 19 (2): 221-278, figs. 1-19. VAN Tor: Malestan species of Leptogomphus Lieftinck, M. A., 1954. Handlist of Malaysian Odonata. A catalogue of the dragonflies of the Malay Peninsula, Sumatra, Java and Borneo, including the adjacent small islands. — Treubia 22 (supplement): i-xiii + 1-202. Lieftinck, M. A., 1960. On the identity of some little known Southeast Asiatic Odonata in European mu- seums described by E. de Selys Longchamps, with de- scriptions of new species. - Memorie della Società entomologica Italiana 38: 229-256, figs. 1-13. Martin, R., 1904. Liste des Neuropteres de l’Indo-Chine. - Mission Pavie Indochine 3: 204-221. Needham, J. G. & M. K. Gyger, 1937. The Odonata of the Philippines. - The Philippine Journal of Science 63 (1): 21-101, figs. 1-2 + plate 1-10. Ris, F., 1927. Odonaten von Sumatra, gesammelt von Edward Jacobson. - Zoologische Mededeelingen 10 (1): 1-49, figs. 1-24. Selys Longchamps, E. de, 1878. Quatriemes additions au synopsis des Gomphines. — Bulletin de l'Académie royale de Belgique (2) 46: sep. 1-106. Selys Longchamps, E. de, 1891. Viaggio di Leonardo Fea in Birmania e regioni vicini. XXXII. Odonates. - An- nali del Museo Civico di Storia Naturale di Genova (serie 2a) 10: 433-518. Williamson, E. B., 1907. The dragonflies (Odonata) of Burma and Lower Siam. — II. Subfamilies Cordulegas- terinae, Chlorogomphinae, and Gomphinae. - Pro- ceedings of the United States National Museum 33 (1571): 267-317, figs. 1-39. Received: 22 March 1990 Accepted: 24 March 1990 105 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 106 W. WITTMER Naturbistorisches Museum, Basel DIE FAMILIE MALACHIIDAE (COLEOPTERA) AUF SULAWESI 43. Beitrag zur Kenntnis der indo-malaiischen Fauna Wittmer, W., 1990. Die Familie Malachiidae (Coleoptera) auf Sulawesi. 43. Beitrag zur Kenntnis der indo-malaiischen Fauna. - Tijdschrift voor Entomologie 133: 107-120, figs. 1-29. [ISSN 0040-7496]. Published 31 July 1990. The species of the family Malachiidae occurring on Sulawesi are revised. Twenty-five species are enumerated, of which nineteen are described as new to science. The genus Luzonotroglops Pic, 1924 is synonymized with Falsolaius Pic, 1917, and the genus Flabellapalochrus Pic, 1923 with Dromanthomorphus Pic, 1921. Carphurus tondanus Champion, 1923 is synonymized with C. rufonotaticeps Pic, 1917. Dr. W. Wittmer, Naturhistorisches Museum, CH-4001, Basel, Schweiz. Key words. - Malachiidae, Sulawesi, new species. FINLEITUNG Von der Insel Sulawesi waren bisher nur 5 Arten der Familie Malachiidae beschrieben worden und zwar Carphurus celebensis Champion, C. rufonotaticeps Pic, C. tondanus Champion, Laius ribbei Pic und Dromanthomorphus cyaneus Pic. Carphurus tondanus musste eingezogen wer- den, weil diese Art Synonym mit C. rufonotaticeps ist. Zwei weitere Arten: Carphurus cf. rubroannu- latus Motschulsky und Latus cyaneus Guérin haben eine weitere Verbreitung und werden heute zum ersten Mal von Sulawesi gemeldet. Weitere 19 Ar- ten stellten sich als neu für die Wissenschaft heraus. Sie stammen von dem reichen Material, das ich Herrn Dr. Peter Hammond, London, verdanke, das im British Museum (Natural History) aufbewahrt wird und vom “Project Wallace”, sponsored by the Royal Entomological Society of London and the Indonesian Institute of Science, Results of Project Wallace No. 85 aufgesammelt wurde. Den Londoner Koleopterologen, den Herren M. J. D. Brendell, Dr. Peter Hammond, Dr. Christo- pher Lyal und Dr. Nigel Stork, die auf Sulawesi so erfolgreich tätig gewesen sind, gratuliere ich für die schönen dort erzielten Resultate. Des weiteren danke ich Herrn G. Hodebert, Pa- ris, für die sorgfältig ausgeführten Zeichnungen und Herrn Dr. Ch. O'Toole, Hope Museum, Oxford, für die ausgeliehenen Typen. Verwendete Abkürzungen: BM = British Mu- seum (Natural History), London; MP = Muséum de Paris, NHMB — Naturhistorisches Museum, Basel. SYSTEMATISCHER TEIL Bestimmungstabelle für die auf Sulawesi vorkommenden Gattungen 1. Die Flügeldecken bedecken das Abdomen voll- kommen. Fühler anscheinend 10-gliedrig, das Glied 2 steckt ganz im 1. Vordertarsen einfach, oder das Tarsenglied 2 ist mit einem Kamm versehen. „tonus Apt SERIE La er CRE 2 — Die Flügeldecken sind verkürzt und lassen mehrere Tergite unbedeckt. Fühler 11-gliedrig, das Glied 2 ist voll sichtbar. Glied 1 der Vorder- tarsen mit einem Kamm (Carphurinae).... 3 . Fühlerglied 3 difform, meist mehr oder weniger stark ausgehöhlt. Vordertarsen ohne Kamm etant Seren 5 RN ER behe Laius Guérin — Fiihler von Glied 3 an gekimmt. Glied 2 der Vordertarsentmit Kamm RAEE bes bapa sia wich cb: Dromanthomorphus Pic 3. Fliigeldecken einfach, ohne Auszeichnungen N — Flügeldecken gegen die Spitze mit einer mehr oder weniger deutlichen Grube oder Eindruck, mit vermehrten Haaren am Rand .......... els enne e Paracarphurus Wittmer 4. Halsschild gegen die Basis schwach bis stark verengt bis eingeschnürt, vor dem Basalrand mässig bis stark quer eingedrückt. Fühler und Korpegnormaljbehaactm nne: 3 — Halsschild vor der Basis schwach gerundet ver- engt, Seiten davor fast parallel, ohne queren Eindruck vor dem Basalrand. Körper und be- sonders die Fiihler, meistens auch die Beine 107 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 RR E NR Telocarphurus Wittmer 5. Halsschild gegen die Basis schwach verengt, vorderer Teil der Scheibe wenig bis kaum auf- gewölbt, Eindruck vor dem Basalrand nur schwach entwickelt. Rippen auf den Flügel- cedken ENENMMMER NE ER eo SE RIO IR RE e Carphurus Erichson — Halsschild gegen die Basis stark eingeschniirt, vorderer Teil der Scheibe stark gerundet aufge- wölbt, Eindruck vor dem Basalrand breit. Rip- pen auf den Flügeldecken fast immer vorhan- ENTE: Falsolaius Pic Carphurus Erichson, 1840 Bestimmungstabelle für die auf Sulawesi vorkommenden Arten 1. Halsschild mit mehr oder weniger feinen Quer- runzeln auf der Scheibe und an den SEIEN. NN NR ION. 2 — Halsschild ohne Querrunzeln auf der Scheibe, diese meistens glatt, schwach matt, oder punk- GENE NN es chp cus RN 8 2. Halsschild und Flügeldecken schwarz, letztere manchmal mit einer mehr oder weniger gros- sen und deutlichen hellen (weisslichen) Auf- hellung, oder vorwiegend weisslich .… 3 — Halsschild und Flügeldecken gelb oder hell- bruno ere AMEN ER MELA 7 3. Flügeldecken einfarbig schwarz, oder mit mehr oder weniger grossen und deutlichen hellen (weisslichen) meistens seitlich gelegenen Auf- hellungen, Naht immer dunkel .......... 4 — Flügeldecken fast ganz weisslich, nur an der Basis schmal angedunkelt, Seiten und Spitzen manchmal, jedoch noch schmäler als an der Basis, angedunkelt ....... C. albipennis sp.n. 4. Fiihler kürzer, letzte 4 bis 8 Glieder angedun- keltsbisischmarzins tn EED a 5 — Fühler länger (Abb. 2) einfarbig gelb, nur die Spitze des letzten Gliedes ist dunkel ........ TR ae a C. pallidicornis sp.n. 5. Vorderschienen dunkel bis schwarz. Flügel- decken glänzender, manchmal mit leichtem violettem Schimmer, Punktierung etwas tiefer, Elaaresschwarzigg ad. aaa mess 6 — Vorderschienen gelb. Flügeldecken weniger glänzend, manchmal leicht matt, ohne Metall- schimmer, Punkte etwas weniger tief, Haare preis Aarden mer. C. dumogaensis sp.n. 6. Grössere Art: 6 mm (9). Fühler (Abb. 6) sehr stark gezahnt (9). Flügeldecken schwarz mit schwachem violettem Schimmer ........... VAER: sine dona sia: C. celebensis Champion — Kleinere Art: 4-5 mm (9). Fühler weniger stark gezahnt (9). Flügeldecken schwarz, ohne Metallschimmer oder heller (weisslicher) Auf- 108 122 155 HEID C. strigilatus sp.n. . Der ganze Körper ist einfarbig gelb. Fühler (Abb. 7) länger, alle Glieder länger als breit, schwach gezahnt (8), einfarbig gelb oder die letzten 1 bis 5 Glieder angedunkelt, selten ist die Mittel- und Hinterbrust dunkel ......... SRI ee C. totopallidus sp.n. Nur die Oberseite ist einfarbig gelb, auf der Unterseite ist die Vorder- und Mittelbrust, mei- stens auch die Hinterbrust schwarz, manchmal auch das Abdomen, Basis der Mittel- und Hin- terschenkel, seltener der Vorderschenkel dun- kel. Fühler (Abb. 9) kürzer, einzelne Glieder so lang wie breit oder breiter als lang (8), einfar- big gelb oder ein paar der letzten Glieder sind EEnansenaeieS Bo tare ee eee ee UMILI AES ARE NEN C. partepallidus sp.n. . Halsschild einfarbig rot oder orange bis gelb heden Pe e O 9 Halsschild einfarbig schwarz oder zum grös- Sn Tes AE RON O NNT 10 . Grosse Art: 5-6 mm. Halsschild rot, glatt, in den Vorderecken befinden sich ein paar grobe Punktes Fligeldeckenischwatz see eee NE C. cf. rubroannulatus Motschulsky Kleine Art: 2.5-3 mm. Halsschild orange bis gelb, fein chagriniert, leicht matt. Fliigeldecken gelblich, basale Hälfte bis ein Drittel dunkel DALE AGREE IE C. basilimbatus sp.n. . Halsschild einfarbig schwarz ........... 11 Halsschild an der Basis schmal gelblich ge- STUER C. testaceolimbatus sp.n. MIK PITCINfARDISISCAWA EZRA EN 12 Kopf rot, an den Schläfen und manchmal an der Stirnbasis schmal schwarz. Flügeldecken schwarz mit einem weisslichen mehr oder we- niger ausgeprägten Langswisch ............ Nana mania ran C. brendelli sp.n. Kopf zum grössten Teil glatt, vereinzelt mit feinen Punkten besetzt. Halsschild ohne PunktelanidentSel ent rt sun Dr or 13 Kopf mit ausgesprochen tiefen und groben Punkten besetzt, besonders an der Basis, Zwi- schenräume glatt. Halsschild an den Seiten punktiert, Punkte an der Basis ineinander flies- send. AN MOENS C. rufonotaticeps Pic Kopf so breit wie der Halsschild, dieser mit feinen Querrunzeln basal an den Seiten des Quereindrucks. Punktierung der Fliigeldecken wenig deutlich, fast erloschen ............. N ne Ken AEN OR TORE C. storki sp.n. Kopf deutlich breiter als der Halsschild, dieser ohne feine Querrunzeln basal an den Seiten des Quereindrucks. Punktierung der Flügeldecken dEUTLICHICODE PETER ARE C. lyali sp.n. WITTMER: Malachiidae auf Sulawesi 1 mm Abb. 1-6. Fühler von Carphurus. — 1, C. albipennis sp.n. 8; 2-3, C. pallidicornis sp.n. (2, 8. 3, 2); 4-5, C. dumogaensis sp.n. (4, 8. 5, 2); 6, C. celebensis Champion 9. Carphurus albipennis sp.n. (Abb. 1) Material. - Holotypus: &, Indonesia, Sulawesi Utara: Dumoga Bone N.P., lowland forest, ca. 200 m, 11.vii.1985 (BM). Paratypen: Gleicher Fundort wie Holotypus, ii, vi, vii, x, xi und xii.1985 (14 BM, 12 NHMB); Fog 13, trays 81, 105, 106 (Plot A), 230/240 m, 11.vii.1985, (2 BM, 1 NHMB). 6. Kopf einfarbig schwarz, von der Mitte nach vorne manchmal teilweise leicht, selten ganz aufge- hellt; Maxillarpalpen gelb; Fühler gelb, die letzten 3 bis 5 Glieder mehr oder weniger dunkel; Hals- schild, Schildchen und Abdomen schwarz; Fliigel- decken weisslich, ringsum schmal, schwach ange- dunkelt, manchmal nur angedeutet; Schenkel nur an der Basis oder fast bis zu den Knien schwarz bis dunkel, Schienen gelb oder ein wenig angedunkelt, Tarsen gewöhnlich gelb, selten schwach angedun- kelt. Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewölbt, Querrunzeln an der Stirnbasis nur angedeutet, restliche Oberfläche fast glatt, zer- streute Haarpunkte sichtbar. Fühler (Abb. 1) die Schultern ein wenig überragend, Glieder:4 bis 10 schwach stumpf gezahnt, einzelne Glieder ein we- nig breiter als lang, 3 zur Spitze verbreitert. Hals- schild kaum merklich breiter als lang, Seiten gerun- det, gegen die Basis ein wenig stärker verengt als nach vorne, Scheibe regelmässig gewölbt, vor der Basis nicht quer eingedrückt und abgesetzt, Ober- fläche dicht mit feinen Querrunzeln bedeckt. Flü- geldecken nach hinten etwas erweitert, 3 bis 4 Ter- gite unbedeckt lassend, fast glatt, zerstreut mit ziemlich grossen, ganz flachen Haarpunkten be- setzt. Behaarung des ganzen Körpers fein und greis, nur auf dem Kopfe und Halsschild ein paar gröbere, dunkle, längere Haare. Länge: Knapp 3 mm. Neben C. dumogaensis Wittmer zu stellen, der nachfolgend beschrieben wird. Carphurus pallidicornis sp.n. (Abb. 2-3) Material. - Holotypus: &, Indonesia, Sulawesi Utara: Gng. Ambang F.R., near Kotamobagu, Gng. Muajat sum- mit area, ca. 1780 m, 30.v.-2.vi.1985 (BM). Paratypen: 109 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 1 mm aan = = I N PESI Abb. 7-12. Fühler von Carphurus. — 7-8, C. totopallidus sp.n. (7, 8. 8, 2); 9-10, C. partepallidus sp.n. (9, 8. 10, 2); 11, C. basilimbatus sp.n. 8; 12, C. testaceolimbatus sp.n. 6. Gleicher Fundort wie Holotypus, iv und xi.1985 (1 BM, 1 NHMB); Clarke Camp, lower montane forest, 1140 m, x.1985 (1 BM, 1 NHMB); Sulawesi Tengah: Mt. Tambu- sisi, 6500’, 01° 38 ’S-121° 23 ’E, 8-9.iv.1980, M.J.D. Bren- dell (1 BM, 2 NHMB). Kopf selten einfarbig schwarz und einer kleinen, schlecht begrenzten hellen Makel zwischen den Au- gen, oder der ganze Vorderkopf bis hinter die Füh- lerwurzeln ist orangebraun; Fühler gelb, nur die Spitze des letzten Gliedes ist mehr oder weniger breit schwarz; Halsschild, Schildchen, Flügeldecken und Unterseite schwarz, nur bei 1 Exemplar ist das zweit- und drittletzte Tergit ein wenig aufgehellt; alle Tibien und Tarsen gelb, bei den Mittel- und Hintertibien manchmal ein wenig angedunkelt, Schenkel schwarz, an der Spitze mehr oder weniger breit gelb, bei den vorderen am breitesten gelb. 6. Kopf mit den stark vorstehenden Augen brei- ter als der Halsschild, Stirne leicht gewölbt, einige Querrunzeln an der Basis, davor grob punktiert, nur in der Mitte eine kleinere fast punktfreie Stelle. Fiihler (Abb. 2) ziemlich lang, ca. 2% mal so lang wie der Halsschild, von Glied 3 an gezahnt, 3 ein wenig länger als die folgenden. Halsschild kaum merklich breiter als lang (22X21), Seiten kaum gerundet, gegen die Basis ein wenig verengt, Scheibe regelmässig gewölbt, vor der Basis nur ganz schwach eingedrückt, Oberfläche dicht mit Quer- runzeln bedeckt. Flügeldecken nach hinten leicht erweitert, 2% bis 3 Tergite unbedeckt lassend, Ober- fläche ziemlich dicht mit feinen manchmal etwas erloschenen Punkten besetzt, Behaarung ziemlich 110 dicht, dunkel. Q. Die Fühler (Abb. 3) sind kürzer als beim &, weniger stark gezahnt, sonst mit dem @ überein- stimmend. Länge: 4-5 mm. Ein wenig grösser als C. dumogaensis Wittmer, mit diesem nahe verwandt, zu unterscheiden durch die längeren, stärker gezahnten Fühler, die fast ein- farbig gelb sind, ausgenommen die Spitze von Glied 11 und die durchwegs gelben Vorderschienen und Tarsen. Carphurus dumogaensis sp. n. (Abb. 4-5) Material. - Holotypus: @, Indonesia, Sulawesi Utara: Dumoga Bone N.P., G. Mogogonipa summit, 1008 m, v.1985 (BM). Paratypen: Gleicher Fundort wie Holoty- pus, v und xi.1985 (15 BM, 12 NHMB); Clarke Camp, lower montane forest, 1140 m, iv, v und x.1985 (4 BM, 5 NHMB). Schwarz bis schwarzbraun, Kopf auf der vorde- ren Hälfte meistens mehr oder weniger rötlich bis gelblich aufgehellt; Fühler gelb, die letzten paar Glieder ein wenig angedunkelt oder ganz dunkel; die letzten Tergite manchmal gelblich; Schenkel meistens schwarz, oder an den Knien kurz gelb, Tibien ein wenig aufgehellt, die vorderen meistens ganz gelb, Tarsen gelb bis braun. 8. Kopf mit den Augen deutlich breiter als der Halsschild, Stirne leicht gewölbt, Stirnbasis mit ein paar deutlichen Querrunzeln, davor und neben den Augen ziemlich grobe Punkte, die gegen die Mitte und nach vorne rasch feiner werden. Fühler (Abb. 4) ungefähr so lang wie die Flügeldecken, Glieder 4 bis 10 ziemlich scharf gezahnt, 3 gegen die Spitze verbreitert. Halsschild kaum merklich breiter als lang, Seiten gerundet, gegen die Basis ein wenig stärker verengt als nach vorne, Scheibe regelmässig gewölbt, vor der Basis nicht quer eingedrückt und abgesetzt, Oberfläche dicht mit Querrunzeln be- deckt. Flügeldecken nach hinten leicht erweitert, 4 Tergite unbedeckt lassend, Oberfläche ziemlich dicht mit feinen, etwas erloschenen Punkten be- setzt, leicht matt wirkend. Behaarung wie bei C. albipennis Wittmer. ®. Fühler (Abb. 5) ein wenig kürzer und weniger breit gezahnt. Länge: 3-3.8 mm. Eine mit C. albipennis Wittmer nahe verwandte Art, die sich durch etwas grössere Gestalt und die dunklen, etwas stärker punktierten Flügeldecken unterscheidet. Der Kopf ist bei dumogaensis im Durchschnitt um ca. 15% breiter als der Halsschild und misst im © 0.8 mm, bei albipennis ist er nur ca. 8-9% breiter als der Halsschild und misst im © 0.7 mm. Carphurus celebensis Champion (Abb. 6) Carphurus celebensis Champion, 1923: 12. Die Art wurde nach 1 9 aus Tondano, Celebes beschrieben, der Holotypus befindet sich im Hope Museum Oxford, ex Wallace. In dem vorliegenden Material ist C. celebensis nicht vertreten. Die Art ist grösser als alle anderen auf Sulawesi vorkom- menden Arten mit quergerunzeltem Halsschild. Für die Abb. 6 des Fühlers diente der Holotypus. Der Fühler erinnert im Bau und in der Grösse an C. pallidicornis sp.n. &, doch ist er bei celebensis schwarz, nur die ersten Glieder sind hauptsächlich auf der Unterseite aufgehellt. Der Kopf und Hals- schild besitzen einen schwachen grünlichblauen und die Flügeldecken einen blauvioletten Schim- mer. Carphurus strigilatus sp.n. Material. — Holotypus: g, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Banks of River Tumpah, i.1985 (BM). Paratypen: Gleicher Fundort wie Holotypus, ii, iv, vi und viii.1985 (16 BM, 12 NHMB); idem lowland forest, ca. 200 m, i, ii, iii, iv, v, vi, vii und viii.1985 (24 BM, 14 NHMB); idem near Edward’s Camp, ca. 500 m, iv und x.1985 (5 BM); idem Fog 9, Tray 46, mixed crops, 1.iii.1985 (1 BM); idem Hog’s Back Camp, lowland forest, 492 m, vii.1985 (1 BM). Kopf vorwiegend schwarz, vorderer Teil bis über die Fühlerwurzeln rötlichbraun, manchmal er- streckt sich die helle Färbung bis zwischen die Au- WITTMER: Malachiidae auf Sulawesi gen; Fühler schwärzlich, erste 3 bis 5 Glieder gelb- lich; Halsschild und Schildchen schwarz; Fliigel- decken einfarbig schwarz oder schwärzlich, oder jede Decke mit einer hellen Längsaufhellung, mehr oder weniger deutlich, meistens mehr an den Seiten gelegen, ohne sie zu berühren; Unterseite schwarz, letzte 1 bis 3 Segmente oft gelbraun; Beine schwarz, oft sind die Tarsen, meistens nur die vorderen, aufgehellt. Die übrigen Merkmale inklusive die Fühler stim- men mit der Beschreibung von C. dumogaensis überein, bis auf die Flügeldecken, die glänzender sind mit gröberer Punktierung, Behaarung etwas kräftiger, schwarz. Länge: 4-4.3 mm. Diese Art ist nahe mit C. dumogaensis Wittmer verwandt, sie unterscheidet sich durch etwas grös- sere Gestalt, die dunkeln Beine bei denen höchstens die Vordertarsen gelblich sind, selten sind die Vor- derschenkel an den Knien und die Vordertibien an der Spitze kurz aufgehellt. Die Flügeldecken sind von variabler Färbung, bei dumogaensis immer dunkel, sie glänzen, haben tiefere Punkte und die Behaarung ist kräftiger, dunkel, bei dumogaensis greis. Carphurus totopallidus sp.n. (Abb. 7-8) Material. — Holotypus: &, Indonesia, Sulawesi Utara: Dumoga Bone N.P., lowland forest, ca. 200/300 m, viii.1985 (BM). Paratypen: Gleicher Fundort wie Holoty- pus, ii, iii, iv, vii, viii, x, xi und xii.1985 (13 BM,8 NHMB); idem Fog 1, trays 2, 5, 12, 87 (Plot A), 230 m, 5.11.1985 (3 BM, 1 NHMB); idem Fog 3, trays 6, 43, 47, 54, 55, 67, 85, 88, 95 (Plot B), 315 m, 8.ii.1985 (9 BM, 2 NHMB); idem Fog 5, trays 8, 10, 17, 24, 27, 30, 31, 41, 43, 46, 47,52, 54, 72, 91, 101, 106 (Plot C), 400 m, 11.ii.1985 (21 BM, 2 NHMB); idem Fog 11. trays 23, 24, 36, 41, 43, 52, 60, 65, 67, 69, 71, 72, 74, 76, 82, 89, 93, 94, 95, 96 (Plot A), 230 m, 10.111.1985 (22 BM, 3 NHMB); idem Fog 13, trays 4, 9, 11, 12, 14, 22, 25, 26, 30, 34, 41, 50, 54, 68, 78, 79, 85, 86, 87, 88, 90, 93, 98, 99, 100, 103 (Plot A) (21 BM, 6 NHMB); idem Fog 15, trays 44, 60, 91 (PLot C) (3 BM); idem Fog 26, trays 27, 74, 76, 77, 86, 102, 107, 112 (Plot A) (10 BM). Einfarbig blassgelb, ausgenommen die Fiihler, bei denen die Glieder 5 oder 6 bis 11 dunkel sind, seltener 8 bis 11, oder nur 11, selten sind die Fühler einfarbig gelb; manchmal ist die Unterseite ange- dunkelt. 8. Kopf mit den Augen breiter als der Halsschild, fast so breit wie die Flügeldecken an den Schultern, Stirne leicht gewölbt, Querrunzeln an der Stirnbasis wenig ausgeprägt, davor wenige Punkte, die gegen die Mitte der Augen erlöschen, von hier ab fast glatt. Fühler (Abb. 7) ungefähr so lang wie die Flügeldecken, Glieder länger als breit, gegen die Spitze leicht verbreitert, 5 bis 10 unter sich unge- fähr gleich lang. Halsschild ungefähr so lang wie din TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 breit, Seiten gerundet, gegen die Basis kurz verengt, Scheibe gewölbt, vor der Basis leicht quer einge- drückt, dieser Teil ein wenig aufgewölbt; Oberflä- che mit feinen Querrunzeln die von der Basis bis kurz über die Mitte oder noch weiter nach vorne sichtbar sind. Flügeldecken nach hinten leicht ver- breitert, ein wenig klaffend, ca. 3 Tergite unbedeckt lassend, Oberfläche bei einzelnen Individuen fast glatt, höchstens Haarpunkte sichtbar, bei anderen mit deutlicher, etwas erloschener Punktierung, Be- haarung sehr fein, hell. Q. Eigentlich nur durch das Fehlen des Kammes am Glied 1 der Vordertarsen vom & verschieden, denn die Fühler (Abb. 8) zeigen wenig Unter- schiede, sie sind kaum merklich kürzer. Länge: 2-3 mm, mit dem Abdomen, das bei den Q oft stark vorsteht. Neben C. testaceipennis Pic zu stellen, der je- doch einen glatten, an der Basis ein wenig stärker eingedrückten Halsschild besitzt. Die Fühlerglieder sind bei testaceipennis zur Spitze weniger verbrei- tert. Carphurus partepallidus sp.n. (Abb. 9-10) Material. — Holotypus: &, Indonesia, Sulawesi Utara: Dumoga Bone N.P., lowland forest, malaise trap, ca. 200 m, ii.1985 (BM). Paratypen: Gleicher Fundort wie Holo- Fog 13, trays 77, 82, 100 (Plot), 11.vii.1985 (3 BM); idem Edward's Camp, lowland forest, malaise trap, ca. 664 m, 26.iv-28.v.1985, 1-16.x.1985 (2 BM); Sulawesi Tengah: near Morowali, Ranu River area, ii und iii.1985 (3 BM). Kopf, Halsschild und Schildchen orange; Fühler gelborange, letzte 1 bis 5 Glieder mehr oder weni- ger angedunkelt, selten einfarbig gelborange; Flü- geldecken gelb; Mittel- und Hinterbrust dunkel, Koxen, Trochanteren und Hinterschenkel manch- mal bis über die Mitte dunkel, Mittelschenkel an der Basis oft kurz dunkel, Vorderschenkel meistens ein- farbig gelb, Tibien und Tarsen gelb; Hinterleib stel- lenweise oft leicht angedunkelt, Rest gelblich, sel- tener ganz dunkel. 8. Kopf mit den Augen breiter als der Halsschild, Surne leicht gewölbt, Querrunzeln an der Stirnbasis kaum erkennbar, diese Stelle ist mit mehreren Punkten besetzt, die an den Seiten zahlreicher sind und bis zu dem Hinterrand der Augen und nach vorne bis zu den Fühlerwurzeln reichen, zwischen den Augen fast glatt. Fühler (Abb. 9) verhältnis- mässig kurz und kräftig, nicht ganz 2% mal so lang wie der Halsschild, breit gezahnt, einzelne Glieder ein wenig breiter als lang, 3 länger als 4. Halsschild ein wenig breiter als lang (18X16), Seiten gerundet, gegen die Basis ein wenig stärker verengt als nach vorne; Oberfläche mit deutlichen Querrunzeln; Scheibe gewölbt, an der Basis kaum angedeutet ein- gedrückt. Flügeldecken nach hinten leicht verbrei- 112 tert, ca. 3 bis 4 Tergite unbedeckt lassend, Punkte deutlich, etwas erloschen, ihr Abstand viel gròsser als ihr Durchmesser, Behaarung sehr fein, weiss- lich. Q. Fühler (Abb. 10) ein wenig kürzer, weniger breit gezahnt, sonstiger Bau und Skulptur wie beim 3. Lange: ca. 3 mm. Diese Art gehört in die Verwandtschaft von C. albipennis Wittmer, sie unterscheidet sich hauptsächlich durch die Färbung, vergleiche Be- schreibung und Bestimmungstabelle. Carphurus cf. rubroannulatus Motschulsky Carphurus rubroannulatus Motschulsky, 1859: 64. Material. — Indonesia, Sulawesi Utara: Dumoga Bone N.P., lowland forest edge, ca. 200 m, malaise trap, ii und iii.1985 (1 BM, 1 NHMB); Sulawesi Tengah: near Moro- wali, Ranu River area, iii.1980, at light, M. J. D. Brendell (1 BM, 1 NHMB). Diese Art ist von Ceylon beschrieben und von Indien, Tenasserim, Siam, Penang, Singapore, Java, Sarawak, Laos, Tonkin, Philippinen, Palawan, Su- matra, Molukken, Ceram, Buru, Borneo in der Lite- ratur erwähnt. Aehnliche Arten sind aus Sumatra (C. atromaculatus Pic), Burma (C. birmanicus Pic) beschrieben. Aus Sulawesi liegen mir 4 9 vor, die ich aufgrund ihrer Aehnlichkeit provisorisch als rubroannulatus bestimme. Carphurus basilimbatus sp.n. (Abb. 11) Material. — Holotypus: 6, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Hog’s Camp, lowland forest, 492 m, vii.1985 (BM). Paratypen: Gleicher Fundort wie Holoty- pus, vii.1985 (1 NHMB); idem Fog 5, trays 1, 5, 46, 49, 52, 69, 70, 88, 100 (Plot C), 400 m, 11.11.1985 (6 BM, 3 NHMB); idem Fog 15, trays 81, 105, 106 (Plot C), 19.vii.1985 (2 BM, 1 NHMB); idem sites 10 and 11, Tumpah Transect, 664 m, 19-25.11.1985 (1 BM). Kopf und Halsschild einfarbig gelborange; Füh- ler gelblich, letzte 3 bis 5 Glieder etwas angedun- kelt; Schildchen und grösster Teil der Unterseite mehr oder weniger aufgehellt; Flügeldecken gelb- lich, Basis zu % bis % schwarz; Beine blassgelb. 8. Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewölbt, Stirne schmal, gegen den Hinterrand der Augen breiter, ziemlich dicht punk- tiert, zwischen den Augen glatt, hier stehen nur ganz vereinzelte Punkte. Fühler (Abb. 11) ziemlich lang, wenig mehr als 3 mal länger als der Hals- schild, Glieder langlich, gegen die Spitze etwas ver- breitert, 2 breiter als 3, ein wenig nach unten ver- längert. Halsschild nur wenig breiter als lang (16X15), Seiten gerundet, kurz vor der Basis ein wenig verengt, Scheibe schwach gewölbt, vor der basis schwach eingedriickt; Oberfläche fein punk- tiert, stellenweise ein wenig gewirkt, dadurch leicht matt wirkend. Flügeldecken nach hinten leicht er- weitert, ca. 3 bis 4 Tergite unbedeckt lassend, fast unpunktiert, leicht gewirkt, Behaarung fein, greis. Letztes Tergit sehr kurz, breiter als lang. Q. Die Fühler sind kaum merklich kürzer als beim 4, Glied 2 ebenfalls nach unten verbreitert. Länge: 2.5-3 mm. Eine durch die Färbung der Flügeldecken sehr charakteristische Art, dem C. semiflavus Wittmer sehr ähnlich und die ausserdem durch das verdickte Fühlerglied 2 auffällig ist, das bei semiflavus nicht verdickt ist. Carphurus testaceolimbatus sp.n. (Abb. 12) Material. — Holotypus: 6, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Fog 3, tray 5 (Plot B), 315 m, 8.11.1985 (BM). Paratypen: idem Fog 12, tray 29 (Plot A), 280 m, 5.v.1985 (1 BM); idem Fog 26, tray 89 (Plot A), 230 m, 2.x11.1985 (1 NHMB). 8. Kopf, Schildchen und Abdomen schwarz- braun; Fühler schwarz, erste 4 bis 5 Glieder gelb; Halsschild schwarzbraun, basales Drittel bis Viertel gelbbraun; Flügeldecken schwarzbraun, mit einem gelblichweissen Querband vor der Mitte, ungefahr so breit wie der Abstand zur Basis; Beine gelblich. Kopf mit den Augen breiter als der Halsschild, Stirne ziemlich flach, Schläfen runzlig gewirkt oder mit Querrunzeln, die manchmal bis zur Stirnbasis zu erkennen sind, Rest des Kopfes glatt mit ganz vereinzelten Punkten. Fühler (Abb. 12) 3.5 mal so lang wie der Halsschild, Glieder 3 bis 10 gezahnt, unter sich ungefähr gleich lang. Halsschild unge- fähr so lang wie breit, Seiten leicht gerundet, von der Mitte zur Basis stärker verengt als nach vorne; Scheibe gewölbt, vor dem Basalrand flach, jedoch praktisch nicht eingedriickt; Oberfläche glatt, glän- zend, wie der Kopf punktiert. Fliigeldecken fast parallel, ca. 4 Tergite unbedeckt lassend, glatt glin- zend, ein paar Haarpunkte kaum sichtbar, Behaa- rung spärlich, schräg aufstehende kleine Borsten, ausserdem 3 bis 4 längere Borsten auf jeder Decke. Länge: 2-2.3 mm. Deise Art unterscheidet sich von C. /yali sp.n. durch den mit einem gelblichen Saum versehenen Halsschild und dem gelblichweissen Querband auf den Flügeldecken; die Fühler sind bei testaceolimbatus ein wenig stärker gezahnt, die Flü- geldecken praktisch glatt, bei C. /yali sp.n. deutlich punktiert. Carphurus brendelli sp.n. (Abb. 13-14) Material. — Holotypus: 8, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Fog 11, tray 24 (Plot A), 10.111. 1985 WITTMER: Malachtidae auf Sulawesi (BM). Paratypen: idem Fog 12, tray 112 (Plot A), 5.v.1985 (1 BM); idem Toraut alluv. forest, 1-2.11.1985 (1 NHMB); idem Plot A, ca. 200 m, lowland forest, 6-13.111.1985 (1 BM); idem Edward’s Camp, lowland forest, 664 m, 26.iv- 7.vi.1985 und x.1985 (1 BM, 1 NHMB). Kopf rot, Wangen und manchmal die Kopfbasis schmal schwarz; Fühler schwarz, erste 4 bis 5 Glie- der gelb; Halsschild, Schildchen, Unterseite und Flügeldecken schwarz, letztere mit einem ver- schwommenen bis deutlichen, weisslichen, mehr oder weniger langen Längsflecken, der weder die Naht noch die Seiten berührt; Beine gelb, beim einzigen vorliegenden @ sind die Hinterschenkel fast bis zu den Knien schwarz, die Hintertibien angehaucht dunkel; bei den 9 sind oft auch die Mittelschenkel fast bis oben, seltener die Vorder- schenkel an der Basis kurz schwarz. 8. Kopf mit den Augen ein wenig breiter als der Halsschild, Stirne leicht gewölbt, Schläfen an der Basis mit Querrunzeln, an der Stirnbasis meistens durch den Halsschildvorderrand verdeckt, davor wenige gröbere Punkte, Rest des Kopfes praktisch glatt. Fühler (Abb. 13) sehr lang, 4 mal so lang wie der Halsschild, Glieder 5 bis 10 zur Spitze regelmäs- sig leicht verbreitert, 4 ein wenig länger als 5, zur Spitze stärker verbreitert als dieses, breit ausgeran- det, 3 ein wenig kiirzer als 4, 2 nach unten leicht gerundet erweitert, fast so breit wie 1. Halsschild ein wenig langer als breit (24X22), Seiten nur schwach gerundet, gegen die Basis verengt; Scheibe in der Mitte ein wenig stärker gewölbt, vor dem Basalrand deutlich quer eingedriickt; Oberfläche glatt, ein paar Haarpunkte sichtbar. Flügeldecken nach hinten leicht erweitert, ca. 5 Tergite unbedeckt lassend, Punktierung spärlich, fast erloschen. Glied 1 der Vordertarsen verlängert und mit einem lan- gen, an der Spitze gerundeten Kamm versehen. ©. Fühler (Abb. 14) einfach, wesentlich kürzer als beim Ó. Linge: 3.5-4.5 mm. Färbung und Körperform wie bei C. rouyeri Pic, verschieden durch den Bau des Fühlers beim Ó, dessen Glied 4 breit ausgerandet ist. Carphurus rufonotaticeps Pic Carphurus rufonotaticeps Pic, 1917: 9. Carphurus tondanus Champion, 1923: 35, syn.n. Material. — Indonesia, Sulawesi Tengah: Ranu River area, 27.i-20.iv..1980, sweeping clearings, M. J. D. Bren- dell, BM 1980-280 (zahlreiche Exemplare BM, NHMB); idem near Kolonodale, Gililana Village, 01°55’S- 121°22’E, 7-8.ii.1980, M. J. D. Brendell (6 BM); Sulawesi Utara: Dumoga Bone N.P., Mogogompa, i, iii und x.1985 (14 BM, 14 NHMB). Der Holotypus von Champion (Q) ist ca. 10% grösser als die meisten mir vorliegenden Exem- plare. Der Bau der Fiihler, Punktierung von Hals- schild und Fliigeldecken stimmmt mit dem Paralec- 115 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 totypus ® von rufonotaticeps Pic überein, so dass ich tondanus Champion als Synonym betrachte. Carphurus storki sp.n. (Abb. 15) Material. — Holotypus: @, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Rothamsted light trap, site 2, 220 m, 11.1985 (BM). Paratypen: idem Edward’s Camp, lowland forest, 664 m, at light, x.1985 (1 NHMB); idem Fog 1, tray 56 (Plot A), 5.ii.1985 (1 BM). Einfarbig schwarz, nur die Fiihlerglieder 1 bis 5 aufgehellt, 1 und 5 nur auf der Unterseite. 8. Kopf mit den Augen so breit wie der Hals- schild, Stirne leicht gewölbt, an der Basis einige = Nn III / | 5 feine Querrunzeln, davor ein paar Punkte, zwischen den Augen und nach vorne glatt, spärliche Punkte. Fühler (Abb. 15) knapp 3 mal so lang wie der Halsschild, Glieder 3 bis 10 gegen die Spitze deut- lich verbreitert, ein wenig länger als breit, 2 schwach nach unten erweitert, kaum merklich schmäler als 3. Halsschild breiter als lang (19X17), Seiten nach vorne schwach verengt, kurz vor der Basis gegen diese verengt; Scheibe leicht gewölbt, vor der Basis quer eingedrückt, an dieser Stelle seitlich mit ein paar kurzen Querrunzeln, Rest glatt, wenige Haarpunkte sichtbar. Flügeldecken nach hinten ein wenig erweitert, 3 bis 3% Tergite unbedeckt lassend, Punkte kaum angedeutet, erlo- schen gewirkt, nicht glatt. Q. Fühler um % kürzer als beim @. Länge: 2.5-3 mm. Die neue Art ist nahe mit C. brunneiventris (Pic) verwandt. Obwohl die Picsche Art nach 1 © be- schrieben wurde, ist C. storki leicht anhand des Halsschildes zu unterscheiden, dessen Querein- druck vor der Basis deutlich tiefer ist als bei brunneiventris. Ausserdem ist er bei brunnet- A mm 17 en Abb. 13-17 — Fühler von Carphurus. 13-14, C. brendelli sp.n. (13, &. 14, Q); 15, C. storki sp.n. 8; 16, C. lyali sp.n. 6; 17, Fühler von Telocarphurus nebulosus sp.n. &. 114 ventris vollständig glatt, bei stork: zeigt er an den Seiten ein paar gröbere, fast erloschene Punkte und 2 bis 3 deutliche kurze Querrunzeln an den Seiten basal vor dem Quereindruck und eine grössere An- zahl dahinter. Der Kopf zwischen den Augen ist bei storki deutlicher punktiert als bei brunneiventris. Carphurus lyali sp.n. (Abb. 16) Material. — Holotypus: @, Indonesia, Sulawesi Utara: Gng. Ambang ER. near Kotamobagu, Fog 7, 1200 m, 18.11.1985 (BM). 8. Oberfläche und Abdomen einfarbig schwarz bis schwarzbraun; Fühler schwarz, Glied gelb, 2 leicht aufgehellt. Beine braun. Kopf mit den Augen viel breiter als der Hals- schild, Stirne ziemlich flach, Stirnbasis gegen die Schläfen und an denselben mit deutlichen Querrun- zeln, diese dringen nach vorne neben den Augen bis fast über die Fühlerwurzeln; zwischen den Augen ganz wenige Punkte, vor der Stirnbasis gröbere und zahlreichere Punkte. Fiihler (Abb. 16) fast 4 mal länger als der Halsschild, Glieder 3 bis 10 schwach gezahnt, 2 ungefähr so breit wie 3. Halsschild ein wenig breiter als lang (13X12), Seiten wenig gerun- det, kurz vor der Basis ein wenig verengt; Scheibe gewölbt, vor der Basis kurz, flach eingedriickt; Oberfläche glatt mit wenigen Haarpunkten. Flügel- decken ca. 4 Tergite unbedeckt lassend, ziemlich grob und dicht, teils etwas erloschen punktiert. Länge: 2.7 mm. Neben C. testaceolimbatus sp.n. zu stellen. Telocarphurus Wittmer, 1939 Telocarphurus nebulosus sp.n. (Abb. 17) Material. — Holotypus: g, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Fog 5, tray 9 (Plot C), 400 m, 11.11.1985 (BM). 8. Kopf, Halsschild, Schildchen und grösster Teil der Unterseite schwarz; Fühler schwarz, 2 erste Glieder gelb; Flügeldecken schwarz, basale °/; ver- schwommen gelblichbraun; Beine braun bis schwärzlich. Kopf mit den Augen breiter als der Halsschild, Stirnbasis sehr schmal und Schläfen gewirkt, Ober- fläche glatt mit ein paar Punkten. Fühler (Abb. 17) ca. 3% mal so lang wie der Halsschild, Glieder 3 bis 10 zur Spitze leicht verbreitert, unter sich ungefähr von gleicher Länge, erst bei 10 deutlich kürzer als 3. Halsschild nur wenig breiter als lang (12X11), Seiten nur schwach gerundet, Oberfläche glatt mit wenigen Punkten, ausser je einer langen Borste in den Basalecken ist die Scheibe mit einer Anzahl WITTMER: Malachiidae auf Sulawesi kürzerer Haare besetzt. Flügeldecken verkürzt, ca. 4 Tergite unbedeckt lassend, fast glatt, leicht durch- sichtig, zerstreut, ungefihr wie der Halsschild be- haart. Linge: 2 mm. Noch ein wenig kleiner und zarter gebaut als T. strigilatus Wittmer, von diesem durch den glat- ten Halsschild und die viel kürzeren Fühler leicht zu unterscheiden. Paracarphurus Wittmer, 1953 Paracarphurus sulawesiensis sp.n. (Abb. 18) Material. — Holotypus: 8, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Clarke Camp, lower montane forest, 1140 m, iv.1985 (BM). Paratypen: Gleicher Fundort wie Holotypus, v.1985 (1 BM); idem Edward's Camp, lower montane forest, 664 m, x.1985 (1 NHMB). 8. Braun, letzte 4 bis 5 Fühlerglieder ganz schwarz, manchmal auch die Spitzen ein paar wei- terer Glieder angedunkelt, Hinterschenkel fast bis zu den Knien und manchmal auch die Basis der Mittelschenkel kurz schwarz; Unterseite schwarz, letzte 3 Sternite orange, Tergite orange, ausgenom- men das viertletzte, das schwarz ist. Kopf mit den Augen kaum merklich breiter als der Halsschild, Stirne nur schwach gewölbt, über jeder Fühlerwurzel ein flacher Eindruck, Oberfläche fast glatt, Stirnbasis mit ein paar Querrunzeln. Füh- ler die Schulterbeulen ein wenig überragend, Glie- der 3 bis 10 gezahnt, 3 ein wenig länger als 2. Halsschild ein wenig länger als breit, Seiten vor der Mitte am breitesten, hier gerundet, dann zur Basis leicht verschmälert, Basis vor dem Basalrand leicht aufgewölbt; Oberfläche glatt, feine zerstreute Haar- punkte sichtbar. Fliigeldecken verkiirzt, etwas mehr als doppelt so lang wie der Halsschild, jede Spitze breit gerundet, fast in der Mitte, etwas vor der Spitze befindet sich ein runder Eindruck mit leicht erhöhtem Rand, vor dem Eindruck eine Anzahl sehr lange weissliche, nach hinten gerichtete Haare (Abb.18), die an der Spitze leicht verbreitert sind; Punktierung mässig grob, Punkte teils zusammen- fliessend, Zwischenräume glatt. Tarsalkamm kurz, so lang wie das Tarsenglied 1. Länge: 5 mm. Die neue Art ist am nächsten mit P. neobrit- tanicus Wittmer verwandt, sie hat ähnlich gerun- dete Spitzen der Flügeldecken, die jedoch mit einem grösseren Eindruck versehen sind, der weiter hin- ten als bei neobrittanicus liegt. Bei neobrittanicus fehlen die langen Haare hinter dem Eindruck, dafiir befindet sich hier eine kurze breite Borste und zwi- schen dem Eindruck und dem Hinterrand eine grosse, fast runde, fast glatte Fläche, die bei sulawesiensis fehlt. 115 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Falsolaius Pic, 1917 Luzonotroglops Pic, 1924 syn.n. Der Vergleich des Holotypus von Luzonotrog- lops carinatus Pic (spec. typ. von Luzonotroglops) mit dem Holotypus von Falsolaius curtipennis Pic (spec. typ. von Falsolaius), beide im MP, hat obige Synonymie ergeben. Die beiden Arten F. carinatus (Pic) und E curtipennis Pic bleiben bestehen. Alle seit 1924 als Luzonotroglops beschriebenen Arten sind zu Falsolaius zu stellen. Die Gattung Falsolaius Pic ist nahe mit der Gat- tung Neocarphurus Lea verwandt, die hauptsäch- lich über Australien verbreitet ist, sie unterscheidet sich von letzterer nur durch den einfachen Kopf, dem Gruben oder sonstige Bildungen fehlen. Falsolaius sulawesiensis sp.n. (Abb. 19, 21) Material. — Holotypus: 6, Indonesia, Sulawesi Utara: Gng. Ambang F.R. near Kotamobagu, lower montane forest, ca. 1750 m, 18-24.v.1985, yellow pan trap (BM). mm 21 22 25 Paratypen: Gleicher Fundort wie Holotypus, 18-24.v.1985 (1 BM - @ in copula mit Holotypus — 2 NHMB); Gng. Muajat, summit area, ca. 1780 m, vi.1985, malaise trap (1 BM). Kopf und Halsschild braun, letzterer an der ein- geschniirten Stelle bis zur Basis meistens mehr oder weniger aufgehellt; Fühler braun, die letzten 2 oder 3 Glieder angedunkelt; Schildchen und Flügel- decken braun, letztere oft mit einem schwach ange- dunkelten Flecken auf jeder Schulterbeule, der stark reduziert sein kann und nur noch neben dem Schild- chen sichtbar ist, bei den Exemplaren mit gut ent- wickelten Schulterflecken ist auch eine dunkle Mit- telbinde vorhanden, die die Seiten und die Naht nicht immer erreicht und je eine quere Makel an den Spitzen; Abdomen dunkelbraun, letztes Tergit heller; Beine braun, Schienen meistens ein wenig angedunkelt, manchmal auch die Oberseite der Schenkel. 8. Kopf mit den Augen breiter als der Halsschild, Stirne schwach gewölbt, Basis kurz fein längsgestri- chelt, Wangen ebenfalls gestrichelt, Rest des Kop- fes gröstenteils chagriniert, zwischen den Augen 23 19 20 24 Abb. 18-25. — 18, Spitze der Flügeldecke von Paracarphurus sulawesiensis sp.n. &; 19-20, Fühler von Falsolaius (19, E sulawesiensis sp.n. Ô, 20, E semicostatus sp.n. 8); 21-22, Letztes Tergit von Falsolaius (21, F. sulawesiensis sp.n. à; 22, E semicostatus sp.n. @); 23-25: Dromanthomorphus cyaneus Pic &. (23, Vordertibie von oben, daneben im Profil; 24, Mitteltibie; 25 letztes Tergit). 116 eine glattere Stelle, ein paar lange aufrechtstehende Borsten vorhanden. Fühler (Abb. 19) fast 3 mal so lang wie der Halsschild, Glieder 3 bis 10 zur Spitze nur ganz wenig verbreitert, 4 länger als 3. Hals- schild länger als breit (20X17), Seiten stark gerun- det verengt, basales Drittel eingeschnürt und dann zur Basis wieder ein wenig verbreitert, Seitenkante vorne gegen den breiten und stark gewölbten Teil am deutlichsten, gegen die Basis verschwindend, der ganze eingeschnürte Teil ist körnig gewirkt, matt, der vordere Teil glatt, vereinzelte Haarbor- sten vorhanden. Flügeldecken nach hinten wenig verbreitert, 4 bis 5 Tergite unbedeckt lassend, Schultern stark vorstehend mit einer Längskante, die nach den Schulterbeulen verschwindet, Spitze jeder Decke schwach beulenartig aufgewölbt, Ober- fläche körnig gewirkt, matt, mit zahlreicheren schwarzen Borsten, besetzt als Kopf und Hals- schild. Letztes Tergit (Abb. 21) ziemlich vorste- hend, Spitze breit gerundet. Q. Wie das 8 gebaut, sogar die Fühler sind kaum merklich kürzer. Die neue Art gehört zu den Arten mit glattem Halsschild, wenigstens dem vorderen Teil und an- gedeuteter Beule vor der Spitze der Flügeldecken. Sie unterscheidet sich von F. unituberculatus (W itt- mer) durch hellere Gestalt, den an der basalen, verengten Stelle des Halsschilds, der fein gewirkt, matt ist und die nur angedeutete beulenartige Ver- dickung vor der Spitze der Flügeldecken, um nur die wichtigsten Unterscheidungsmerkmale aufzuzäh- len. Falsolaius semicostatus sp.n. (Abb. 20, 22) Material. — Holotypus: 8, Indonesia, Sulawesi Utara: Dumoga Bone N.P., Fog 5, trays 13, 14, 18, 26, 32, 54, 97 (Plot C), 11.11.1985 (BM). Paratypen: Gleicher Fundort wie Holotypus, 11.11.1985 (4 BM, 3 NHMB); idem Fog 8, trays 7, 11, 13, 15, 16, 18, 19, 20, 22, Mangrove, 3 km S. of Labuanika, 0°52’N-23°57’E, 25.11.1985 (7 BM, 3 NHMB); idem Plot C, ca. 400 m, lowland forest, malaise trap up trees, 18-25.1x.1985 (1 NHMB); idem Plot B, ca. 300 m, lowland forest, malaise trap up trees, 6-13.iii.1985 (1 BM); idem Edward's Camp, lowland forest, 664 m, malaise trap, 26.iv-7.vi.1985 (1 BM). Kopf dunkelbraun, vorderer Teil mehr oder we- niger aufgehellt; Halsschild dunkelbraun, seltener hellbraun, der eingeschnürte Teil an der Basis auf- gehellt (gelblich); Fühler gelb, letzte 2 bis 5 Glieder angedunkelt; Schildchen und Flügeldecken braun, dunkelbraun oder seltener hellbraun, variabel auf- gehellt, z.B. mit einer queren Aufhellung vor der Mitte, die an der Naht unterbrochen sein kann und einer breiteren dahinter, oder die hintere ist schmä- ler als die vordere, bei dunkelbraunen Exemplaren sind Aufhellungen kaum sichtbar oder nur ganz schwach, manchmal nur hinten sichtbar; Abdomen WITTMER: Malachiidae auf Sulawesi dunkelbraun, die beiden letzten Tergite oder nur das letzte kann hell sein; Beine dunkelbraun, oder heller braun bis gelb, dann können die Schenkel, manchmal auch die Schienen einen dunkieren Längswisch aufweisen. 8. Kopf mit den Augen ein wenig breiter als der Halsschild, Stirne leicht gewölbt, Oberfläche cha- griniert (64X). Fühler (Abb. 20) fast doppelt so lang wie der Halsschild, Glieder 3 bis 10 zur Spitze nur ganz wenig verbreitert, 4 ein wenig länger als 3. Halsschild länger als breit (20X15), Seiten ge- rundet verengt, basales Drittel eingeschnürt und dann zur Basis wieder ein wenig verbreitert, Seiten- kante auf dem vorderen Teil der Einschnürung kurz oder nur angedeutet; Oberfläche am eingeschnür- ten Teil fein chagriniert, Rest glatt. Flügeldecken fast parallel, Schulterbeulen vorstehend, mit einer Längskante, die ungefähr nach % verschwindet, vor jeder Spitze ein Querwulst; Oberfläche fast glatt, Behaarung fein gelblich nicht dicht, dazwischen ganz vereinzelte, kurze aufstehende dunkle Bor- sten. Letztes Tergit (Abb. 22) sehr kurz und breit, Spitze ausgerandet. Q. Fühler ein wenig kürzer als beim @, sonst wie dieses gebaut. Länge: 2-2.3 mm. Nahe mit F sulawesiensis Wittmer verwandt, Gestalt kleiner, die Seitenkanten an der Einschnü- rung auf dem Halsschild sind weniger ausgeprägt, die Querwulste an den Spitzen der Flügeldecken sind grösser und das letzte Tergit ist viel kürzer, Spitze ausgerandet. Dromanthomorphus Pic, 1921 Flabellapalochrus Pic, 1923 syn. n. Die Gattung Dromanthomorphus wurde von Pic nach 1 ® aus Nord-Celebes, Toli-Toli, Nov.-Dez. 1895, H. Fruhstorfer, beschrieben, Holotypus im MP. Im Material des BM fand sich 1 @, das zu D. cyaneus Pic gehört. Pic hat bei der Beschreibung der Gattung Flabellapalochrus nach 1 & übersehen, dass er diese Gattung bereits nach 1 ® unter dem Namen Dromanthomorphus beschrieben hatte, so- dass erstere Gattung in Synonymie verfällt. Alle als Flabellapalochrus beschriebenen Arten sind in die Gattung Dromanthomorphus Pic zu stellen. Dromanthomorphus cyaneus Pic (Abb. 23-25) Dromanthomorphus cyaneus Pic, 1921: 7. Material. — Indonesia, Sulawesi Utara: Dumoga-Bone N.P., Plot B. ca. 300 m, lowland forest, malaise trap, ix.1985 (1 & BM). Da das & noch nicht beschrieben wurde, folgt hier dessen Beschreibung: 117 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 6. Kopf blau, Vorderkopf bis kurz über den Füh- lerwurzeln, Oberlippen und Maxillarpalpen gelb; Fühler gelb, die kammartigen Fortsätze der Glieder 4 bis 6 schwarz, 7 bis 11 ganz schwarz; Halsschild und Schildchen blau; Flügeldecken weisslichgelb, Basis und Spitzen ein wenig breiter, Seiten und Naht schmal blau; Abdomen gelblich, letztes Tergit braun, Spitze schwach, schmal angedunkelt; die 4 Vorderbeine gelb, Hinterbeine dunkel, an den Knien leicht aufgehellt. Kopf mit den Augen schmiler als der Halsschild an der Basis, Stirne leicht, regelmässig gewölbt, zwischen den Augen eine kleine Grube, Oberfläche ziemlich grob punktiert. Fühler von Glied 4 an lang gekämmt, 3 stark nach innen gerundet erweitert. Halsschild breiter als lang, Seiten gerundet, ein wenig nach vorne verengt, Scheibe gewölbt; Ober- fläche unregelmässig, wenig dicht punktiert. Flü- geldecken nach hinten leicht erweitert, fast ganz erloschen punktiert, Behaarung doppelt, greis, kurz auf der hellen Fläche, abstehende Borsten dunkel. Letztes Tergit (Abb. 25) stärker sklerotisiert als die nachfolgenden, lang, Spitze vorstehend, an den Sei- ten ausgeschnitten. Vordertibien (Abb. 23) von oben gesehen, vor der Spitze sehr schmal, wie zu- sammengedriickt, im Profil gesehen (Abb. 23) nach oben gerundet verbreitert; Mitteltibien (Abb. 24) stark verbreitert, von oben gesehen ohne Aushöh- lungen. Länge: ca. 4.3 mm. Dromanthomorphus ranuensis sp.n. Material. — Holotypus: 8, Indonesia, Sulawesi Tengah: near Morowali, Ranu River area, 27.i-20.iv.1980, Vert. series 30 m actinic code, 13.111.1980, also at flight, M. J. D. Brendell (BM); Paratypen: Gleicher Fundort wie Holoty- pus (17 BM, 10 NHMB). Kopf, Halsschild, Schildchen und Flügeldecken schwarz mit mehr oder weniger deutlichem blauem Metallschimmer, auf letzteren manchmal eine längliche, schlecht begrenzte Aufhellung; Fühler schwarz, erste 6 Glieder gelb, bei 4 bis 6 sind die Kamme zunehmend angedunkelt; Beine schwarz, Vordertarsen gelblich; Abdomen schwarz, die letz- ten 2 bis 3 Sternite gelblich. 8. Kopf mit den halbkugelförmigen Augen brei- ter als der Halsschild; Stirne schwach gewölbt; Oberfläche dicht punktiert. Fühler um ca. ‘4 kürzer als die Flügeldecken, von Glied 4 an in zunehmen- der Lange gekämmt, 3 länglich, zur Spitze verbrei- tert, diese stumpf, ein wenig gerundet. Halsschild breiter als lang, Seiten gerundet, nach vorne etwas verengt, Vorderecken etwas stärker gerundet als die Basalecken; Oberfläche uneben, weniger deutlich als der Kopf punktiert. Flügeldecken langgezogen, nach hinten schwach verbreitert; Oberfläche teils schwach runzlig, teils fast matt. Alle Beine ohne Auszeichnungen. 118 Q. Augen normal, nicht vergrössert, Kopf mit den Augen praktisch gleich breit wie der Hals- schild. Fühler um ‘4 kürzer als beim @, stumpf gezahnt. Länge: ca. 3.3 mm. Neben D. saigonensis (Pic) und D. impressitho- rax (Pic) zu stellen, die beide ebenfalls einfache Beine besitzen (8). Die neue Art ist durchwegs grösser als die beiden erwähnten Arten von Pic, die nur 2.5 und 2.8 mm messen. Laius Guérin, 1830 Bestimmungstabelle für die auf Sulawesi vorkommenden Arten 1. Kleine Arten, weniger als 3 mm messend; Flü- geldecken mit 2 oder 4 kleinen weissen Flecken — Grosse Art, 4 mm messend; Flügeldecken ein- farbig blauviolett ........ L. cyaneus Guérin 2. Wangen einfach, ohne Gruben; Flügeldecken mit 4aweissenthlecken) see CSO 3 — Wangen mit einer tiefen Grube, die sich bis zum Hinterrand der Augen erstreckt; Flügeldecken mit 2 weissen Flecken vor der Mitte. Fühler- gliederail'-3; Abby I ER SARETE AE aen miettes Har L. fossigerus sp.n. 3. Seiten des Halsschilds ungefähr in der Mitte mit einem kräftigen Zahn ……. sanar: 4 — Seiten des Halsschilds einfach, ohne Zahn. Füh- lerglieder 1-3, Abb. 27 ..... L. ranuensis sp.n. 4. Vorderkopf breit, über den Fühlerwurzeln leicht abgesetzt aufstehend, Wangen flach; Fühlerglied 3 kürzer, oben nicht längseinge- drucke@Abbs26)i aa EEE L. ribbet Pic — Vorderkopf nach vorne verschmälert, über den Fühlerwurzeln nicht erhöht, Wangen ausge- höhlt; Fühlerglied 3 länger, oben längseinge- drucke(Albbs28) isan L. denticollis sp.n. Laius fossigerus sp.n. (Abb. 29) Material. — Holotypus: 8, Indonesia, Sulawesi Tengah: near Morawali, Ranu River area, 27.1-20.iv.1980, M. J. D. Brendell (BM). 8. Kopf, Halsschild, Schildchen und Flügel- decken schwarz, letztere mit 2 weissen ovalen, iso- lierten Querflecken auf der vorderen Hälfte; Fiihler schwarz, Glieder 2 bis 7 ganz, 8 basale Hälfte gelb; Beine schwarz, alle Tarsen leicht aufgehellt. Kopf mit den stark vorstehenden Augen breiter als der Halsschild, Schläfen neben den Augen tief ausgehöhlt, zwischen dem Hinterrand des Auges und der Aushöhlung ragt eine schmale, wenig hohe Leiste auf (von vorne gesehen gut sichtbar), Längs- leiste zwischen den Augen am deutlichsten sicht- bar, Wangen breit, flach; Oberfläche fein gewirkt, matt. Fühler (Abb. 29) die Schulterbeulen nur we- nig überragend, Glied 1 gegen die Spitze verbrei- tert, 2 kaum sichtbar, 3 länglichoval, vor der Spitze ein ovaler Eindruck, vor der Basis ein langer, schmaler, schräg nach aussen gerichteter Fortsatz. Halsschild ein wenig länger als breit, Seiten leicht gerundet, Scheibe gewölbt, an der Basis quer einge- drückt; Oberfläche fein chagriniert, matt. Flügel- decken nach hinten leicht erweitert, ziemlich dicht gewirkt, matt. Länge: 2.5 mm. Dies ist die einzige mir bekannte schwarze Lazus- Art mit weissen Flecken auf den Flügeldecken, de- ren Wangen tief ausgehöhlt sind. Laius ranuensis n.sp. (Abb. 27) Material. — Holotypus: 8, Indonesia, Sulawesi Tengah: near Morawali, Ranu River area, vert. series 20 m actinic, 14.11.1980, M. J. D. Brendell (BM). 6. Kopf schwärzlich, der vordere Teil der Wan- gen und der Vorderstirne gelb; Fühler gelb bis gelb- braun, die letzten 4 Glieder leicht angedunkelt; Halsschild, Schildchen und Fliigeldecken schwärz- lich, die letzteren mit 4 queren, weissen Flecken, die vorderen erreichen die Seiten, die hinteren errei- chen sie nicht ganz; Beine dunkel, die Spitze der Vorderschenkel, die Vordertibien und alle Tarsen gelb. Kopf mit den Augen ungefähr so breit wie der Halsschild, Stirne zuerst leicht gewölbt, dann bis zur Vorderstirne fast glatt, Lingsleiste vorhanden, 26 Di WITTMER: Malachtidae auf Sulawesi an der Kopfbasis beginnend, nicht sehr deutlich sichtbar, Wangen sehr breit; Oberfläche gewirkt, matt. Fühler (Abb. 27) die Schulterbeulen ein wenig überragend, Glied 1 keulenförmig, gegen die Spitze nach aussen erweitert, 2 versteckt, 3 länglichoval, an der Basis innen in eine kleine stumpfe, behaarte Spitze ausgezogen, Oberfläche längseingedrückt. Halsschild kaum merklich breiter als lang, Seiten fast in gerader Linie gegen die Basis verengt; Scheibe gerundet, vor der Basis schwach quer ein- gedrückt, Oberfläche wie der Kopf gewirkt. Flügel- decke nach hinten nur wenig erweitert, wie der Halsschild gewirkt. Länge: ca. 2.5 mm. Neben L. denticollis Wittmer zu stellen, ver- schieden durch die fehlenden Dorne an den Seiten des Halsschilds und die anders geformten Fühler. Laius ribbei Pic (Abb. 26) Laius ribbei Pic, 1910: 84. Der Holotypus im MP trägt folgende Etikette: S. Celebes, Pangie, C. Ribbe, 1882. Die Abb. 26 zeigt die Fühlerglieder 1 bis 3. Laius denticollis n.sp. (Abb. 28) Material. — Holotypus: @, Indonesia, Sulawesi Tengah: near Morawali, Ranu River area, 27.1-20.1v.1980, M. J. D. Brendell (BM). Paratypen: Gleicher Fundort wie Holoty- pus (8 BM, 4 NHMB); Sulawesi Utara: Dumoga-Bone N.P., lowland forest, ca. 200 m, 11.1985. mm 28 29 1 Abb. 26-29. — Fühlerglieder 1 bis 3 von Latus @. — 26, L. ribbei Pic; 27, L. ranuensis sp.n.; 28, L. denticollis sp.n.; 29, L. fossigerus sp.n. 119 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 6. Kopf, Halsschild, Schildchen und Flügel- decken schwarz, letztere mit 4 weissen, isolierten, fast runden Flecken, die 2 vorderen vor der Mitte, die 2 hinteren hinter der Mitte; Fühler schwärzlich, Glied 1 auf der Unterseite, 3 fast vollständig und die folgenden 2 bis 4 auf der Innenseite bräunlich; Beine schwärzlich, Tarsen ganz leicht aufgehellt. Kopf mit den Augen kaum merklich schmäler als der Halsschild an den seitlichen Zähnen, Stirne gewölbt, eine feine Längsleiste beginnt an der Stirnbasis und erstreckt sich bis ungefähr zur Mitte, die Stirne ist vorne neben den Augen leicht gerun- det aufgewölbt und dazwischen flach eingedrückt, der grösste Teil der Wangen ist breit ausgehöhlt, so dass die Vorderstirne sehr schmal ist; Oberfläche gewirkt, matt. Fühler (Abb. 28) die Schultern ein wenig überragend, Glied 1 keulenförmig gegen die Spitze nach aussen erweitert, 2 fast ganz versteckt, 3 fast länglichoval, in der Mitte längseingedrückt, an der Basis mit einem kurzen, leicht aufwärts ge- richteten Zahn. Halsschild breiter als lang, Seiten zur Basis verengt, jederseits fast in der Mitte ein kräftiger Zahn, Oberfläche wie der Kopf gewirkt. Flügeldecken nach hinten nur wenig erweitert, wie der Halsschild gewirkt, matt. 120 Q. Wie das 8 gefärbt. Kopf einfach; Fühlerglied 1 zur Spitze verbreitert, jedoch nicht nach innen erweitert, 3 so lang wie beim J, oben nicht einge- drückt, Zahn an den Halsschildseiten kleiner. Linge: 2.5-2.8 mm. Neben L. celebensis Pic und L. dentatithorax Pic zu stellen, die beide verschieden gebauten Kopf und Fiihler besitzen. LITERATUR Champion, G. C., 1923. A revision of the Malayan and Indian species of the melyrid subfamily Carphurinae represented in the Hope Collection at Oxford and in the British Museum in London (Coleoptera). - Annals and Magazine of Natural History 9(12): 1-54. Motschulsky, V. de, 1859. Etudes Entomologiques 8: 1-64. Pic, M. 1910. Sur les “Laius” Guer. à quatre macules blanches élytrales. - Echange 26: 83-84. Pic, M., 1917. Descriptions abrégées diverses. - Mélanges exotico-entomologiques 25: 1-24. Pic. M., 1924. Coléoptères nouveaux des Philippines. — Bulletin de la Société Entomologique de France: 1924: 230-231. Received: 2 January 1190 Accepted: 30 April 1990 Tijdschrift voor Entomologie Volume 133, no. I | Editorial Articles 3 A. Asquith & J. D. Lattin Nabicula (Limnonabis) propinqua (Reuter) (Heteroptera: Nabidae): dimorphism, phylogenetic relationships and biogeography. 17 A. Bedos & L. Deharveng New species of Superodontella Stach (Collembola: Odontellidae) from Thailand. 27 ). Belle Progomphus nigellus and Phyllocycla hamata, two new dragonflies from Brazil (Odonata: Gomphidae). 3l K. A. Monk & R. K. Butlin A biogeographic account of the grasshoppers (Orthoptera: Acridoidea) of Sulawesi, Indonesia. 39 T. Pape Two new species of Sarcophaga Meigen from Madeira and mainland Portugal (Diptera: Sarcophagidae). 43 T. Pape Revisionary notes on American Sarcophaginae (Diptera: Sarcophagidae). 75 V. Sruoga Seven new species of Elachistidae (Lepidoptera) from the USSR. 85 Br. Theowald & P. Oosterbroek Zur Zoogeographie der Westpaläarktischen Tipuliden. IX. Die Tipuliden des Vorderen Orients. 97 J. van Tol Key to the Malesian species of Leptogomphus Selys, with the description of a new species from Sabah (Odonata, Gomphidae). Descriptions and records of Malesian Odonata, |. 107 W. Wittmer Die Familie Malachiidae (Coleoptera) auf Sulawesi. 43. Beitrag zur Kenntnis der indo-malaiischen Fauna. Book review 96 R. Johansson et al., The Nepticulidae and Opostegidae (Lepidoptera) of North West Europa [R. T. A. Schouten] © Nederlandse Entomologische Vereniging, Amsterdam Published 31 July 1990 ISSN 0040-7496 Volume 133, no. 2, 1990 … ae ke i ISSN 0040-749 Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 . Published by the Nederlandse Entomologische Vereniging Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. |. van Nieukerken (elected 1986) and |. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address c/o National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Correspondence regarding membership of the society, subscriptions and possibilities for exchange of this journal should be addressed to: Nederlandse Entomologische Vereniging c/o Instituut voor Taxonomische Zoölogie Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands Subscription price per volume Hfl. 300.- (postage included). Special rate for members of the society. Please enquire. Instructions to authors May be obtained from the editors. Graphic design Ontwerpers B.V., Aad Derwort, ’s-Gravenhage BEREND AUKEMA MCZ LIBRARY Biological Station, Wijster (Communication no. 421) TAXONOMY, LIFE HISTORY AND DISTRIBUTION JAN (ee 1991 OESREIREE CLOSELY INELALED SPECIES OEZTEIE RVARD A a GENUS CALATHUS (COLEOPTERA: CARABIDAE) ito, ERSITY Aukema, B., 1990. Taxonomy, life history and distribution of three closely related species of the genus Calathus (Coleoptera: Carabidae). — Tijdschrift voor Entomologie 133: 121- 141, figs. 1-18, tabs. 1-15 [ISSN 0040-7496]. Published 14 December 1990. After cross-breeding experiments between Calathus melanocephalus, C. mollis mollis and C. mollis cinctus and analysis of morphological data of these taxa, Calathus cinctus is established as a good species. An identification key for the three species is given, main morphological differences are depicted and data on their biotope, life history, distribution and time trends are provided and discussed. Berend Aukema, Pomona 66, 6708 CC Wageningen, The Netherlands. Keywords. - Carabidae; Calathus, cinctus, melanocephalus, mollis, cross-breeding; identi- fication key; life history; distribution maps; time trends. In the past the status of Calathus melanocephalus (Linnaeus, 1758) and C. mollis (Marsham, 1802) as different species has been questioned many times, mainly because of the frequent occurrence of mor- phologically intermediate specimens. Schatzmayr (1937), Lindroth (1943) and Wiebes-Rijks (1959) treated them as distinct species, whereas Friede- richs (1907) and Gersdorf (1937) could not find any distinguishing character between both taxa at all. Furthermore the existence of a morphologically intermediate form cinctzs Motschulsky, 1850 (= erythroderus Gemminger & Harold, 1868), estab- lished as a subspecies of mollis by Lindroth (1945), masks the status of these taxa. This so-called melanocephalus group belongs to Neocalathus, established as a subgenus of the genus Calathus Bonelli by Ball and Negre (1972) with Calathus melanocephalus as the type species. For nomenclature in the melanocephalus group see Au- kema (1990) and Aukema & Luff (1990). The discovery at Wijster of populations of a cinc- tus-like Calathus on abandoned agricultural fields in the surroundings of the Biological Station of the Agricultural University Wageningen (van Dijk 1978) offered a good opportunity for studying its taxonomic relationship with both melanocephalus and mollis, and its life history. Moreover, in the present study cross-breeding experiments between these three taxa were carried out to elucidate their taxonomic status. Morpholog- ical data (including biometrics) were used to sup- port the resulting classification. The taxonomic characters given in literature to distinguish melanocephalus and mollis from each other are: l. The coloration of the pronotum in comparison to the head (and elytra): clear rufous, contrast- ing with the black head and elytra in melanoce- phalus, and piceus to brown and not or little contrasting in mollis. In the typical form of melanocephalus and in mollis all appendages are pale. However, a varying degree of mela- nism may occur in melanocephalus, in wich the pronotum is more or less infuscated, from pos- sessing only a faint cloud at the centre to being black with only narrowly translucent margins (in that case also the appendages are melanistic, at least tarsi brown, often also the main part of legs and palpi, as well as middle antennal seg- ments (Jeannel 1942, Lindroth 1974, 1986, Ve- reshchagina 1985). The shape and size of the right paramere: wi- dened at the tip and hooked at the extreme apex in melanocephalus and not widened and un- armed or with a very small hook at the apex in mollis (Lindroth 1974, 1986, Freude 1976, Ve- reshchagina 1985). Differences in internal and external morphology of the vesica are indicated by Lindroth (1943) and Vereshchagina (1985). 3. The dimensions of the metepisterna: short in melanocephalus (ca. 1.6 times as long as wide in both brachypterous and macropterous spec- imens), and longer in macropterous specimens of mollis (ca. 1.8 times as long as wide) (Jeannel 1942, Lindroth 1974, Freude 1976, Vereshchag- ina 1985). N 1941 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 4. The shape of the pronotum: more slender, with the greatest width usually before the middle and clearly narrowed towards the base in mol- lis, and less slender, with the greatest width in or behind the middle and sides less convergent in the basal half in melanocephalus (Freude 1976, Lindroth 1986, Vereshchagina 1985). 5. The shape of the elytra: longer oval (more stretched), with lateral margin curved and gradually tapering apically in mollis, and shor- ter oval, with sides almost parallel and broadly rounded apically in melanocephalus (Freude 1976, Lindroth 1986). According to both Lindroth (1974, 1986) and Freude (1976) the subspecies mollis cinctus (at the time known as mollis erythroderus) can be distin- guished from mollis mollis by the entirely rufous pronotum, contrasting with the piceus to brown head and elytra. Van Dijk (1978) furthermore men- tions a difference in the colour of the sternites between cinctus and melanocephalus. Material col- lected in the field was studied to evaluate the use- fulness of these characters for identification. Fig. 1. Collecting sites of Calathus cinctus, C. melanoce- phalus and C. mollis in The Netherlands. - 1, Drenthe (Dwingeloo, Kraloo, Nuil); 2, Ameland (Buren, Nes); 3, Texel ('t Horntje); 4, Oost-Flevoland (Ecological Re- serve); 5, Amsterdam (Bijlmermeer), 6, Wassenaar (Meijendel); 7, Schouwen (Haamstede). 122 Finally museum collections were revised in order to get an impression of the distribution of the spe- cies involved in The Netherlands. Due to misiden- tification in the past, existing distribution maps of the taxa concerned (Turin, Haeck & Hengeveld 1977) have to be replaced. In this paper the taxonomic status of the three taxa mentioned above is unravelled and the data presented lead to the conclusion that Calathus cinc- tus can be considered as a good species. Data on habitat, life history and distribution of cinctus, me- lanocephalus and mollis are given. MATERIAL AND METHODS Cross-breeding experiments Single-pair crosses between all possible combina- tions of the three taxa were carried out during 1978/1979 and 1979/1980 under approximately outside conditions in an outdoor insectary. In 1978/1979 selected pairs were kept either in glass 50 km jars on sieved peat litter (group 1) or in petri dishes in ground peat litter (group 2). In the first case newly emerged larvae were gathered from the peat litter as described by van Dijk (1979a), whereas for the second group eggs were separated from the peat according the sieve-wash method of Mols et al. (1981). Eggs were incubated at a constant temper- ature of 19°C. Larvae were moved to glass tubes in which they were reared individually in peat litter in the outdoor rearing facility. During the winter months (from the end of November until the end of March) the larvae were moved to a climate room at 5°C under short day (LD 10/14) conditions. Single-pair crosses of males and females of the same taxon served as control groups. Crosses and origin of the beetles used are given in table 1. Offspring of crosses performed by van Dijk (1978) in 1977/1978 and specimens collected in the field before the start of the reproduction period (tenerals in the case of females; cinctus from Dwin- geloo, melanocephalus from Kraloo, and mollis from Voorne) were used as starting material (table 1). Furthermore, offspring of females inseminated in the field (cznctzs from Nuil and Dwingeloo, and melanocephalus from Oost-Flevoland) were reared in order to get virgin females as starting material for the next crossing programme. In 1979/1980 crosses were carried out compara- ble to those of group 2 of the preceding year. The food, however, was changed from pieces of meal- worm to small maggots. Again offspring of females inseminated in the field (mollis from the West Frisian Island of Ameland) was reared. Information on these crosses is summarized in table 2. Collecting sites are depicted in fig. 1. Fig. 2. Measurements. BL: body length, BW: body width, PL: length of pronotum, PW: greatest width of prono- tum, EL: length of elytron, EW: greatest width of elytron, WL: greatest length of wing, WW: greatest width of wing, TIL: length of hind tibia, TAL: length of hind tarsus. AUKEMA: Three species of Calathus Morphology Material collected in the field was studied to check the validity and usefulness of characters given in literature for identification of the species con- cerned. Samples of the following localities were used: Calathus cinctus: Drenthe: Dwingeloo (Noorden- veld), Ruinen (Nuil); Friesland: Ameland (Buren, Nes); N-Holland: Texel Ct Horntje); Zuid-Hol- land: Wassenaar (Meijendel) and Zeeland: Schouwen (Haamstede). Calathus melanocephalus: Friesland: Ameland (Buren, Nes); Drenthe: Dwingeloo (Noordenveld), Ruinen (Nuil, Kraloo); Flevoland: Oost-Flevoland (Oecologisch Reservaat); N-Holland: Texel Ct Horntje), Amsterdam (Bijlmermeer). Calathus mollis: Friesland: Ameland (Buren, Nes); Z-Holland: Wassenaar (Meijendel) and Zeeland: Schouwen (Haamstede). Measurements were made with a Wild M5 ste- reo-microscope provided with a calibrated ocular micrometer. The following measurements were taken (magnification between brackets): TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 1. Single-pair crosses between and within Calathus cinctus, C. melanocephalus and C. mollis in 1978/1979. cin: cinctus; mel: melanocephalus, mol: mollis; lw: long winged; sw: short winged; N: number of crosses; NL: ibid, producing larvae; L: number of larvae produced and reared; LR: number of larvae reared; B: number of beetles reared. PARENTS Species/Wing-morph Origin Males Females Males Females N NL LAON Group 1 mel sw cin lw Kraloo (28-6-1978) F;-offspring 1978!) 4 - - - mel sw cin lw Kraloo (28-6-1978) Dwingeloo (19-7-1978, teneral) 1 - - = mel sw cin sw Kraloo (28-6-1978) F -offspring 1978 5 = = = cin lw mel sw F,-offspring 1978 Kraloo (28-6-1978, tenerals) 5 Lai - cin sw mel sw F,-offspring 1978 Kraloo (28-6-1978, tenerals) 5 192 = Group 2 cin lw mel sw Nuil (4-8-1978) F,-offspring 1978 6 - - - cin lw mel sw F,-offspring 1978 F,-offspring 1978 6 - - - cin sw mel sw Nuil (4-8-1978) F,-offspring 1978 2 - - - cin sw mel sw F -offspring 1978 F -offspring 1978 7 = - - mol lw mel lw Voorne (end July 1978)°) F,-offspring 1978 1 - - - mol lw mel sw Voorne (end July 1978) F;-offspring 1978 5 = = = cin lw mol lw Voorne (end July 1978) Voorne (end July 1978) 5 2 2Y - mel sw mol lw Kraloo (4-8-1978) Voorne (end July 1978) 5 22 - LR Controls cin lw cin lw Parents 1977!) F -offspring 1978 6 DS) cin sw cin lw F,-offspring 1978 Parents 1977 6 30 149759 cin lw no males Dwingeloo (23-8/22-9-1978) 16 16 734 471 cin lw no males Nuil (23-8/22-9-1978) 5 >) 259.198 cin lw cin sw F;-offspring 1978 Parents 1977 6 4 114 48 ein sw cin sw F -offspring 1978 F,-offspring 1978 6 6 295 1182 cin sw no males Dwingeloo (7-9-1978) 1 I 49 27 cin sw no males Nuil (23/31-8-1978) 4 3 147 86 mel sw mel sw Kraloo (30-8-1978) Kraloo (30-8-1978) 6 6 60 40 mel lw no males O-Flevoland (20-9-1978) 14 13 489 304 mel sw no males O-Flevoland (27-9-1978) gy CIO syle 236 '): parents and offspring of crosses carried out by Th. S. van Dijk *): material collected by A. van Tiggele ML1 Fig. 3. Measurements of met-episternum. ML1, ML2: length; MW1, MW2: width. Scale 0.5 mm. 124 - length and width of the body (12): length between tip of mandibles and apex of elytra and greatest width over elytra (fig. 2: BL and BW). - length and width of the pronotum (25): median length and greatest width (fig. 2: PL and PW). - length and width of the (right) elytron (12): EL and EW in fig. 2. EW was calculated as half the greatest body-width (0.5*BW). - length and width of the (right) wing (12): greatest length and greatest width (fig. 2: WL and WW). - length and width of the (right) met-episternum (50) as shown in fig. 3 (length: ML1, ML2; width: MW1, MW2). - length of the (right) hind tibia and of the (right) hind tarsus (25) (fig. 2: TIL and TAL). Elytra, wings, met-episterna and hind-legs were removed from the body for proper measurement in a flat plane. Wings were put on a slide in alcohol and unfolded with a fine brush. From these measurements ratios between length and width of the body (BL/BW), width and length | AUKEMA: Three species of Calathus Table 2. Single-pair crosses between and within Calathus cinctus, C. melanocephalus and C. mollis (1979/1980). cin: cinctus; mel: melanocephalus, mol: mollis; lw: long winged; sw: short winged; N: number of crosses; E: number of eggs laid; NL: number of crosses producing larvae; L: number of larvae produced and reared; LR: number of larvae reared; B: number of reared beetles. PARENTS Species/Wing-morph Origin Males Females Males mel lw cin lw Oost-Flevoland (11-10-1978) mel sw cin lw F,-offspring 1979 (O-Flevoland) mel sw cin lw F,-offspring 1979 (Kraloo) mel lw cin sw Oost-Flevoland (11-10-1978) mel sw cin sw F,-offspring 1979 (O-Flevoland) mel sw cin sw F,-offspring 1979 (Kraloo) mol lw cin lw Ameland (28-6-1979) mol lw cin sw Ameland (28-6-1979) cin lw mel lw F -offspring 1979 cin lw mel lw Ameland (28-6-1979) cin Sw mel lw F -offspring 1979 cin sw mel lw Ameland (28-6-1979) cin lw mel sw F,-offspring 1979 cin lw mel sw Ameland (28-6-1979) cin lw mel sw F -offspring 1979 cin lw mel sw Ameland (28-6-1979) cin sw mel sw F -offspring 1979 cin sw mel sw Ameland (28-6-1979) cin Sw mel sw F,-offspring 1979 cin sw mel sw Ameland (28-6-1979) mol lw mel lw Ameland (28-6-1979) mol lw mel sw Ameland (28-6-1979) mol lw mel sw Ameland (28-6-1979) Control groups mel lw mel lw F -offspring 1979 mel sw mel lw F,-offspring 1979 (O-Flevoland) mel sw mel lw F;-offspring 1979 (Kraloo) mel lw mel sw F,-offspring 1979 mel sw mel sw F -offspring 1979 (O-Flevoland) mel sw mel sw F;-offspring 1979 (Kraloo) mel lw mel sw F -offspring 1979 mel sw mel sw F -offspring 1979 (O-Flevoland) mel sw mel sw F -offspring 1979 (Kraloo) cin lw cin lw F -offspring 1979 cin lw cin lw Ameland (28-6-1979) cin sw cin lw F -offspring 1979 cin sw cin lw Ameland (28-6-1979) cin lw cin sw F -offspring 1979 cin lw cin sw Ameland (28-6-1979) mol lw no males Females = NIEREN TE F,-offspring 1979 4 422 5) 4168 - F,-offspring 1979 4 591 - - - F -offspring 1979 4 320 = È _ F,-offspring 1979 4 91 - - - F,-offspring 1979 4 219 = = È F,-offspring 1979 AN u Bs à F,-offspring 1979 4 184 = 2 = F -offspring 1979 4 254 = = = Oost-Flevoland (25-10-1978) Oost-Flevoland (25-10-1978) Oost-Flevoland (25-10-1978) Oost-Flevoland (25-10-1978) F,-offspring 1979 (O-Flevoland) F,-offspring 1979 (O-Flevoland) NNNNNNNNNNNN WN O0 F,-offspring 1979 (Kraloo) 218 DE - F,-offspring 1979 (Kraloo) 14 - = - F,-offspring 1979 (O-Flevoland) 15 = = - F,-offspring 1979 (O-Flevoland) 50 2 - F,-offspring 1979 (Kraloo) 87 - F,-offspring 1979 (Kraloo) 4 - - Oost-Flevoland (25-10-1978) 4 181 25 - F,-offspring 1979 (O-Flevoland) 4 57 - - F,-offspring 1979 (Kraloo) 459 = = = LR F,-offspring 1979 4 1173 4 238 85 F,-offspring 1979 4 1084 4 240 135 F,-offspring 1979 4 1279 A Zilk 9) F,-offspring 1979 (O-Flevoland) 4 1113 4239 92: F,-offspring 1979 (O-Flevoland) 4 959 4 240 130 F,-offspring 1979 (O-Flevoland) 4 665 4 224 136 F,-offspring 1979 (Kraloo) 4 682 4 160 96 F,-offspring 1979 (Kraloo) 4 1079 4 240 132 F;-offspring 1979 (Kraloo) 4 920 4 240 112 F,-offspring 1979 2 468 21207243 F,-offspring 1979 157232 17607520 F;-offspring 1979 2 639 2585) el F -offspring 1979 1 509 IGOR F,-offspring 1979 2 496 271167168 F,-offspring 1979 152216 122607729 Ameland (28-6-1979) 28 573 9 309 194 of the pronotum (PW/PL), pronotum width and body width (PW/BW), length and width of the elytra (EL/EW), length of the elytra and body length (EL/BL), length and width of the wings (WL/WW), wing-length and length of the elytra (WL/EL), length and width of the met-episterna (ML2/MW2), and the ratio between the lengths of the hind tarsus and hind tibia (TAL/TIL) were estimated. Special attention was paid to the coloration of the beetles, the size and shape of the right paramere and to the number of dorsal punctures in the third elytral interval. 125 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Habitat and life history Details of the habitat and the life history of the species were derived from the analysis of field data (mixed populations of cinctus and melanocephalus from Dwingeloo, Ruinen and the West Frisian Is- land of Texel and populations of cinctus and mollis from the West Frisian Island of Ameland), from literature (populations of melanocephalus studied by Vliim & van Dijk (1967), Vlijm et al. (1968) and van Dijk (1972, 1973)) and from breeding experi- ments with the three species under outside condi- tions carried out in 1981/1982 and 1982/1985. Distribution Calathus-material of this group from Dutch col- lections was revised to be able to get an impression of the distribution of the species in The Nether- lands. Material from the following collections is included: Museums and institutions: Zoölogisch Museum, Amsterdam; Rijksmuseum van Natuurlijke Histo- rie, Leiden; Vakgroep Entomologie, Landbouwuni- versiteit, Wageningen; Biologisch Station, Land- bouwuniversiteit, Wijster. Private collections: K. den Bieman, Bennekom; E. H. M. Bouvy, Beek; H. T. Edzes, Nijmegen; T. van Gijzen, Arnhem; Th. Heijerman, Wageningen; S. van Heijnsbergen, Naarden; Y. Jongema, Wage- ningen and A. P. J. A. Teunissen, Vlijmen. Additional material was supplied by K. Alders, Arnhem; M. A. Baars, De Koog, Texel (Loopkever- werkgroep Texel); Th. S. van Dijk, Wijster; P. J. M. Mols, Heteren and H. Turin, Renkum. Time trends in the occurrence of the species (i. e. numbers of occupied 10 X 10 km squares of the distribution maps per decade since 1870) were es- timated according to the method of Turin & den Boer (1988). RESULTS Cross-breeding experiments In total 123 single-pair crosses were carried out: 59 between cinctus and melanocephalus, 47 be- tween cinctus and mollis and 17 between melanoce- phalus and mollis (tables 1 and 2). None of these crosses produced adult offspring although in 5 (8.5 %), 3 (6.4 %) and 3 (17.6 %) cases respectively larvae were produced, which, however, never reached the third larval stage. Table 3. Measurements (in mm) of short winged males of Calathus cinctus and C. melanocephalus from Texel. N: number of measurements. cinctus melanocephalus MEASUREMENT N Mean 95% cl. N Mean 95% cl. Body Length (BL) 20 6.79 6.59-6.98 20 6.66 6.51-6.82 Width (BW) 20 2.71 2.64-2.77 20 2.67 2.61-2.73 BL/BW 20 DL 2.47-2.55 20 2.50 2.45-2.55 Pronotum Length (PL) 20 1.64 1.61-1.68 20 1.68 1.65-1.72 Width (PW) 20 199) 1.95-2.04 20 2 2.06-2.15 PW/PL 20 1.21 1.20-1.23 20 1825 1.23-1.27 PW/BW 20 0.74 0.73-0.74 20 0.79 0.78-0.80 Elytron Length (EL) 20 8 4.07-4.29 20 4.09 4.00-4.18 Width (EW) 20 195 1.32-1.38 20 1.99) 1.30-1.36 EL/EW 20 3.09 3.05-3.12 20 3.07 3.03-3.11 EL/BL 0 0.62 0.61-0.62 20 0.61 0.60-0.63 Wing Length (WL) 20 1.93 1.85-2.01 20 1.93 1.86-2.00 Width (WW) 20 0.48 0.45-0.51 20 0.51 0.47-0.54 WL/ WW 20 4.05 3.86-4.24 20 3.86 3.65-4.07 WL/EL 20 46 0.45-0.47 20 0.47 0.46-0.49 Met-episternum Length (ML2)') 20 0.76 0.74-0.78 20 0.70 0.68-0.73 Width (MW2)!) 20 0.57 0.55-0.59 20 0.57 0.55-0.58 ML2/MW2 (ME2) 20 1.34 1.32-1.36 20 1.24 1.22-1.27 Hindleg Length tarsus (TAL) 20 2.07 2.01-2.13 20 2.02 1.98-2.06 Length tibia (TIL) 20 2.02 1.98-2.07 20 1.98 1.94-2.01 AE DIE 20 1.02 1.01-1.03 20 1.02 1.01-1.04 Dseertie 8; All control crosses (42 of melanocephalus and 30 of cinctus) crosses that produced larvae (respec- tively 100 % and 82 % for both years together), also produced adult offspring (tables 1 and 2). In 1979/1980 all crosses of both species produced offspring (table 2). In the case of mollis not suffi- cient material to carry out control crosses in the same way was available. However, of 28 females collected in the field (Ameland, Buren, 28.vi. 1979) and kept without males under the same conditions as single-pair crosses, 9 produced larvae and off- spring (table 2: 194 beetles from 309 larvae). The other 19 females (ten of which were tenerals at the time of capture) produced no or only a few unfer- tilized eggs, and were considered not to have been inseminated at all. Nevertheless it is clear from these results that also in the case of mollis the failure of single-pair crosses with cinctus and me- lanocephalus cannot be attributed to the rearing conditions used. In 1979/1980 egg production was also estimated. It appeared that, although unfertilized females may produce a considerable number of eggs, the egg production of fertilized females generally is much higher (table 2). It is concluded that cinctus, mela- nocephalus and mollis show mutual reproductive isolation, and, therefore, have to be considered as distinct species. AUKEMA: Three species of Calathus Morphology Biometric data, especially the ratio length/ width of the met-episterna, have been used in attempts to separate the taxa of this group of closely related species (Gersdorf 1937, Wiebes-Rijks 1959). The establishment of cinctus as a valid species as the result of the breeding program necessitated a new attempt to find morphological differences between the three species. Concerning biometrics, however, two points have to be taken into account. Firstly general differences both between sexes (on average females are larger than males) and between wing- morphs (on average long winged specimens are larger than short winged specimens and differently shaped) exist in these species. Secondly, the way measurements are made should be described (and figured) in detail, which apparently is not normal practice. Lindroth (1974) and Freude (1976), for instance, use the ratio length/width of the met- episterna without indicating how to measure it, and do not point out the differences between sexes and between wing-morphs in a proper way. Tables 3-6 list measurements and ratio's of both males and females of short as well as long winged Calathus cinctus and C. melanocephalus collected on Texel by M. A. Baars and of long winged C. mollis collected in the coastal dunes near Haam- Table 4. Measurements (in mm) of short winged females of Calathus cinctus and C. melanocephalus from Texel. N: number of measurements. cinctus melanocephalus MEASUREMENT N Mean 95% cl. N Mean 95% cl. Body Length (BL) 20 1559 7.23-7.56 20 732 ~7.18-7.46 Width (BW) 20 3.01 2.94-3.09 20 2.95 2.91-3.00 BL/BW 20 2.46 2.42-2.50 20 2.48 2.44-2.52 Pronotum Length (PL) 20 177 1.73-1.81 20 1.79 1.76-1.82 Width (PW) 20 2,19) 2.13-2.24 20 2.28 2.24-2.32 PW/PL 20 1.23 16222125 20 1.27 1.26-1.28 PW/BW 20 0.73 0.72-0.74 20 0.77 0.76-0.78 Elytron Length (EL) 20 4.54 4.44-4.64 20 4.45 4.36-4.53 Width (EW) 20 Lil 1.47-1.54 20 1.48 1.46-1.50 EL/EW 20 3.02 2.98-3.05 20 3.01 2.97-3.04 EL/BL 20 0.61 0.61-0.62 20 0.61 0.60-0.62 Wing Length (WL) 20 2.14 2.08-2.20 20 2.06 1.99-2.13 Width (WW) 20 0.60 0.57-0.62 20 0.60 0.55-0.64 WL/WW 20 3.61 3.48-3.75 20 3.49 3.32-3.65 WL/EL 20 0.47 0.46-0.48 20 0.46 0.45-0.48 Met-episternum Length (ML2) 20 0.83 0.80-0.85 20 0.76 0.74-0.77 Width (MW2) 20 0.64 0.62-0.65 20 0.62 0.61-0.64 ML2/MW2 (ME2) 20 1.30 1.27-1.33 20 1.22 1.20-1.24 Hindleg Length tarsus (TAL) 20 2.08 2.03-2.12 20 197 1.92-2.01 Length tibia (TIL) 20 2.07 2.03-2.12 20 2.02 1.98-2.06 TAL/TIL 20 1.00 0.99-1.02 20 0.97 0.96-0.98 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 5. Measurements (in mm) of long winged males of Calathus cinctus and C. melanocephalus from Texel and C. mollis from Haamstede. N: number of measurements. cinctus melanocephalus mollis MEASUREMENT N Mean 95% cl. N Mean 95% cl. N Mean 95% cll. Body Length (BL) IRR 52-7057 TMD OSB) U ee TEN Width (BW) 1 CAMP 55-2573 0 ar 2:39, aD 70) 29) Do BL/BW AZ, IED 2.52 244-260 20 2.63 2.61-2.66 Pronotum Length (PL) 160 AE Co LOL TO STE 20,169: 1:61.69 Width (PW) U Oe ESS 15997 D'LA AT SI 205. 2205) 2.002209, PW/PL This ZINN TENS Zed LMD TE ALD) PW/BW RI OT. 072220, 74 7) 0.79 0.78-0.79 20 0.73 0.73-0.74 Elytron Length (EL) 1197 212705 3.98-4257 432 419-445 20 438 427-449 Width (EW) br SZ AEN 7 PASS SEAT 20771995 31362142 EL/EW LIN 31030, 31030 EON 20 ESE EL/BL Ue OHG OEE 97 0.60 0.58-0.62 20 0.60 0.59-0.60 Wing Length (WL) 1.2629 SIL 7.10 686-785 20 6.61 6.39-6.83 Width (WW) I 12.36 223250 7 2.68 263-274 20 2.46 237-254 WL/WW Il m 2.097 Pos DOSE 258-25) 220i 2:69 ae 66-27/8, WL/EL u 147157007 SA 16216720 IODS Met-episternum Length (ML2) U OT 07420797 0.77 0.73-0.81 20 082 0.80-0.84 Width (MW2) STe LOOD 0.60 0.58-0.62 20 058 0.56-0.59 ML2/MW?2 (ME2) IE > E o 7 1.28 121-135 20 143 140-145 Hindleg Length tarsus (TAL) UO OEE 6 DEF NEP WA DO ZD ZLD DS) Length tibial (TIL) 110 221:96.591.902203556 2.13 2.04-2.22° 20. 2:09), 2.04-2.15 TAL/TIL 11 102 101-104 6 102 1.00-1.05 20 1.05 1.04-1.06 Table 6. Measurements (in mm) of long winged females: Calathus cinctus and C. melanocephalus from Texel and C. mollis from Haamstede. N: number of measure-ments. cinctus melanocephalus mollis MEASUREMENT N Mean 9%cl N Mean 95% cl. N Mean 95% cl. Body Length (BL) 11 927:.04678-7.50 2077.85 78722797 a201 8:03: EBEN, Width (BW) D287 274300 20775157 52341870202 3.187) 3:103:25 BL/BW 1782462241251 2072497 24525108208 25372517255 Pronotum Length (PL) Ll ve EGT 1:60-1.74 20) E89" 71:86:19 rn ZON TIEKE Width (PW) Hy 32.0977731:99 220207 245 2402747 002.0 22802222354 PW/PL LUPE Or SEE 20 IE 127215198205 219 825419721050 PW/BW 11 0.73 0.72-0.74 20 0.77 0.76-0.78 20 0.72 0.71-0.72 Elytron Length (EL) 11 434 418-450 20 480 472-488 20 481 471-492 Width (EW) ILS ESES 0! 220) ES NS CES OMZ OM 7159915521762 EL/EW 110 0302083 0820 05 NS 003 00205 0 SUIS 06 EL/BL 11 062 061-063 20 061 0.60-0.62 20 0.60 0.60-0.60 Wing Length (WL) 1190649 CNE C2 07 SIN 5827 COR 2 00727 OTETEZG Width (WW) MSI 923822:642 2.07 92:85 227/7528902 ON ETE 68-285 WL/WW 1192258 255222164. 201 265 2.62.2269 208 2656020 WL/EL TIRA 1451541207157. LES ATS OAN 207 151149152 Met-episternum Length (ML2) 11 0.82 0.77-0.88 20 083 081-084 20 092 0.88-0.94 Width (MW2) 11 0.61 0.58-0.63 20 0.68 0.66-0.69 20 0.65 0.63-0.66 ML2/MW?2 (ME2) MET 367 MESSE 20 AIDE ZOR 2395 Hindleg Length tarsus (TAL) 110292:00 7 7188921069207 22.092220 7A 2 0,7291 52711672775 Length tibial (TIL) 9719222108 0727187723529 252 O ZEE 2152225 TAL/TIL 11 101 099-104 20 096 094-097 20 1.01 1.00-1.02 128 AUKEMA: Three species of Calathus Figs. 4-6. Habitus of females of Calathus. — 4, C. melanocephalus; 5, C. cinctus, 6, C. mollis. Scale 1 mm. stede (Province of Zeeland) by the author. The following conclusion can be drawn. Although there are some differences between the means (e. g. in the case of the ratio length/ width of the metepisterna), these measurements (or ratios estimated from them) do not allow unambiguous identification due to significant individual variation and wide overlap in the ranges of measurements. Table 7 summarizes length/width ratios (MEI: ML1/MW1 and ME2: ML2/MW2, see fig. 3) of long and short winged males and females of cinctus and melanocephalus, as well as of long winged males and females of mollis (all specimens collected or reared of the latter species turned out to be long winged) from localities mentioned above, as well as from a number of other localities too. Although both ratios show a clear trend mela- nocephalus 50 %) from the standard number of three punctures in the third elytral interval was found, whereas the deviation in melanocephalus and mollis was relatively low (< 40 %). Both the number and position of the punctures are also rather variable and certainly not of diagnos- tic value. General coloration, shape of the elytra and shape of the right parameres are very constant and can be used to identify most specimens according to the key presented hereafter. Only a few long winged females of cinctus and mollis may give some prob- lems in identification. 129 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 7. Measurements of met-episterna: ratio length/ width. MEI: ML1/MW1; ME2: ML2/MW2 (see fig. 3); sw: short winged; lw: long winged; N: Number of measurements. MALES Wing-morph Species SW SW lw FEMALES SW SW lw Locality cinctus melanocephalus cinctus melanocephalus mollis cinctus melanocephalus cinctus melanocephalus mollis Nuil Noordenveld!) Texel Oost-Flevoland Nuil Noordenveld!) Kraloo Bijlmermeer Texel Nuil Noordenveld!) Texel Oost-Flevoland Kraloo Bijlmermeer Texel Meijendel Haamstede Nuil Noordenveld!) Texel Oost-Flevoland Nuil Noordenveld!) Kraloo Bijlmermeer Texel Nuil Noordenveld!) Texel Oost-Flevoland Kraloo Bijlmermeer Texel Meijendel Haamstede 1): measurements M. G. J. Oude Wesselink MEI ME2 N Range Mean 95% cl. Range Mean 95% cl. 30 106-118 1.11 110-112 1.28-1.46 136 1.35-1.38 27 10522221711 1.09-1.15 “119-47 134 131137 20 0.97-1.10 1.04 1.03-1.06 125-143 134 1.32-1.36 30 09921916, 1:0777:05-108 EO 3 ONE 29 PT 19 0.96-1.10 1.04 1.02-1.06 1.15-1.31 1.23 1.21-1.25 30 OVE AOS NO = POS NONO 25930 9) 0.98-1.10 104 1.01-1.06 1.26-1.29 1.24 1.20-1.28 gf 103-114 1.07 1.06-1.09 1.23-1.40 1.27 1.25-1.30 20 0.95-1.07 1.02 1.00-103 115-132 1.24 1.22-1.27 30 ROIO MERE EDA OO ees ele al 30 1.04-1.26 1.14 IMO AO PRINS SES JIS 1.33-1.40 11 ROO OIO DINO OIS TIEL ES Seles E139 13 1.00-1.15 1.08 105-111 114-134 124 121-128 2 1.09-1.12 1.39-1.45 3 1.07-1.09 1.21-1.29 7 1.00-1.09 1.05 1.02-1.08 1.20-1.40 1.28 1.21-1.35 14 IS 1 25 INO El EE) OA 20 alien AO) A6 MEET 1.40-1.45 20) 1.02-1.18 1.11 1.09-1.12 124-149 1.35 15352197 29 1.00-1.20 1.08 1.07-1.10 1.17-144 130 1.28-1.33 20 0.921.070 10120991025 114313027130 21272133 30 OOG OD MIDI OI ES 125 DET 30 1.00-1.10 1.04 1.03-1.05 1.14-139 1.25 1.23-1.27 30 0.98-1.22 106 104-109 1.16-139 127 1.24-1.29 12 1.00-1.13 105 103-108 118-137 1.24 1.18-1.30 10 0.98-1.18 1.09 1.05-112 1.23-1.33 1.27 1.24-1.30 20 0.92-1.07 1.01 099-102 114-130 1.22 1.20-1.24 30 KONZ 1.11-1.13 1.26-1.44 1.35 1.34-1.37 29) DOS OMAN WIG rele ASS SANA CI 31850) 11 LOB 1.08-1.14 1.25-1.49 136 131-141 30 0.97-1.13 1.06 1.04-107 1.12-141 1.26 1.24-1.28 18 0:98212187 17.087 7705-1117 1482 180551905154 7 NOMA WO, BETT LS 005112. 6216577 20 095-108 1.03 101-104 117-130 1.22 121-124 17 1.13-1.28 1.19 1.17-1.21 133-154 143 140146 20 1.05-1.24 116 115-118 129-161 142 1.39-1.45 Tabel 8. Number of punctures in 3rd elytral striae. N: number of observations; D: deviation (%) from standard number of puntures (3/3) in third striae; sw: short winged; lw: long winged. Punctures in 3rd elytral stria (right/left elytrum) Species Sex Wing morph N cinctus males sw 20 lw 11 females sw 20 lw 11 melanocephalus males Sw 20 lw 7 females sw 20 lw 20 mollis males lw 20 females lw 20 130 WO DB DL BR BIB Bie LB fin ids bale =p Think Sue SOR? Sin hed RT i Rc eRe) en RR Delbo hal sl TU À à e x = 5 2 E A Rt Ind HEISA AA on dee ee D RZ e aan ce Pre NE ka Md A CALL ll 4/4 DW NR be 4/5 5/3 5/4 D = Dp) - 545 90.0 - 63.6 = ap - 286 | SI) = 300m =, 15.00 550 Figs. 7-9. Right parameres of Calathus.—7,C. melanoce- phalus; 8, C. cinctus; 9, C.mollis. Scale 0.5 mm. In all three species the females are less shining than the males, due to a denser microsculpture of usually isodiametric meshes, which is most obvious on the elytra. Key to the species 1. Elytra bicoloured, with epipleura, base and dor- sal punctures clear rufous, contrasting with the black dorsal surface. Sides of elytra almost pa- rallel and broadly rounded apically (fig. 4). Pronotum more or less rufous, clearly contrast- ing with the black head and with the elytral surface. Sternites dark, contrasting with the ru- fous meso- and meta-thoracical structures. Males: right paramere widened and hooked at the tip (fig. 7). Vesica: fig. 10. Wings reduced or fully developed (rarely specimens with only one fully developed wing are found) (body IENE th (6-8 IMM) ER melanocephalus — Elytra unicoloury rufous to piceus brown, with sides evenly rounded (figs. 5-6). Little or no contrast in colour between pronotum and head and elytra, nor between sternites and meso- and meta-thorax. Males: right paramere not wi- dened near the apex and unarmed or with a very small hook apically (figs. 8-9) ................. 2 2. Pronotum slightly contrasting with the darker head and the elytra. Males: right paramere hooked, tip unarmed or nearly so (fig. 8). Ves- ica: fig. 11. Wings either fully developed or strongly reduced (body length 6-8.5 ON) EEEN IE n OM Aert hd cinctus - Pronotum not contrasting with the head and the elytra. Males: right paramere almost straight with a very small hook apically (fig.9). Vesica: fig. 12. Wings always full (body length CINTO Aaa MER a RR OM Was mollis AUKEMA: Three species of Calathus Habitat and life history Habitat Calathus melanocephalus is mainly a species of open country, where it lives on different kinds of moderately dry soil with sparse vegetation, achiev- ing its greatest abundance on sandy soils. It is a common inhabitant of dry meadows, grassland dunes and heaths; also on agricultural land and in thin forest, mainly of Pinus (Lindroth 1986) or Betula (den Boer pers. comm.). Den Boer (1977) in his subdivision of carabids in habitat groups classi- fied melanocephalus as an F-species, i.e. mainly a species of heath and peat moor. Calathus cinctus is mainly found in mixed pop- ulations with the preceding and/or the following species in dry meadows, dune grassland, agricultu- ral land and waste land. Van Dijk (1986) classified cinctus as an H-species sensu den Boer (1977), i.e. an eurytopic species, which in most cases prefer localities significantly influenced by human activi- ties (arable fields, pastures, farm yards, gardens and Figs. 10-12. Vesica of Calathus. - 10, C. melanocephalus; 11, C. cinctus; 12, C. mollis. Scale 0.5 mm. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 9. Numbers and ratio of Calathus cinctus (cin) and C. melanocephalus (mel) in mixed populations. NUIL SCHOONGELEGEN TEXEL Species Species Species Year cin mel %cin cin mel %cin cin mel %cin 1973 11 30 26.8 8 71 10.1 1974 13 29 31.0 3 54 5.2 1975 35 64 35.4 6 23 20.7 1976 16 20 44.4 22 39 36.1 1977 34 24 58.6 39 31 Doll 1978 59 67 46.8 172 112 60.6 1979 12 34 26.1 3 62 4.6 1980 2 19 9.5 — 40 & 1981 = 16 - - 30 = 1982 = 6 = 72 = 1983 1 = = 84 = 1984 - 6 = 50 - 82 19 81.1 1985 - 4 - 91 = 264 265 49.9 1986 - 18 - 102 = 133 310 30.0 1987 32 176 155 1988 6 74 U Total 183 DO 253 861 Sly 844 Table 10. Weekly catches of Calathus cinctus and C. melanocephalus at Nuil, Drenthe in 1976. N: number of beetles; ND: number of females dissected, MNE: mean number of eggs in the ovaries of dissected females. cinctus melanocephalus Males Females Males Females Date N N ND Young Old MNE N N ND Young Old MNE 7 Jul 1 1 1 1 AUTO 14 Jul 5 4 4 3 1 4.3 21 Jul - 1 l 1 - 8 12 8 6 2 Wile 28 Jul = = = = = 14 18 10 9 115.9) 4 Aug - 1 1 1 2 15.0 7 10 9 8 a 11 Aug = = = = 5 11 25 ll 7 LO IS 18 Aug 1 2 2 2 - 23.0 35 26 10 7 3 22.6 25 Aug 3 2 2 1 1 35.0 46 17 10 9 1 23.0 1 Sep 13 15 15 7 8 20.2 42 34 10 8 2 25.8 8 Sep 10 16 16 9) 7 33.9 11 24 10 10 - 2165 15 Sep 21 20 20 15 5 26.5 18 21 10 10 - 1726 22 Sep 27 18 18 16 2 17.8 23 18 13 12 1 18.8 29 Sep 16 8 8 5 3 18.3 22 4 4 3 1 10.8 6 Oct 17 10 10 9 1 15.4 9 2 2 0 2 16.5 13 Oct 12 20 18 10 8 111152 6 10 10 9 Il 2.0 20 Oct 4 6 6 4 2 9.8 2 1 1 1 - - 27 Oct 2 6 6 3 3 8.0 2 4 2 2 - 3 Nov - 1 Il 1 - 1 = - 10 Nov 2 3) 1 1 - 3 1 - Total 128 129 125 85 40 262 231 125 105 20 roadside verges). In three cases a mixed population of cinctus and melanocephalus was observed during a period of several years: populations at Nuil and Dwingeloo (fields withdrawn from agricultural practice) and at the West Frisian Island of Texel (artificial dune grassland) (table 9). In all three cases, however, cinctus disappeared or nearly disap- peared after a number of years of coexistence with melanocephalus. These data might suggest that 152 cinctus is better adapted to changing, unstable hab- itats than melanocephalus. At Nuil and Dwingeloo the treatments for impoverishing the amount of nutrients (annual mowing and removal of vegeta- tion) most likely caused the decrease of cinctus (see | also van Dijk 1986), while on the dune grassland at | Texel a similar treatment might be responsible for | the same kind of shift. | Calathus mollis is strictly confined to dry sandy habitats in open country. It lives in sparse xerophi- lous vegetation, common on sand dunes along the coast, where it occurs in tufts of Elymus and Am- mophila in company with Demetrias monostigma Samouelle and Dromius linearis (Olivier) (Lind- roth 1986; own observations), and only incidentally inland, e.g. on blown sands and other large sand deposits. In den Boer's subdivision (1977) mollis would be classified as an E-species (species of blown sands). In the coastal dune region as well as in inland blown sands mixed populations of mollis and cinctus and occasionally with both cinctus and melanocephalus occur. Life history Annual rhytms. - All three species show a similar type of development. They belong to the group of autumn breeders with a thermic hibernation para- pause in the larvae and a photoperiodic aestivation parapause in the adults (Thiele 1977). As in most autumn breeders, adults of melanocephalus are al- most strictly night active (Thiele 1977). Larvae as well as a varying part of the adults overwinter (Lindroth 1986). In northern and alpine areas, however, melanocephalus is a semivoltine spring breeder, instead of a univoltine autumn breeder, adapted to the shorter growing season of a cooler climate (Forsskähl 1972, DeZordo 1979, Refseth 1988). Under these circumstances larvae of mela- nocephalus need almost two years to develop. Reproduction period and age-composition. - De- tailed information about the life history of mela- nocephalus is given by Vlijm & van Dijk (1967), Vlijm et al. (1968) and van Dijk (1972, 1973). Adults emerge in June and July, reproduce from August onwards and a varying part of these beetles hibernate to become active again in May-June, re- producing for the second time from July onwards. The total reproduction period covers the months July, August and September. Vlijm & van Dijk (1967) and Vlijm et al. (1968) estimated the second year category as about one third of the adult pop- ulation at Schiermonnikoog, but this figure seem to show significant annual and local variation (van Dijk 1972, Baars & van Dijk 1984). Even three and four year old beetles were found by a mark-recap- AUKEMA: Three species of Calathus mean number of eggs laid o cinctus O melanocephalus A mollis 30 35 40 45 50 August September October November December week number/month Fig. 13. Mean egg production of Calathus cinctus (29 females), C. melanocephalus (23 females) and C. mollis (16 females) and mean weekly temperature in 1981 under outside conditions. ture experiment in a populations of melanocepha- lus (van Dijk 1979b), representing 26.5 and 0.5 % of the total catch during four years respectively. Calathus cinctus was found to reproduce in Drenthe somewhat later than melanocephalus (Au- gust-November) (van Dijk 1978). From weekly pit- fall samples of cinctus collected in 1976 at Nuil, 165 females caught between 21 July and 17 November were dissected and checked for their ovarian devel- opment. The main reproductive period, defined Table 11. Reproduction period of Calathus cinctus, C. melanocephalus and C. mollis in outdoor breeding experiments. N: number of females. REPRODUCTION PERIOD Year Species N Egg-laying period Mean number of eggs> 10 1981 cinctus 29 12 Aug - 9 Dec (18 weeks) 26 Aug - 21 Oct (9 weeks) melanocephalus 36 31 Jul —6 Nov (15 weeks) 7 Aug - 9 Oct (10 weeks) mollis 14 3 Aug - 7 Dec (19 weeks) 31 Aug - 19 Oct (8 weeks) 1982 cinctus 30 16 Aug - 31 Jan (25 weeks) 30 Aug - 22 Nov (13 weeks) melanocephalus 8 30 Jul - 22 Oct (13 weeks) 13 Aug - 15 Oct (10 weeks) mollis 17 6 Aug- 3 Jan (23 weeks) 20 Aug- 5 Nov (12 weeks) 133 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 SIT IG OÙ G € MO vy I OI G SIL CO OV CO O Sy € O- BI [B10], \O N = oO £6 EY hii DC Ce GA Va, O¢ OL 1861 das LI ce N cn CON 4 Con 0£0 1861 das 91 9 1 1 N = N en HN = ON GS ON — Naso N 068 1861 das SI (11815) 0€8I 1861 das I QUI L 312 OBMOOOODODODOS LL (nese) een) a N [ELO SG ln Oe Gevangen Brela ee U — = A SU] OUL su» darourjau SHIIUII ‘(anqa) porad 1481 :] ‘Gpejg) porsad yep :q ‘(sImoYy 00:61-06 9 :porsad 1481) Yyasuay Kep jermeu Ajsyewrxordde pure ) „GI JO sanıesadurs IUEISUOI e je {ep ayı Furmp Syjow ‘9 pue suypgdaroupjpou ‘7 ‘sm19u19 snqipje Jo vOnonpord 33q ‘ZI algeL 134 mean number of eggs laid 30 35 40 45 50 1 August September October November December January AUKEMA: Three species of Calathus Fig. 14. Mean egg production of Cala- thus cinctus (30 females), C. melanoce- CI - 1993 ne DIA rome) and < mollis au females) and mean weekly tempera- tures in 1982/1983 under outside con- ditions. o cinctus O melanocephalus 4 mollis February week number /month here as the period in which the mean number of eggs in the ovaries is 10 or more, fell between the first week of August and the third week of October (table 10). Forty females (24.2 %) were classified as old females, reproducing for the second year. Comparable data for melanocephalus collected at the same site resulted in a main reproductive period between the third week of July and the first week of October and 16 % old females (table 10), of which only a single female caught on 13 October was not or no longer reproductive. Calathus mollis was not studied in detail in this respect but hand catches on the West Frisian Island of Ameland on 27 May 1981 comprised old beetles only (29), whereas on 28 June 1979 about 12 % of the 248 beetles collected at the same site were just emerged tenerals. These data suggest a similar type of age composition of populations of this species. In the laboratory a small part of the females of all three species kept individually even survived a second hibernation period and laid eggs during a third year. Outdoor breeding experiments carried out in 1981 and 1982 also gave an impression of the re- productive period of the three species (table 11). Also in these experiments melanocephalus showed an earlier start and finish of the egg-laying period than both cnctus and mollis. Differences between the results for the two years are attributed to differ- ent temperature conditions. In 1981, for instance, all females stopped egg production in the first third of December because of a sudden frost period (fig. 13). Due to the later start, cinctus and mollis suf- fered more from this event than melanocephalus. Under the more favourable weather conditions of 1982/1983 melanocephalus showed about the same reproduction period, whereas both cinctzs and mol- lis were able to prolong their reproduction periods substantially under these conditions (fig. 14). Data on egg production and larval development of the three species in relation to wing develop- ment, temperature and availability of food will be published elsewhere. Daily rhythms. — Information about the daily activity of the species was derived from a small experiment in which the egg production was esti- mated every two hours under fixed temperature and daylength conditions. On 14 September 1981 eight reproductive females of cinctus, melanocephalus and mollis, collected at Ameland on 14 July, were transferred from the outdoor breeding house to an incubator adjusted at 19°C and a daylength of 14.75 hours (6.15-19.00 h). They were kept there with 135 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 males in single pairs for three days. In all three species egg production took place mainly during the dark hours (18.30-6.30 h); a very few eggs were also laid during the early morning hours (table 12). Emergence of adults. - The outdoor breeding experiments also gave an impression of the emer- gence period of the adults (table 13). In 1982 almost all cinctus, melanocephalus and mollis emerged in the first two thirds of June, and in 1983 in the latter half of the same month. Van Dijk (1973) trapped teneral females of melanocephalus in the field as well as in enclosure experiments during a short period from 15 June onwards. Samples of cinctus and mollis collected on 28 June 1979 on the West Frisian Island of Ameland consisted of 75 % (102 of 138 specimens) and 11.7 % (26 of 248 speci- mens) tenerals respectively. Feeding. - Feeding behaviour and prey spectrum of melanocephalus were studied by Hengeveld (1980) by identifying food remains in the gut con- tents of beetles trapped in the field and killed in- stantly. From the diet components found, it ap- peared that melanocephalus is a polyphagous, but not exclusively carnivorous species (see also Melber 1983). Similar results were found by Smit (1957), Skuhravy (1959) and Hengeveld (1980) for the closely related Calathus ambiguus (Paykull), C. er- ratus Sahlberg and C. fuscipes (Goeze). Dispersal. - Den Boer (1977) developed criteria to classify carabid beetles in three dispersal groups using data on wing development, wing-morph fre- quencies and actual flight (catches of flying beetles in window traps). A-species are those with a rela- tively low power of dispersal (monomorph bra- chypterous or wing-polymorphic species with a very low frequency (< 1%) of macropterous bee- tles). B-species have a relatively high power of dispersal due to their capability of flight (macro- pterous and wing-polymorphic species caught in window traps, see also van Huizen 1980), whereas C-species are the remaining species (macropterous Fig. 15. Distribution of Calathus cinctus in The Nether- lands. Symbols on this and the following maps: open circles: recorded before 1930 only; small dots: recorded between 1930 and 1960; large dots: recorded since 1960. or wing-polymorphic with a high frequency of ma- cropterous beetles, but never or rarely caught fly- ing) with uncertain dispersal power. The wing-dimorphic cinctus was caught in win- dow traps (van Huizen 1980; den Boer et al. 1980) and consequently belongs to the group of B-species (van Dijk 1986). The wing-dimorphic melanocephalus and the monomorph macropterous mollis were never caught flying, but laboratory-bred beetles of both species developed flight muscles and flew fre- quently in a laboratory test (to be published else- where). For the time being both species therefore have to be considered C-species. Den Boer (1977) Table 13. Emergence of adult beetles in outdoor breeding experiments. N: number of beetles. SPECIES cinctus melanocephalus mollis Year Decade N N N 1982 1-10 June 3 ( 1.8 %) 113 (43.3 %) 158 (98.7 %) 11-20 June 136 (80.5 %) 101 (38.7 %) 2 (I Gy) 21-30 June 29 (17.2 %) 40 (15.3 %) = = 1-10 July l ( 0.6 %) 7 (27%) = = 1983 1-10 June 2 > = 3 a = 11-20 June 69 (53.9 %) 12 (34.3 %) 32 (68.1 %) 21-30 June 58 (45.3 %) 23 (65.7 %) 15 (31.9 %) 1-10 July 1 ( 0.8 %) = = = = 136 Fig. 16. Distribution of Calathus melanocephalus in The Netherlands. and van Dijk (1986) treated melanocephalus as A- species, based on the extremely low frequency of macropterous beetles in populations investigated in the Dutch province of Drenthe (0.23 %). Popula- tons in the recently reclaimed Flevopolders, how- ever, showed a considerable higher freqency of ma- cropterous beetles (25-40%: den Boer 1970; Auke- ma 1986). During breeding the flight muscles are autolysed in these species and flight only occurs before breed- ing. Data on the development of flight muscles and the occurrence of actual flight from the field and from breeding experiments in relation to temper- ature and availability of food will be treated else- where (Den Boer et al. (1980) already gave some preliminary data for cinctus). Distribution Distribution in The Netherlands Available data were used to prepare distribution maps on the UTM 10 X 10 square km scale as used by the European Invertebrate Survey in The Ne- therlands (figs. 15-17). Calathus cinctus (fig. 15) and C. melanocephalus (fig. 16) are widely distributed over the whole coun- try, whereas C. mollis (fig. 17) is almost restricted to the coastal dune region, with only a few inland populations on blown sands or other large sand deposits. Desender (1985, 1986) and Moncel (1972) AUKEMA: Three species of Calathus Fig. 17. Distribution of Calathus mollis in The Nether- lands. published comparable maps and data for Belgium and France respectively. A closer examination of the data of cinctus (fig. 15) shows, that this species has been caught regu- larly in The Netherlands since 1864 (Overveen, vi, 1 female, leg. J. Kinker, coll. Zodlogisch Museum, Amsterdam). For melanocephalus and mollis re- spectively the first records are Overveen, x.1864, J. Kinker, 1 male (coll. Zodlogisch Museum, Amster- dam) and Driehuizen, 23.vii.1882, 1 male (coll. Zoölogisch Museum, Amsterdam). Time-trends in the number of 10 X 10 km squares occupied per decade by the three species were estimated according to the method of Turin & den Boer (1988). Fig. 18 shows the time-trends after correction of the number of occupied squares for differences in collecting intensity between the decades (table 14). Calathus melanocephalus shows a significantly increasing time-trend, whereas both cinctus and mollis show rather indifferent time trends (fig. 18). When considering the 20th century only (table 14: decades 4-12) all three species show negative correlation coefficients, but only in the case of cinctus was this significant, largely due to the high number of squares occupied in the years 1910- 1930. Ranges Concerning their general distribution, revision of material on a larger scale is necessary to get a 15% TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 14. Number of 10 x 10 km squares occupied per decade by Calathus cinctus, C. melanocephalus and C. mollis. TO: total number of squares occupied by all carabid species per decade; F: correction factor calculated from the total number of squares covering The Netherlands (F=448/TO); O: actual number of occupied squares; OF: corrected number of occupied squares (OF=O%F). Decade cinctus melanocephalus mollis Nr Years TO F O OF O OF O OF 1 1870 - 1879 72 6.22 2 12 0 0 2 12 2 1880 - 1889 81 399) 1 6 1 6 3 17 3 1890 - 1899 108 4.15 5 21 7 29 5 21 4 1900 - 1909 150 3.03 16 48 18 55 6 18 5 LOOS IOI 1972 255 26 61 24 56 11 26 6 1920-1929 214 2.10 22 46 28 59 9 19 7 1950219 95215 2.10 18 38 29 61 15 32 8 1940 - 1949 246 1.82 12 22 26 47 7 15 9) 1950-1959 264 1.70 16 27 35 59 5 22 10 1960 - 1969 312 1.44 9 13 20 29) 10 14 Ul 1970-1979 336 135 23 31 35 47 12 16 12 1980 - 1989 350!) 1.28 21 27 33 42 9) 11 ') Estimated value. Up till now 14714 records from 321 squares were recorded, but many have to be added (H. Turin, personal communication) reliable picture (the data in Turin (1981) are only based on literature). In the Dutch collections men- tioned and/or in the collections of the British Mu- seum (Natural History), London, the Zoologisches Museum, Berlin, the Zoologische Staatssammlung, Miinchen, and M. Baehr, Miinchen, I have seen material from the following localities: C. cinctus: South-England (south of the line River Seven -The Wash), Wales, Ireland, The Ne- therlands, Belgium, Germany, Poland, the Baltic States (Estonia), Russia, France, Spain, Sardinia, Corsica, Italy, Sicily, Malta, Yugoslavia, Austria, Hungary, Romania, Bulgaria, Albania, Greece, Cy- prus, Turkey, Crimea, Ukraine, the Caucasus, Iran, Syria, Israel and Morocco; C. melanocephalus: Finland, Sweden, Norway, Denmark, Iceland, England, Scotland, Wales, Ire- land, The Netherlands, Belgium, Luxembourg, Ger- many, Poland, the Baltic States (Estonia), France, Spain, Sardinia, Corsica, Italy, Sicily, Yugoslavia, Switzerland, Austria, Hungary, Romania, Albania, Greece, Turkey, Ukraine and the Caucasus; C. mollis: Denmark, England, Wales, Scotland, Ireland, The Netherlands, Belgium, Germany (the East Frisian Islands), France, Portugal, Spain, Mal- lorca, Italy, Sicily, Yugoslavia, Greece, Turkey, Gi- braltar, Malta, Algeria and Morocco. Lindroth (1986) indicated the occurrence of cırc- tus in South-Sweden and Denmark (see also Bangs- holt 1983) and of mollis in Norway. Since Lindroth listed the right combination of characters for both taxa, his findings, although not confirmed, are con- sidered reliable. Anderson (1987, in litt. 1990) furthermore found cinctzs to be sporadically dis- tributed around the Irish coast south of the line from the Shannon Estuary to Dublin. Calathus melanocephalus obviously has a Euro- 138 number of corrected squares 60 cinctus 40 20 melanocephalus 60 40 20 mollis 60 40 20 oO N - (CA oo 2 decade Fig. 18. Plots of the corrected time trends of Calathus cinctus, C, melanocephalus and C. mollis; decades: 1870- 1879 (1), 1880-1889 (2), 1890-1899 (3), etc. pean distribution, ranging from Iceland and Scan- dinavia in the North to the Mediterranean in the South. It is the only species of this group also oc- curring at higher altitudes in mountain areas. The subspecies alpinus Dejean is restricted to the east- ern Alps (Koch 1989). Ball & Negre (1972) list a probably introduced specimen of melanocephalus from North America. Calathus cinctus does not occur as far north as melanocephalus, but in the Mediterranean it reaches further south, occurring as far as Morocco and Algeria in the west and Syria and Iran in the east. Calathus mollis seemingly is restricted to the Atlantic and Mediterranean coastal areas. DISCUSSION The establishment of Calathus (Neocalathus) cinctus as a good species has solved a major iden- tification problem in the genus Calathus, although even now long winged females of cinctus are diffi- cult to distinguish from those of mollis. On the other hand, a new problem arises in the interpre- tation of existing literature on the species of the melanocephalus-group. It is quite clear from the data presented here that cnctus, although in liter- ature still considered to be mainly distributed in Eastern Europe (southeast and east of the river Weser) (Lienemann 1981; Koch 1989), is a very common species in Western Europe too. For in- stance in England circtus was already present at the time of Stephens, because in his collection, bought in 1853 by the British Museum (Natural History), London, and kept in its original state (Hammond 1972) four out of the 13 specimens in the box of melanocephalus belong to cinctus (1 male and 3 females, all short winged). Furthermore Jeannel (1942) already indicated the occurrence of cinctus in western France (Vendée). Lindroth (1949) used data on wing development in Calathus mollis as one of the main illustrations for his theory of postglacial recolonization of Fen- noscandia. Although Lindroth indicated in the same publication that there were two taxa involved, his data were never revised. Also the data from the following studies will have to be reconsidered con- cerning species identity: for larval taxonomy Table 15. Spearman rank correlation coefficients be- tween the corrected number of squares occupied and the decade rank Species All decades 20th Century!) cinctus 021 -0.74* melanocephalus +0.72* -0.54 mollis -0.20 -0.60 *) 0.02 RP G Su ) < I, dj m) a Figs. 47-55. Pupa of Stictochironomus obscurus (Guha & Chaudhuri) comb.n. 47, Frontal apotome; 48, antennal sheath of male; 49, antennal sheath of female; 50, thoracic horn; 51, abdomen; 52, tergite II; 53 anal fin and genital sac of male; 54, anal fin and genital sac of female; 55, caudolateral spur of segment VIII. 188 CHAUDHURI & CHATTOPADHYAY: Chironomids > AW „ul = 7 64 Figs. 56-66. Larva of Stictochironomus obscurus (Guha & Chaudhuri) comb.n. 56, Head capsule; 57, antenna; 58, S I; 59, pecten epipharyngis; 50, premandible; 61, mandible; 62, prementohypopharyngeal complex; 63, mentum; 64, ventromental plate; 65, posterior abdominal segments; 66, claws of posterior parapods. 189 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 67-73. Adult of Clinotanypus fuscosignatus (Kieffer). 67, Wing; 68, fore tibial spur; 69, mid tibial spur; 70, abdomen; 71, male hypopygium; 72, gonostylus of male; hypopygium; 73, female genitalia. 190 CHAUDHURI & CHATTOPADHYAY: Chironomids VA, MR À. Zi Z ZA SS Figs. 74-80. Pupa of Clinotanypus fuscosignatus (Kieffer). 74, Frontal apotome; 75, antennal sheath of male; 76, antennal sheath of female; 77, thoracic horn; 78, abdomen; 79, anal fin and genital sec of male; 80, anal fin and genital sac of female. WO TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 81-90. Larva of Clinotanypus fuscosignatus (Kieffer). 81, Head capsule; 82, antenna; 83, apex of antenna; 84, mandible; 85, maxilla, 86, apex of maxilla; 87, ligula and paraligula; 88, mentum and M appendage; 89, posterior abdominal segments; 90, claws of posterior parapods. 192 CHAUDHURI & CHATTOPADHYAY: Chironomids Figs. 91-97. Adult of Procladius noctivagus (Kieffer). 91, Wing; 92, fore tibial spur; 93, mid tibial spur; 94, hind tibial comb; 95, abdomen; 96, male hypopygium; 97, female genitalia. 195 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 98-104. Pupa of Procladius noctivagus (Kieffer). 98, Frontal apotome; 99, antennal sheath of male; 100, antennal sheath of female; 101, thoracic horn; 102, abdomen; 103, anal fin and genital sac in male; 104, anal fin and genital sac in female. 194 CHAUDHURI & CHATTOPADHYAY: Chironomids Figs. 105-112. Larva of Procladius noctivagus (Kieffer). 105, Head capsule; 106, antenna; 107, mandible; 108, maxilla; 109, ligula and paraligula; 110, mentum and M appendage; 111, posterior abdominal segments; 112, claws of posterior parapods. 195 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 196 HENK vAN MASTRIGT Wamena, Indonesia NEW (SUB)SPECIES OF DELIAS FROM THE CENTRAL MOUNTAIN RANGE OF IRIAN JAYA (EEPID OPMER A EIB REID ANE) Mastrigt, H. J. G. van, 1990. New (sub)species of Deltas from the central mountain range of Irian Jaya (Lepidoptera: Pieridae). - Tijdschrift voor Entomologie 133: 197-204, figs. 1- 17. [ISSN 0040-7496]. Published 14 December 1990. Delias sinak sp. n., D. sigit sp. n., D. bobaga sp. n. and D. catocausta eefi subsp. n. are described from the western part of the central mountain ranges in Irian Jaya, Indonesia. The recent acquisition of new Delias material from Abmisibil in the Star Mountains of Irian Jaya, leads to the recognition of the following new taxa: D. oktanglap sp. n., D. wollastoni abmistbilensis subsp. n., D. luctuosa kuning subsp. n. and D. nieuwenhuisi poponga subsp. n. Deltas nieuwenhuisi nom. nov., a replacement name for D. leucias roepkei Nieuwenhuis & Howart (nec Benneth, 1956), is raised to specific rank. Henk van Mastrigt, Kotak pos 25, Wamena 99501, Indonesia Keywords. - Lepidoptera; Pieridae; Delias; new species; Irian Jaya; Indonesia. From the beginning of this century quite a few expeditions succeeded to enter the interior of Irian Jaya (former Dutch New Guinea). The butterflies of the genus Delias collected during these expedi- tions provided material for the description of new species from the Arfak Mountains, Weyland Moun- tains, Wissel Lakes, the surroundings of the Car- stensz Peak, the Baliem Valley, Mt. Goliath and the Star Mountains. The Star Mountains in Irian Jaya are one of the central mountain ranges of which no reports on Deltas have been published. The results of the Dutch expedition to the Star Mountains in 1959 were poor and never published, as far as Delias is concerned. In fact, most information about Delias from the Star Mountains concerns the Papua New Guinea part of this range, where the late Ray Straat- man and others collected at Telefomin, close to the Indonesian border. Results from the Papua New Guinea part of the Star Mountains are found in RMNH, BMNH and several private collections. Since 1978 I have collected in the Star Mountains of Irian Jaya (fig. 17). The weather in these moun- tains is often cloudy and rainy, so that only small numbers of Deltas can be collected in a short time, contrary to more western ranges of the central mountains where the weather is usually sunnier. Besides that, the mountains are rough and steep, which does not make travelling easy. Further exploration of the extensive central mountain range of Irian Jaya, with its isolated val- leys led to the discovery of new taxa. In this paper three new species and one new subspecies from the western part of the central mountain ranges are described. My recent acquisi- tions of Delias from the Irian Jaya Star Mountains and study of additional museum material led to the recognition of one new species and three new sub- species from this area. Finally, a previously des- cribed subspecies is raised to specific rank. Depositories The abbreviations given below have been used in the list of material and throughout the text. BT - Private collection of Bernard Turlin, Andrésy, France; CT - Private collection of Collin Treada- way, Limbach, BRD.; DM - Private collection of David Mannering, Staplehurst, Kent, U.K.; DR — Private collection of David Rushen, E. Sussex, U.K.; ERII - Private collection II of Eduard Reissinger, Kaufbeuren, BRD.; GG - Private collection of Fred Gerrits, Brisbane, Australia; HM - Private collec- tion of the author, Jayapura, Indonesia; MZB - Museum Zoologicum Bogoriense, Bogor, Indone- sia.; RG - Private collection of Robert Gotts, Sid- ney, Australia.; RMNH - Nationaal Natuurhisto- risch Museum (Rijksmuseum van Natuurlijke Historie), Leiden, The Netherlands; ZMA - Insti- tuut voor Taxonomische Zoölogie (Zoölogisch Mu- seum), Amsterdam, The Netherlands. 197 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 1-4. New (sub)species of the genus Delias; uppersides of male holotypes. 1, D. simak; 2, D. sigit; 3, D. bobaga; 4, D. catocausta eeft. Figs. 5-8. New (sub)species of the genus Delias, undersides of male holotypes. 5, D. sinak; 6, D. sigit; 7, D. bobaga; 8, D. catocausta eeft. 198 VAN MASTRIGT: New Delias from Irian Jaya Figs. 9-12. New (sub)species of the genus Delias, uppersides of male holotypes. 9, D. oktanglap; 10, D. wollastoni abmisibilensis; 11, D.luctuosa kuning; 12, D. nieuwenhuisi poponga. Figs. 13-16. New (sub)species of the genus Delias; undersides of male holotypes. 13, D. oktanglap; 14, D. wollastoni abmisibilensis; 15, D.luctuosa kuning; 16, D. nieuwenhuisi poponga. 199 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 DESCRIPTIONS Delias sinak sp. n. Ges 1D 17) Type material. - Holotype @: ‘Irian Jaya, route Mulia — Sinak, vii.1989; don. Fr. Gerrits’, ZMA. - Paratypes: same data as holotype, 2 6, GG; same data, but vii.1990, 6 3, GG; 4 6, HM; Mulia, ix. 1990, 7 6, GG; 3 6 MZB. Diagnosis This species belongs to a subgroup of rather small Delias, consisting of Deltas sagessa Fruh- storfer, 1910 from East Papua New Guinea (includ- ing its subspecies straatmani Schröder, 1977 from Telefomin, Star Mountains and anjae Schröder, 1977 from the Arfak Mountains) and Deltas abro- phora Roepke, 1955 from the Paniai-district. These species are characterized by the black border of the hind wing underside, bearing five rounded red spots. The new species differs from these by the absence of the red spot at the base of the yellow (and white) discal band in the hind wing underside. Description Male. - Upperside fore wing white. Black border, with regularly curved inner edge, from discal cell at vein M, to margin border at 2A. Upperside of hind wing translucent white with narrow (1.5 mm) black border. Underside of fore wing black with a white area (some grey-black diffusion), from middle of tornus to vein CU,, sometimes entering discal cell, three subapical yellow spots followed by two mar- ginal ones. Underside of hind wing black, 5 mm wide, yellow discal band from middle of tornus to vein 2A, becoming narrower from vein M,, not reaching anal border. Terminal border with five large, more or less round, red spots, slightly variable in size and shape. Length of fore wing: 22 mm. Female unknown. Derivation of name. - ‘Sinak’: a noun in apposi- tion. This butterfly occurs along the route from Mulia to Sinak and received its name from the latter locality. Delias sigit sp. n. (igs) 25, Oxley) Type material. - Holotype @: ‘Irian Jaya/Centr. Berg- land/Ilaga, River Jila/ 7.vi.1988/Henk van Mastrigt’ ZMA. - Paratypes: same data as holotype, 2 6, HM; same data but 9.iv.1988, 5 g, ZMA; 6 4, MZB; 22.v.1988, 4 4, HM; 2 6, BT; 27.viii.1988, 2 4, MZB; 2 6, HM; 1 6, BT; 2.v.1990, 2 6, HM; 4.v.1990, 3 6, HM; Ilaga, River Mum, 22.1.1989, 1 8, ZMA; Ilaga, 10.viii. 1988, 1 6, MZB; 1 4, HM; 1 6, BT; 1 4, GG; Mulia-Sinak, vi-vii.1989, 2 8, GG; Beoga, x11.1989, 7 8, GG; 1 6, HM; Mulia, ix. 1990, 10 6, GG. 200 Diagnosis This species is similar to Delias mariae Joicey & Talbot, 1916 and D. bobaga because of its its bright and broad distal band on the hind wing underside, which is white with large orange patches. It differs from D. bobaga in lacking the large number of yellow scales in cells 2A and 3A of hind wing un- derside. It differs from D. mariae in having a yellow basal streak, which is connected to the white costal border. The basal streak of D. mariae sometimes with a small whitish edge, but the costa is black instead of white. Description Male. - Upperside of fore wing milky white with broad black costal border, entering discal cell, black LDC. Black border along termen, much broader at apex than at tornus and has slightly serrate inner edge, often with 1 or 2 subapical spots; second one very small. Upperside of hind wing milky white and translucent, narrow black border; whole pattern of underside visible from above. Underside of fore wing milky white with black border narrower than on upperside, three yellow subapical and one or two very small terminal spots. Large anterior part of discal cell black, passing LDC. Base of wing with some black and some blackish diffusion. Underside of hind wing yellow to yellowish orange with white veins, rounded black submarginal spot reaching LDC. Broad black anal part with some greyish green diffusion, connected to white costa, cutting off a yellow basal streak. Black border from SC+R, to 2A increasing in width from less than 1 mm to 3 mm. Length of fore wing: 25-29 mm. Female unknown. Derivation of name. — ‘Sigit’, a noun in apposi- tion. ‘Sigit-sigit’ is the vernacular word for but- terfly, used by the Dani people in the Ilaga valley, where this butterfly occurs. Delias bobaga sp. n. (figs. 3, 7, 17) Type material. - Holotype @: ‘Kamu Valley, 1700 m, ix.1989, don. Henk van Mastrigt, ZMA. — Paratypes: same data, 1 g, MZB; 1 8, GG; 1 6, DM; Kamu Valley: Ekemanida, River Tuka & Enau, 1700-1800 m, 23- 30.x.1990, 3 &, HM; River Mouna, 1.11.1990, 5 À, GG; River Epugewo, 2.11.1990, 1 6, GG; Idadagi, River Migu, 21.11.1990, 1 @, HM; Idadagi, River Bedonaka, 20.iv.1990, 1 4, HM; River Budau, 10.xi.1989, 1 4, GG; River Ode, 2.11.1990, 3 8, GG; Mapia area: River Okagou, 7.xii. 1989, 1 8, GG; 17.xii.1989, 1 4, HM; River Otika, 8.xii. 1989, 1 6, HM; River Yatuwou, 9.xii.1989, 1 6, GG; River Gudopena, 11.xii.1989, 1 6, GG; River Ude, 13.xii.1989, 1 6, GG; River Piyakoywa, 1 8, GG; River Okagou & Yatuwou, 25-26.xi.1989, 1 6, HM; Mapia, River lyage, 13.xii. 1989, 1 6, HM. Diagnosis This species is closely related to D. sigit sp. n., Delias mariae Joicey & Talbot, 1916 (including its subspecies D. mariae boschmai Roepke, 1955) and D. mariae menooensis Joicey & Talbot, 1922, with which it occurs together in the Kamu Valley. How- ever, it differs from those in the large number of yellow scales in the cells 2A and 3A and in a much larger basal spot on the underside of the hind wing. Description Male. - Upperside of fore wing white to creamy white with broad, black costal border, entering dis- cal cell, black LDC. Black border along termen, much broader at apex than at tornus and has a strongly serrate inner edge, sometimes absorbing black LDC. Upperside of hind wing creamy white and translucent with narrow black border; whole pattern of underside visible from above. Underside of fore wing milky white with black border, more reduced than on upperside, three red subapical spots and a red terminal one. Anterior part of discal cell black, via M, connected to black border, separ- ating a small, yellowish spot. Underside of hind wing yellow with white veins and black submargi- nal spot just outside discal cell. From middle of costa a broad, curved, black line extends through inner part of discal cell along yellow green anal area to the bottom of cell CU, where it is connected by a thin black line to the broad, black border, which runs from RS to 3A. Inner part of costa white. Base yellow with small black triangle at inner part. Length of fore wing: 27-29 mm. Female unknown. Derivation of name. - ‘Bobaga’, a noun in appo- sition, is the vernacular word for butterfly used by the Ekagi people in the Paniai district, where this butterfly occurs. Delias catocausta eefi subsp. n. (figs. 4, 8, 17) Type material. - Holotype @: Irian Jaya/Tembagapura 2000 m/17.xi. 1979/Van Mastrigt’, ZMA. - Paratypes: same data, 1 8, DR; same data but 17.xi.1979, 1 6, HM; 1 4, MZB; Tembagapura, 2000 m, 1980, 2 6, RG; 1983, 1 8, GG; Tembagapura, River Wanogong, 2000 m, 1989, 16, HM. Diagnosis Apart from the nominate form D. catocausta catocausta Jordan, 1912, only D. catocausta niger- rima Roepke, 1955 is known, from the Andrea River, just 50 km west of Mt. Goliath (c. 140° E.). The newly described subspecies is also from the southern side of the central mountain range of Irian Jaya, but occurs much more to the west: on the south sides of the Puncak Jaya (Carstensz Peak), c. 137° VAN MASTRIGT: New Delias from Irian Jaya E. It differs from the two above mentioned ones in the grey (instead of brown) ground colour on the underside of fore and hind wing, in the red (instead of yellow) colour of the basal spot on hind wing underside and in having white veins. Description Male. — Upperside of fore wing milky white, black tornus, thick, black LDC and black border along termen; this black border is much broader at apex than at tornus, has serrate inner edge and bears 2 poorly developed subapical spots. Basis of fore wing with some blackish diffusion. Upperside of hind wing milky white with very narrow black border. Underside of fore wing grey at innerside; the grey colour is connected by black veins to black border along termen; black border with three red subapical spots and two very small terminal ones. Inner edge of border serrate. A white band with some grey diffusion separates dark inner part from black border, widening from less than 1 mm on top to 7 mm along tornus. Underside of hind wing grey with white veins and small black spot close to discal cell on vein M,. Cell along costa, inner part of discal cell and cells 2A and 3A form a black hook with small red basal spot and greyish green diffusion along anal area in 3A. Narrow black border from cell RS to anal area is undulate at inner edge and separated from grey discal part by white line in cell RS, getting broader in M,, M,, CU, and somewhat narrower again in CU. Border between this white line and grey innerpart vague and diffuse from M, LCOS, Length of fore wing: 27 mm. Female unknown. Derivation of name. - This subspecies is named in honour of my father, the late Mr. Eef van Mas- trigt, who awakened my interest in nature and es- pecially in butterflies. Delias oktanglap sp. n. (figs. 9, 13) Type material. - Holotype @: ‘Irian Jaya/Sterren Ge- bergte/ Abmisibil 1920 m/ River Oktanglap/27.ii.1989/- Henk van Mastrigt’, ZMA. - Paratypes: same data, 3 @, ZMA; same data, but 11.ix.1985, 4 &, HM; 12.ix.1985, 1 6, HM; 14-16.ix.1985, 6 @, ZMA; 2 6, HM; 1 6, ZMA; 6.v1.1986, 4 4, RMNH; 29.iv.1986, 1 4, HM; 2.v.1986, 1 3, HM; 22.x.1986, 4 &, RMNH; 2 3, HM; 17.vii.1987, 3 6, HM; 11.iii.1988, 2 4, MZB; 16-19.iii.1988, 7 6, ZMA; 3 d, BT; 20.11.1988, 2 6, HM; Batimban, River Okkim, 1850 m, 14-16.11.1987, Van Mastrigt, 6 @, RMNH; same data, but ix.1987, 2 @, ZMA; 19-21.iii.1988, 2 &, ZMA; 13- 15.iv.1988, 1 4, BT; v.1988, 2 6, MZB; 19-20.viii.1990, 2 8, HM; Abmisibil, River Takpalngi, 17-25.ix.1988, 2 3, MZB; 1 8, BT; Abmisibil, River Okpeti, 1800 m, 14.ix.1986, 1 g, HM. 201 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Diagnosis This new species belongs to the cuningputi group, consisting of D. cuningputi (Ribbe, 1900), D. konokono Orr & Sibatani, 1986, D. chimbu Orr & Sibatani, 1986, D. fascelis fascelis Jordan, 1912, D. fascelis ibelana Roepke, 1955, D. citrona Joicey & Talbot, 1922 and D. jordanı Kenrick, 1909, as de- fined by Orr and Sibatani (1986). It is distinguished from all those by its very reduced black border on the fore wing upperside and from cuningputi and konokono by its continuing submarginal light band on the hind wing underside. Description Male. - Upperside of fore wing white, black border from 1.5 mm outside discal cell to tornus, inner edge straight and slightly serrate. Very thin bar at top of discal cell. Upperside of hind wing white with black border from SC+R, to CU,, 3 mm wide in M,, narrowing to both ends. Underside of fore wing white in bottom 1/3, with black diffusion. Large black border connected with nearly totally black discal cell, enclosing three subapical yellow spots and three yellow to white marginal spots — with thin white line to margin border. Veins be- tween discal cell and border partly black. Underside of hind wing black, divided by an irregular, 2-6 mm wide, curved white discal band, broken in M,. Ter- minal border white with a yellow basal streak. Inner part of hindwing black with two yellow spots and some yellowish diffusion. On white discal band one yellow spot just above black connection, five yellow streaks in underpart. Black border with undulate edge at innerside, having a row of six yellow spots in a line with the above mentioned yellow spot and streaks on white band. Length of fore wing: (25) 28 — 29 mm. Female unknown. Derivation of name. - ‘Oktanglap’ (a noun in apposition) is the name of the creek from where the holotype originates. Delias wollastoni abmisibilensis subsp. n. (figs. 10 & 14) Type material. - Holotype &: ‘Irian Jaya/Abmisibil 1920 m/River Oktanglap/ 11.1x.1985/Van Mastrigt’, ZMA. - Paratypes: same data as holotype, 1 8, HM; same data, but 1880-1920 m, 15.ix.1985, 1 4, HM; 22.x.1986, 1 3, MZB; 1 4, HM; 17-19. vii.1987, 1 6, ZMA. Diagnosis Adding this new subspecies from the Star Moun- tains in Irian Jaya, three separated populations of Delias wollastoni Rothschild, 1915 are now known, the other two being: D. wollastoni wollastoni from the southside of the Puncak Jaya (Carstensz Peak) (1 male in BMNH) and Beoga (1 male in GG) and D. wollastoni bryophila Roepke, 1955 from the 202 surrounding of Mt. Trikora (Mt. Wilhelmina) (good series of males in RMNH and ZMA and two males in HM). The new subspecies differs from the other two subspecies by its golden yellow colour of the fore wing underside and more vivid coloration of hind wing underside. Description Male. - Upperside of fore wing white, greyish black veins and some grey diffusion. Costal border black. Black border from top of discal cell to tornus encloses black LDC, two white subapical spots, two or three small terminal ones. Border reduced in cells M,, CU, and CU, and slightly entering along dor- sum. Upperside hind wing white with grey veins and some grey diffusion, particularly at inner edge of broad black border in cell M, and M,. Underside of fore wing golden yellow, black border much nar- rower than on upperside, regularly curved at inner edge, encloses three large yellow subapical spots and five smaller yellow terminal ones. Costal border black; at top of LDC large black triangle which is narrowly connected to black border. Underside of hind wing with base and basal half of costa grey, bordered by SC+R,. Large basal maroon coloured patch below SC+R,, oblong patch on LDC and bowed, undulate postmedian band of same colour. Postmedian band consists of intervernal patches from RS to CU,. Anal area densely covered with golden yellow scales; grey becoming pinkish grey in outer third of discal cell and in adjacent cells. A strong and rather straight median band extends from costa to 1A+2A, including the maroon brown streak on LDC. Space between this median and postmedian band filled with light pinkish grey, den- sely covered with dark grey-brown diffusion. Sub- terminal area light pinkish grey with darkened ter- men, more or less dusted with golden-yellow scales mainly on tips of veins. Length of fore wing: 29 mm. Female unknown. Derivation of name. - Abmisibil is the name of the village in the Star Mountains, which is close to the sites where this butterfly occurs. Delias luctuosa kuning subsp. n. (figs. 11 & 15) Type material. - Holotype @: ‘Irian Jaya/Star Moun- tains/ Abmisibil 1800 m/River Oktero/ 11.1x.1985’, ZMA. — Paratypes: Abmisibil, River Okmi, 31.iii.1985, Sam Mo- nareh; 1 6, HM; Abmisibil, River Oktanglap, H. van Mastrigt, 1 @, HM; Langda, River Bibwe, 1800 m, 6.iv.1989, H. van Mastrigt, 1 d, HM; same data, but 17- 19.iv.1989, 1 6, HM. Diagnosis Three populations of D. lwctwosa Jordan, 1912 have been named previously, viz. D. /uctuosa arch- boldi Roepke, 1955 from the Baliem Valley (also from the more western Ilaga Valley), D. lwctuosa luctuosa from Mount Groome, Papua New Guinea, and D. luctuosa mizukamit Okana, 1989 from Simbu Province, Papua New Guinea. With the new population from Langda and Abmisibil a bridge is made between the population from central Irian Jaya and the two populations from Papua New Guinea. The new subspecies differs from the three above mentioned in the golden-yellow colour of the male fore wing underside. Description Male. - Upperside of fore wing white with black costal border, slightly entering discal cell. Terminal border very broad in cells M,, M, and M;, narrower towards tornus, where it slightly curves along dor- sum, provided with irregular inner edge, enclosing two small white subapical spots. Veins visible, par- ticularly those on underside of discal cell. Upperside of hind wing translucent white with black border, which is narrow in cell RS and M,, undulate at vein M, and M; and broad in cells CU,, CU, and 2A. Underside of fore wing deep yellow, turning to white along dorsum, black border along inner edge in cell 2A. Costal border black. Large LDC con- nected to black border in cell M;, demarcating two yellow spots with black diffusion in cells M, and M,. Black border encloses three dark yellow subapical spots and two or three small terminal spots. Under- side of hind wing with white ground-colour. Black costal border connected to broad black anal border, including small yellow pear-shaped basal spot. Postdiscal black band (width 3 mm) curving from anal border to costa at vein SC+R, and entering broadly cell RS; outer edge bordered with small red un + è > => f Se AG € S. LE N: CH 3 c , de b 8 Ò o h eN @ EN ©. VAN MASTRIGT: New Delias from Irian Jaya line. White discal part with pink diffusion, darker at innerside; veins white. Discal border consists of five black triangles at veins, touching red line and demarcating six white, triangular to v-shaped, spots. Length of fore wing: 27-29 mm. Female unknown. Derivation of name. — A noun in apposition. ‘Kuning’ is the Indonesian word for yellow and refers to the yellow underside of the fore wing, the characteristic feature of this butterfly. Delias nieuwenhuisi nom. nov. Delias leucias roepkei Nieuwenhuis & Howart, 1969: 86. Holotype 4: Mandated New Guinea, Telefomin, 1700 m, 5.viii. 1962 (Natuurhistorisch Museum, Rotterdam) (preoccupied by D. mira roepkei Benneth, 1956). Discussion Since the subspecific name roepkez in Delis leu- clas roepkei Nieuwenhuis & Howarth is preoccu- pied by Benneth’s name, the replacement name meuwenhuist is proposed here, in honour of Mr. E. J. Nieuwenhuis, who was the first to describe this taxon. Further, since nzeuwenhuisi and leucias s. str. are sympatric in the Star Mountains along river Okkim at Abmisibil, Irian Jaya and at Telefomin, Papua New Guinea, they cannot be regarded as subspecies of one species, therefore the first one is raised to specific rank: D. nieuwenhuisi. The differences between the specimens from Telefomin (types of nieuwenhuisi) and those from Abmisibil lead to the recognition of two subspecies: D. nieuwenhuisi nieuwenhuisi from Telefomin and D. nieuwenhuisi poponga subsp. n. from Abmisibil (see below). Fig. 17. Distribution of new (sub)species of the ge- nus Delias. a, Kamu valley (type locality of bobaga); b, Tembagapura (type locality of catocausta eeft); c, llaga (type locality of sgit); d, Sinak (type locality of sinak); e. Langda (para- types of /uctuosa kuning); f, Abmisibil (type locality of oktanglap, wollastoni ab- misibilensis and luctuosa kuning). 159) 03 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Delias nieuwenhuisi poponga subsp. n. (figs. 12, 16) Type material. - Holotype 8: ‘Irian Jaya/Sterren- gebergte/Batimban/ca 1900 m/ River Okkim, v.1988/ H. van Mastrigt’, ZMA. - Paratypes: same data, 9 8, HM; 1 é, ZMA; same data, but 24.xii.1985-7.1.1986, 3 6, HM; same data, but 1-5.v.1986, 14 6; 2 6, DR; 6 g, RMNH; same data, but 14-16.ii.1987, 6 @, ZMA; same data, but ix.1987,7 6, HM; 1 8, BT; 6 g, ZMA; same data, but 19- 21.iii 1988, 14 À, HM; 6 @, BT; same data, but 13- 15.iv.1988, 2 6, FG; 1 6, HM; 1 4, CT; same data, but 22.v- 5.vi.1988, 2 6, HM; same data, but 11-13.viii.1988, 4 3, HM; same data, but vi.1986, 2 6, ERII; same data, but 19- 20.viii.1990, 4 6, HM; 4 4, MZB; Takpalngi, 17- 25.ix.1988, H. van Mastrigt, 6 4, HM; Ferohir, 5.v.1986, H. van Mastrigt, 1 @, ZMA. Diagnosis This subspecies differs from nominate nieuwen- huisi in three characters. Upperside and underside of fore wing having narrow black borders (even narrower than in leucias); having upperside of fore wing with a black vein, connecting border with midcell and underside of hind wing with a large posterior red strike and a thin anterior white line in the basal spot. Description Male. - Upperside of fore wing white with nar- row black border, 5 mm in cell R, to 1 mm in cell CU. Inner edge slightly serrate, not curved from cell R, to tornus. Black vein R, connects border with discal cell, demarcating a small white spot. Apical and marginal spots absent. Upperside of hind wing white, 1-2 mm wide, black border. Underside of fore wing yellow, narrow black border as broad as on upperside, having four yellow apical spots. Ground- colour orange yellow in cells R,, M,, M,, M;, and CU,, lemon yellow in discal cell and pale yellow to creamy white in cells CU, and 2A. Underside of hind wing with large creamy discal area, covering nearly whole discal cell and large inner parts of cells RS, M,, M,, M; and CU,, just entering cell CU. Discal area surrounded by black line, which is less than 1 mm in cell RS, 2 mm in cell CU, and 3.5 mm in discal cell. Red subbasal mark usually oblong, sometimes more like an upside down ‘v. Al mm wide red line runs from cell SC+R, to CU,, parallel to and connected with black line which surrounds discal area. Discal border consists of a, 1.5 mm wide, white line, with at its outer edge a narrow black line from cell M, to CU,, which is more than 1 mm in cells M; and CU, only. Basal spot anteriorly white, posteriorly red. Anal margin appearing dirty pale yellow due to low number of black scales (compared to other species of this subgroup). Concentration of white scales along vein 3A divides yellow anal area. Length of fore wing: 22-24 mm. Female unknown. Derivation of name. - A noun in apposition. ‘Poponga’ is the vernacular word for butterfly used by the people in the western part of the Star Moun- tains, where this butterfly occurs. ACKNOWLEDGEMENTS I gratefully acknowledge the help of Dr. Fred Gerrits, Mr. David Mannering, Mr. Bernard Turlin and Mr. Collin Treadaway, Mr. David Rushen, who allowed me to study their private collections and the assistance of Dr. Eduard Reissinger and Dr. Atuhiro Sibatani. Many thanks are extended to the Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Amsterdam, for the good and helpful cooperation, especially to Mr. L. van der Laan for the preparation of the photographs. The cooperation with the Museum Zoologicum Bogo- riense, Bogor and the Department of Forestry, Irian Jaya, in the study of butterflies from Irian Jaya is gratefully acknowledged. REFERENCES Bennett, N. H., 1956. A new Deltas from the central highlands of New Guinea (Lepidoptera, Pieridae). — The Entomologist 89: 81-83, pl. 3. Fruhstorfer, H., 1910. Familie: Pieridae, Weisslinge. - In: A. Seitz, Die Gross-Schmetterlinge der Erde 9: 123- 135, 183-190, (1927) 1108. Joicey, J.J. & G. Talbot, 1916. New lepidoptera from Dutch New Guinea. - Annals and Magazine of Natural His- tory (series 8) 17: 68-72. Joicey, J. J. & G. Talbot, 1922. New forms of the genus Deltas (Pieridae) from New Guinea, Ceram and Buru. — Bulletin of the Hill Museum 1 (3): 303-319. Jordan, K., 1912. On new or little-known Delias from New Guinea. - Novitates Zoologicae 18 (1911): 580- DOSI Nieuwenhuis, E. J. & T. G. Howart, 1969. On some but- terflies from the Indo-Pacific region. - Entomolo- gische Berichten, Amsterdam 29: 85-88. Kenrick, G. H., 1909. Descriptions of some new species of the genus Delias from North New Guinea, recently collected by Mr. C. E. Pratt. - Annals and Magazine of Natural History (series 8) 4: 176-183, pls 6, 7. Okana, K., 1989. Descriptions of four new butterflies of the genus Deltas (Lep.: Pieridae), with some notes on Delias. - Tokurana 14 (3/4): 1-6. Orr, A. G. & A. Sibatani, 1986. A revision of the Delias aroae-cuningputi complex (Lepidoptera, Pieridae) 1. The D. cuningputi group. - Tyô to Ga 37 (1): 1-14. Ribbe, C., 1900. Neue Lepidopteren aus Neu-Guinea. — Insektenbörse 17: 329-330. Roepke, W. K. J., 1955. The butterflies of the genus Delias Hübner (Lepidoptera) in Netherlands New Guinea. — Nova Guinea, n. s. 6 (2): 185-260. Rothschild, W., 1915. — In: W. Rothschild and J. H. Dur- rant, Lepidoptera of the British Ornithologist's Union and Wollaston Expeditions in the snow mountains, southern Dutch New Guinea. - 168 pp. London, Ha- zell, Watson & Viney, Ltd. Schröder, H., 1977. Zwei neue Delias der sagessa-Gruppe aus Neuguinea (Lep., Pieridae). - Entomologische Zeitschrift Frankfurt am Mainz 87: 265-271. Received: 18 July 1990 Accepted: 14 September 1990 204 ERIK J. vAN NIEUKERKEN National Museum of Natural History, Leiden THE TRIFURCULA SUBNITIDELLA GROUP (LEPIDOPTERA: NEPTICULIDAE): TAXONOMY, DISTRIBUTION AND BIOLOGY Nieukerken, E. J. van, 1990. The Trifurcula subnitidella group (Lepidoptera: Nepticulidae): taxonomy, distribution and biology. — Tijdschrift voor Entomologie 133: 205-238, figs. 1- 108, tab. 1. [ISSN 0040-7496]. Published 14 December 1990. The nominal subgenus Trifurcula Zeller, 1848 is divided into two species groups: the subnitidella group and the pallidella group. The subnitidella group is described and com- prises T. subnitidella (Duponchel, 1843), and eight new species: austriaca (from Austria), luteola (France), puplesisi (Soviet Union), coronillae (Spain), victoris (Spain), josefklime- schi (Italy, southern Europe), iberica (Spain) and silviae (France). Males of all species and females of six species are described and diagnosed, and for four species the larvae, mines and biology are described as well: they make mines in stems of herbaceous or shrubby Fabaceae (tribes Loteae, Coronilleae). The group is widespread in the western Palaearctic, but most species occur in Southwest Europe. T. subnitidella is recorded for the first time from 14 countries. The phylogeny of the group and subgenus are discussed, the pallidella group is considered to be its sistergroup. A lectotype for T. orientella Klimesch, previously confused with some species here described, is designated. Dr. E. J. van Nieukerken, Rijksmuseum van Natuurlijke Historie, Postbus 9517, 2300 RA Leiden, Netherlands. Keywords. - Nepticulidae, stem-miners, taxonomy, phylogeny, Palaearctic, Fabaceae. The genus Trifurcula Zeller, 1848 constitutes an important part of the nepticulid fauna of the Me- diterranean region with the total number of species probably exceeding 60 (31 described species: van Nieukerken 1986a). It is especially abundant in dry open habitats, such as grasslands, maquis and gar- rigue vegetations, where the larvae make leaf- or stem-mines in shrubby and herbaceous plants, in particular Lamiaceae (Labiatae) and Fabaceae (Le- guminosae). A few species extend their ranges northwards into northern and western Europe. Tri- furcula remains the main nepticulid taxon in Eu- rope which needs extensive taxonomic revision. The genus was fully redescribed by van Nieu- kerken (1986b), who recognized three subgenera: Glaucolepis Braun, Levarchama Beirne and Trifur- cula Zeller. The ten Northwest European species have been treated by van Nieukerken & Johansson (1990), but no comprehensive work for the remain- ing European species is currently available. The present paper is the first in a series of revi- sions of Trifurcula, which follows a few papers dealing with single species (van Nieukerken & Jo- . hansson 1986, van Nieukerken 1987). The subge- nus Trifurcula was revised by Klimesch (1953), who recognized eight species, two of which are now considered to be junior synonyms. Only two species were added later, T. griseella Wolff, 1957 (later recognized to be a junior synonym of T. subnitidella (Duponchel, | 1843 ])) and 7. beirnei Puplesis, 1984, resulting again in a number of eight described spe- cies (van Nieukerken 1986a). These species only represent a minority of the actual number, which probably exceeds 25. The described species are almost all relatively uniformly coloured moths, without striking exter- nal characters, and associated with various species of brooms (Fabaceae, tribe Genisteae). T. subniti- della forms an exception in having a conspicuous external character — a brand of bright yellow scales on the male forewing underside — and having an association with herbaceous Fabaceae (Lotus: van Nieukerken & Johansson 1990). In unidentified and misidentified material from southern Europe and western Asia eight species with similar external features have been found, and following recent field trips three of these species and T. subnitidella itself, have been reared from larvae collected on plants belonging to the tribes Loteae and Coronilleae. In this paper these species are assembled into the T. subnitidella species group, whereas the other pa- laearctic species in Trifurcula s. str. (with the type species T. pallidella (Duponchel, [1843])) are con- sidered to form the T. pallidella species group. 205 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 When studying the material, representing the new species below, several specimens were found to be labelled as T. orzentella Klimesch, 1953. This led me to re-examine T. orientella and select a lectotype for this species, which clearly belongs to the pa//- della group. Other specimens treated here were found amongst material labelled as ‘T. immundella (Zeller)’, ‘T. pallidella Zeller’, ‘T. serotinella Her- rich-Schaffer’ and ‘Nepticula cryptella Frey’. Ap- parently, previous workers rarely examined the un- derside of the forewings of these tiny moths, which exhibit such good characters in this group. MATERIAL AND METHODS Larvae were collected by cutting portions of the stems of the hostplant, containing the mines. The 206 Figs. 1-6. Trifurcula spp., male habitus. - 1, T. austriaca, holotype; 2, T. luteola, paratype from type-locality; 3, T. coronillae, holotype; 4, T. victoris, holotype; 5, T. subnitidella, Spain, Camino de Ojen; 6, T. josefklimeschi, holotype. stems were put in plastic boxes with a layer of paper tissue, in which the larvae could spin their cocoons. Next the cocoons or full-grown larvae were trans- ferred into glass jars with a layer of soil and some moss. In the case of species from southern Europe, these jars were kept indoors (in The Netherlands), at room temperature, until emergence. Autumn col- lected larvae of subnitidella were, after spinning their cocoons, left in an outhouse for overwintering, and were brought indoors in March. Methods for preparation of the genitalia are lar- gely the same as in van Nieukerken (1985). In this group it is often very difficult to separate the aedea- gus from the genitalia capsule and valvae, because of the tight connection with membranes. The ae- deagus has therefore only been severed in some specimens, in order to study the complex morphol- VAN NIEUKERKEN: Trifurcula subnitidella group DEN Wh Ws Figs. 7-10. Trifurcula spp., males. - 7, T. silviae, habitus, holotype; 8, T. austriaca, underside, holotype; 9, T. subnitidella, underside, Spain, Camino de Ojen; 10, 7. coronillae, underside, holotype. Andraconial patch arrowed in figs. 8-10. ogy of aedeagal processes and cornuti; in some spec- imens also the vesica was removed from the aedea- gal tube to show the cornuti better. For identifica- tion purposes, removing of the aedeagus is not always necessary. Larvae were treated with KOH 10% prior to cleaning and mounting with Euparal on slides. They were stained with Chlorazol black. Line figures of genitalia were prepared with a Zeiss Standard microscope or Zeiss Axioskop, with camera and drawing apparatus, both from genitalia in glycerin and permanent mounts. In most cases cornuti are drawn separate from the aedeagus; only the characteristic larger cornuti are depicted, the smaller ones being almost similar in all species. Line drawings of mines were made from herbarium specimens, after boiling for some minutes in dis- tilled water: frass may be better visible in such soaked mines than in fresh ones. Photographs of genitalia and larval head capsules were taken with a Zeiss Axioskop microscope and camera. SEM micrographs were taken with a Jeol JSM 840A scanning electron microscope. Specimens were air-dried, mounted on stubs and gold-coated. Measurements of genitalia are taken with a Zeiss Axioskop or Universal Microscope, at 200 X, and are accurate at the nearest 5 um. All measurements based on sample size of at least five specimens are accompanied by mean, standard deviation and sam- ple size in brackets. Distribution maps are based on material exam- ined, but for T. subnitidella also literature data and personal communications have been used. Locality names are spelled in accordance with the Times Atlas of the World (Comprehensive edition, 1975 and later), deviating names on labels are cited in brackets. For all localities the appropriate UTM grid references (10 X 10 km, or 1 X 1 km for authors’ records) are given. These are used for plot- ting the distribution on the 50 X 50 km squares of the maps provided by the ‘European Invertebrate Survey’. The UTM grid references were obtained directly from ‘Tactical Pilotage Charts 1:500,000', topographical maps, or calculated from geographi- cal coordinates (obtained from atlases, various maps or US gazetteers), using the computer pro- gram ‘UTM’ by Rasmont & André. Most of the British records, however, are an approximation of the correct UTM square, since they are based on vice-county records and dots on a vice-county map, provided by A. M. Emmet (in litt.). Nomenclature of hostplants follows Heywood & Ball (1968), but the family name Fabaceae is used in stead of Leguminosae. Abbreviations (codens) for depositories follow Arnett & Samuelson (1986), with the addition of: ETO (for collection E. Traugott-Olsen, Marbella, 207 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 11-16. Trifurcula subnitidella, male, details of wings, scanning micrographs. - 11, forewing underside with yellow scale patch, some normal wing scales in upper right corner; 12, 13, details of androconial scales; 14, hindwing with frenulum and group of ‘black’ scales; 15, idem, detail; 16, ultrastructure of ‘black’ scale. bs = black scales; ns = normal scale. Spain), GBA (for collection G. Baldizzone, Asti, Italy) and ZKVV (Zoologijos Katedra VPI, Vilnius, Lithuania). Note that in the index of Arnett & Samuelson the coden NHMW is misspelled as NHMV. RMNH remains the coden for the Na- tional Museum of Natural History, Leiden, Nether- lands (formerly Rijksmuseum van Natuurlijke His- torie). Genitalia slide numbers refer to the numbering system of the author, unless accompan- ied by another abbreviation. 208 TAXONOMIC PART For descriptions of the genus Trifurcula Zeller, 1848 and its typical subgenus refer to van Nieu- kerken (1986b) and van Nieukerken & Johansson (1990). This subgenus is here divided into two spe- cies groups, recognized according to the following key: 1. Forewing underside in male with a patch of yellowish androconial scales near base (except in T. silviae); in addition hindwing costa often with row or group of dark brown or black spe- cial scales. Male genitalia: gnathos often atyp- ical: either with serrations, keels, projections or completely asymmetrical, occasionally simple and symmetrical. Aedeagus usually with dorsal lobe at right side; large cornuti (3) not curved and horn-like (except in awstriaca). Larvae stem-miners in tribes Loteae or Coronilleae (Rabaccao) enn: subnitidella group - Forewing underside and hindwing costa in male without androconial or special scales. Male genitalia: gnathos simple, symmetrical. Aedeagus usually with dorsal lobe at left side; at least one or two large cornuti strongly curved, horn-like. Larvae stem-miners in brooms: tribe Genisteae..... pallidella group The following nominal species are here assigned to the pallidella group: Trifurcula pallidella (Du- ponchel, [1843]) (= incognitella Toll, 1936), T. immundella (Zeller, 1839), T. serotinella Herrich- Schäffer, 1855 (= confertella Fuchs, 1895), T. orien- tella Klimesch, 1953, T. aurella Klimesch, 1953, T. beirnet Puplesis, 1984 and T. squamatella Stainton, 1849 (= maxima Klimesch, 1953). Also the undes- cribed species from Sumatra (see van Nieukerken 1986b) belongs here. This group will be the subject of a future revision. The Trifurcula subnitidella group Description Adult (figs. 1-7). - Forewing length 2-3.5 mm. Antenna with 27-48 segments, in male with more flagellar segments than in female. Forewing upper- side uniformly pale or irrorate by dark tipped scales, mixed with pale scales. No other colour pattern present. Venation as described for subgenus (van Nieukerken 1986b, van Nieukerken & Johansson 1990). Underside of forewing in male with a distinct patch of lamellar androconial scales near wing-base (figs. 8-10), usually yellow or yellowish white, oc- casionally with various tinges of yellow (this patch is absent in T. silviae). The patch consists of scales, which are ultrastructurally different from normal wing scales (fig. 11) (see also van Nieukerken et al. 1990). In szbnitidella these scales have almost no ultrastructure (figs. 12, 13), although remnants of scale-ribs are visible on some scales. In coronillae the three differently coloured areas also appear to have different ultrastructure: the basal area is much similar to the patch of szbnitidella (not illustrated), but the scales in the central area have an irregular pattern of ribs (fig. 19). Forewing costa often with VAN NIEUKERKEN: Trifurcula subnitidella group distinct costal fold on underside, in josefklimeschi extremely large (figs. 20, 58). Hindwing underside in male with terminal velvet-like patch of raised scales (generic character) (illustrated by van Nieukerken 1986b and van Nieukerken et al., 1990). Costa of hindwing in sev- eral species slightly enlarged (fig. 21), in some spe- cies provided with a short or longer row of distinct black androconial scales (figs. 57-60). These scales also differ ultrastructurally from normal wing scales (figs. 15, 16, 22). In fig. 15 both the black scales and normal wing scales are shown, the ultra- structure of the normal scale is illustrated in fig. 17. The androconial scales are completely jet-black, or dark-brown in sz/viae, and have spiny ribs, with the intercalary area almost devoid of structure, apart from some indistinct bulges (fig. 16). Male without costal bristles, but often with long white hair-scales instead. Male genitalia (figs. 23-88 in part). - Vinculum anteriorly rounded, truncate or slightly concave. Uncus with central process pointed or terminally widened, occasionally bilobed. Gnathos occurs in two forms: either symmetrical, with rounded or triangular central element, or asymmetrical and complex, provided with keel(s) or with serrate mar- gin or with a combination of both; anterior apo- demes usually present, but not always well sclero- tized. Valva elongate, more or less triangular, or in some species with broader basal part. Aedeagus joined by juxta (ventral process) to valvae and cap- sule. Aedeagal tube asymmetric, complicate, taper- ing posteriorly both in lateral and ventral view; ventrally ending in a pointed or serrate (fringed) medial carina, which hinges to the juxta, dorsally ending ina pointed or spatulate tip, often indistinct, and in addition usually with a dorsal lobe at right- hand side, in austriaca at left-hand side. Vesica with complex set of cornuti: a group of about 10-20 long needle-like cornuti at right-hand side (dorsally in lateral view), many smaller cornuti of variable shape spread throughout vesica and usually three larger cornuti near phallotrema: a long smooth spine, often protruding from phallotrema, basally connected to a smaller cone-shaped cornutus; in addition another large, often irregular cornutus is often hidden amongst the group of spine-like cor- nuti, it may also protrude from the aedeagal tube. Sometimes both large cornuti are very similar, and other cornuti may be more reduced in size and number. In coronillae the dorsal lobe has a spiny process, which in normal slides is inseparable from the cornuti (fig. 49). Female genitalia (figs. 89-100). — In general as in (sub)genus. Terminal segments either broad and blunt, or slightly pointed. Ductus bursae relatively wide; corpus long, covered with minute pectina- tions; signa usually long and narrow: a thickened margin encloses some rows of complete cells, out- 209 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 17-22. Trifurcula spp., male, details of wings. - 17, T. subnitidella, ultrastructure of normal wingscale from fig. 15; 18, T. coronillae, forewing underside with yellow patch, different areas numbered from wingbase; 19, idem, ultrastructure of scale from area 2; 20, T. josefklimeschi, forewing underside with costal fold and androconial scale patch underneath; 21, idem, hindwing with humeral lobe and ‘black’ scales, costal margin below; 22, idem, detail. bs = black scales; f = fold; hl = humeral lobe; hs = hairscales. side this margin there is a row of incomplete larger cells. Ductus spermathecae with less than 5 coils. Final instar larva (figs. 101-104). — Body ex- tremely elongate, deep yellow in colour. Head with frontoclypeus almost rectangular, tentorial arms short. Labrum with two pairs of setae: a medial and a lateral. Labial palpi three segmented. Antenna with sensilla not cross-wise. Prothorax with small ventral sclerite and a pair of narrow dorsal sclerites. Thorax with respectively 13, 12 and 10 setae pairs, 210 abdominal segments 1-8 with 6, segment 9 with 3 and 10 with 2 setae pairs in all species examined. Integument with reduced spinosity, usually only spines ventrally and laterally upwards to the spir- acles; spines usually short and inconspicuous. Anal rods posteriorly bifid. The four species described below can hardly be distinguished as larva, except by size, length/width ratio of the headcapsule and slight differences in spinosity. Diagnosis Males of the subnitidella group are recognized by the presence of a yellow androconial patch on the forewing underside, in some species in combination with black scaling along the hindwing costa. In 7. josefklimeschi the patch is hidden under a large costal fold, in s#/vrae the patch is absent, but there are rows of dark brown to black scales along the margin of the forewing costal fold and along the hindwing costa. Males of several species in the subgenus Glaucolepis Braun (such as T. stoecha- della (Klimesch) and T. saturejae (Parenti)) also have patches of androconial scales on the underside of the forewing: they are either farther away from the wing base or extend over almost the whole wing surface; these species also differ in venation and genitalia (see van Nieukerken & Johansson, 1990). Females cannot be differentiated externally from other Trifurcula species. The female genitalia differ from other Trifurcula s. 1. species by a combination of the typical shape of signa as described above and the relative short ductus spermathecae with few coils. However, some species of the Trifurcula pal- lidella group have similar female genitalia. Biology The biology of this group has not been described prior to the recent discovery of the biology of T. subnitidella (van Nieukerken & Johansson 1990). The hostplant and larva of four of the species des- cribed here are known through rearing. The larvae of these species make stem-mines in herbaceous or shrubby species of Fabaceae, belonging to the genera Lotus, Dorycnium, Anthyllis or Coronilla, see table 1. The stem-mining species of this group are often sympatric with and feeding within cen- timeters of leaf-mining species of the subgenus Trifurcula (Levarchama), which have a very similar host-range: on Lotus we find T. subnitidella to- gether with either 7. (L.) eurema (Tutt) or T. (L.) cryptella (Stainton); on Dorycnium, T. josefklime- schi occurs together with T. (L.) eurema and on Antbyllis cytisoides we found T. victoris on the same plants as T. (L.) anthyllidella Klimesch. Hi- therto all species of the subnitidella group are found on a single hostplant genus only. The egg is deposited on the stem of the host, in one species on a leaflet. The larvae make relatively straight gallery mines in the green bark of the stem, usually changing direction of feeding several times (figs. 105-107). Larvae feed with their venter to- wards the epidermis. The full-grown larva quits the mine through a semicircular slit, and spins a cocoon on the soil or in leaf-litter. The life history is still incompletely known. It is assumed here that most species are univoltine, al- beit with a long flight period, but bivoltinism is not excluded, and well possible for T. sxbnitidella and josefklimeschi. The mediterranean species have VAN NIEUKERKEN: Trifurcula subnitidella group Table 1. Hostplants of Trifurcula subnitidella group, all in family Fabaceae (Papilionaceae). Hostplant species of Trifurcula Tribus Loteae Dorycnium hirsutum (L.) Ser. D. pentaphyllum Scop. Lotus corniculatus L. Anthyllis cytisoides L. 7. T. josefklimeschi 7. T. josefklimeschi 6. T. subnitidella 5. T. victoris Tribus Coronilleae Coronilla juncea L. 4. coronillae been collected as larvae in January, February and April, and T. subnitidella larvae have been found in September and October in The Netherlands. Most specimens of subnitidella and josefklimeschi emerged within a short period, but T. coronillae and victoris specimens emerged over a period of several months. Although the hostplant and immature stages of the other five species are unknown, it is tentatively assumed that they feed on related hostplants (her- baceous Fabaceae) and make stem-mines as well. The first assumption is supported by the localities of several of these species: they have been found in grassland habitats without any species of broom, the most likely alternative hosts in this subgenus. T. iberica and silviae have been found in alpine meadows, where herbaceous Fabaceae are abundant and T. puplesisi occurs in western Asia, beyond the natural range of broom species. Adults of the subritidella group have been col- lected by sweeping in grasslands and at light. They are, however, easily overlooked, and the search for mines is a more secure way to obtain records. Distribution The group has a wide distribution in the western Palaearctic region, but only T. subnitidella is wides- pread. The other species center around the mediter- ranean, currently with the highest number of spe- cies in Spain (5), southern France (3) and eastern Austria (3). One species, T. puplesisi, has a much more eastern distribution: around the Caspian Sea and in the western part of the Central Asian deserts, it is the easternmost Palaearctic species of Trifur- cula s. str. The scarcity of records from Greece and northern Africa presumably is biassed by collecting activities. Checklist of the Trifurcula subnitidella group Trifurcula Zeller, 1848 subgenus Trifurcula s. str. 211 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 submitidella species group - Underside of forewing with large costal fold, l. austriaca Sp. n. hiding the distinct androconial patch (figs. 20, 2. luteola sp. n. 58). Hindwing costa with enlarged lobe, a small 3. puplesisi sp. n. group of black scales just behind frenulum (fig. 4. coronillae sp. n. DO io SEEN à 7. T. josefklimeschi 5. victoris sp. n. - Underside of forewing without yellow patch, 6. subnitidella (Duponchel, 1843) but margin of costal fold with row of dark- griseella Wolff, 1957 brown or black special scales; a row of similar 7. josefklimeschi sp. n. scales along hindwing costa (fig. 60). Pale al- 8. iberica sp. n. PINEISPECICS APE 9. T. silviae 9. silviae sp. n. 2. Yellow patch divided into three well-separated areas with different tinges: from yolk yellow, through yellowish white to grey-brown (figs. Key to the males of the Trifurcula subnitidella 10, 18). Hindwing costa without black scaling ERO UP Ee OTT si A O EN ei Ge ANNE ei 4. T. coronillae 1. Underside of forewing with a well exposed - Yellow patch with only one colour. Hindwing patch of yellow(ish) androconial scales near costa with or without black scaling........... 3 wing base (figs. 8-10). Hindwing costa with or 3. Hindwing costa with black scales extending Wichoutsomeiblacksis cal es rn ee 2 from about % to % (fig. 59)....... 8. T. iberica Figs. 23-28. Trifurcula austriaca, male genitalia holotype, slide 2591 (except fig. 28). - 23, Capsule, valva outlined; 24, Juxta; 25, Valva, dorsal aspect; 26, Larger cornuti from paratype, slide Klimesch 757, slightly squashed; 27, Aedeagus, ventral aspect; 28, Idem, lateral aspect. Scale 0.1 mm. 2112 © coronillae © iberica B luteola A silviae EA W austriaca . Small species (forewing length 2.6-2.9 mm) Hindwing either with very small group of black scales near frenulum or without such scales 4 . Hindwing costa with a small row of black scales just behind frenulum (fig. 57) (difficult to de- tect). Antenna with 31-36 segments. Gnathos asymmetrical, with a small keel at right side (COSO eno 6. T. subnitidella Hindwing costa completely without black scal- ing. Antenna with 34-48 segments. Gnathos either asymmetrical with two keels, or symmet- GIGAN ate ore Peng aa o Ne ea 5 with 34-37 antennal segments. Androconial patch (fig. 8) yellowish white, similar scales also present on hindwing upperside. Aedeagus with two curved cornuti and one spine-like cor- OUEUSK(H ONDO) par ne 1. T. austriaca Larger species (2.8-3.5 mm), with 40-48 anten- nal segments. Androconial patch a darker yel- low. Aedeagus without curved cornuti, but with one or two spine-like and a conical cornutus . Gnathos asymmetrical, with two large keels (fig. 51, 52). Aedeagus with two very similar spine-likercornuti nen 5. T. victoris Gnathos simple, symmetrical (figs. 30, 37). Ae- deagus with three differently shaped larger cor- MUD en Maus art a Ser Een le 7 . Very pale species: scale tips yellow (fig. 2). Valva almost triangular, gradually narrowing towards apex (fig. 33). Species from western Krane tee. tente coco Darker species: scales with darker tips. Valva basally with almost parallel margins, distally suddenly tapering towards apex (fig. 38). Spe- GIES (HHONIN WIESE JAG i 3. T. puplesisi VAN NIEUKERKEN: Trifurcula subnitidella group Fig. 29. Distribution of Trifwrcula spe- cies, see legend. Mapped on 50 X 50 km UTM squares. 1. Trifurcula austriaca sp. n. (figs. 1, 8, 23-28, 29) Trifurcula n. sp.; van Nieukerken im Kasy 1985: 5. Type material. - Holotype 8, Austria: Fürbachwiesen, E. of Gramatneusiedl (Niederöst.), UTM: 33U XP12, 15.v.1968, F. Kasy, Genitalia slide EvN 2591 (NHMW). - Paratypes, 2 &: same data, but 4.ix.1973, 31.vii.1978 (RMNH, NHMW). Description Male (fig. 1). - Forewing length 2.6-2.9 mm, wingspan 5.6-6.7 mm. Head: frontal tuft yellowish orange, collar yellowish white. Antenna with 34-37 segments; scape yellowish white, sometimes with a few dark scales. Forewing and thorax covered with fuscous tipped scales, mixed with white scales, giv- ing a grey-brown impression; terminal cilia white beyond relatively distinct cilia-line. Underside of forewing with basal patch buff yellow, extending to ‘4 (fig.8). Hindwing with a row of white hair-scales in stead of costal bristles; basally on upperside with some yellowish white to buff yellow scales, similar to forewing underside. Female unknown. Male genitalia (figs. 23-28). — Capsule length 330-350 um. Vinculum anteriorly slightly nar- rowed, truncate; lateral arms near gnathos with minute spines. Uncus with medial process pointed, with pair of lateral setae. Gnathos symmetrical, central element triangular, with wide apical angle; anterior processes present. Valva length 245-255 um, basally broad, with parallel margins, posterior half suddenly narrowed towards short pointed tip, transverse bar of transtilla less than twice the length of sublateral process. Aedeagus 310-385 um PAS) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 30-36. Trifurcula luteola , male genitalia. - 30, Capsule, valva outlined, slide 2342; 31, Gnathos, holotype, slide 2503; 32, juxta, holotype, slide 2303; 33, Valva, dorsal aspect, slide 2342; 34, Aedeagus, ventral aspect, holotype, slide 2303; 35, Aedeagus, ventro-lateral aspect, slide 2304; 36, Larger cornuti, holotype, slide 2303. Scale 0.1 mm. long, with ventral carina posteriorly ending in two ventrally curved processes; dorsal lobe at right side indistinct, a large lobe at left side, ventrally enlarged and strongly sclerotized (fig. 27, 28). Vesica with three large cornuti (fig. 26): one long pointed (-135 um) and two curved (respectively 85 and 60 um); further a serrate cornutus lower in vesica, in addi- tion to the group of needle-like and the small cor- nuti. Juxta fig. 24. 214 Diagnosis Externally most similar to sabnitidella, but with- out black scales near frenulum; austriaca also has special scales on hindwing upperside. Other species without black scales are larger and/ or paler (luteola, puplesisi, victoris). Male genitalia resem- ble more those of the T. pallidella group than other species in this group: the two curved cornuti and the triangular gnathos are diagnostic. Biology Immature stages and hostplant unknown. The adults were all taken at light, from May to Sep- tember, in a grassland nature reserve, with a very rich Lepidoptera fauna (Kasy 1985). Distribution (fig. 29) Eastern Austria: only from the type locality: the ‘Pischeldorfer Fischawiesen’ (named Fürbach- wiesen on labels). After finishing this description, Z. Lastúvka showed me in August 1990 a drawing of male genitalia undoubtedly belonging to this species, from a specimen from Czechoslovakia: Prostéjov Hamry [UTM: 33U XQ48], collected in 1990; this record is added on fig. 29. Possibly a species with an eastern distribution, like several other Lepidoptera species occurring in this area. Etymology An latinized adjective, from the type locality Aus- tria. 2. Trifurcula luteola sp. n. (figs. 2, 29, 30-36) [Trifurcula pallidella Duponchel [1963]: 1209. Misidentification.] partim; Lhomme Type material. - Holotype &: France, Molières[-sur- l’Alberte] (Aude), UTM: 31T DH57, 6.viii.1903, Chré- tien, Genitalia slide EvN 2303 (MNHN). - Paratypes: 4 6. France: 1 ¢ (abdomen missing), data as holotype (MNHN); 1 6, Arten. [St. Pons, montagne d’Artenac] (Hérault), UTM: 31T DJ81, 19.vii.1904, Chrétien (MNHN); 2 3, Nesp. [Nespouls, near St. Pons] (Hé- rault), UTM: 31T DJ81, 2.viii.1904, (MNHN, RMNH). Description Male (fig. 2). - Forewing length (2.4) 3.2-3.4 mm, wingspan (5.4) 7.2-7.6 mm. Head: frontal tuft yellowish orange, collar paler. Antenna with 40-45 segments; scape white with few yellow scales. Fore- wing and thorax pale yellowish white, tips of scales yellow; terminal cilia concolorous. Underside of forewing with orange yellow basal patch to %, cos- tal fold with row of brown scales along edge. Hindwing white, humeral lobe on upperside near frenulum with a row of yellowish-orange special scales. Female unknown. Male genitalia (figs. 30-36). — Capsule length 370-375 um. Vinculum anteriorly truncate, broad and short. Uncus with medial process terminally truncate, not widened, several lateral setae. Gna- thos symmetrical, central element approximately triangular, slightly angular or more rounded (figs. 30, 31); long anterior apodemes present, but not very distinct. Valva length 280-305 um, roughly triangular, with short pointed tip; transverse bar of transtilla 2.5-3 X as long as sublateral process. Ae- VAN NIEUKERKEN: Trifurcula subnitidella group deagus 405-435 um long, with long, terminally fringed, ventral carina; vesica with one long, pointed cornutus (225-250 um), a short conical (75 um), + 10 long needle-like and many small cornuti. Aedeagal tube with dorsal lobe at right side not very pronounced, slightly serrate. Juxta fig. 32. Diagnosis Externally easy to separate from most other spe- cies by pale colour and size. Resembles Trifurcula pallidella (Duponchel), but male easily distin- guished by yellow patch on forewing underside. Male genitalia characterized by triangular valva and triangular, symmetrical gnathos, but see puplesisi. Biology Immature stages and hostplant unknown. Adults collected in July and August. Distribution (fig. 29) Only known from Southwest France, in mediter- ranean region. Remarks The four specimens of this species were found amongst material labelled as Trifurcula pallidella in the Chrétien collection. Although the species has only been collected in the beginning of this century, it is very well possible that it still can be found, but has until now been overlooked, as was the case with silviae. Etymology A Latin adjective, from /uteolus = yellow, orange, named after the pale forewing colour. 3. Trifurcula puplesisi sp. n. (figs. 37-44, 89, 95) Type material. - Holotype 8: Soviet Union: Turkmen- istan, Sandykachi (Sandy Katschi), UTM: 41S MA55, 1.v.1986, Puplesis, Genitalia slide EVN 2760 (ZKVV). — Paratypes: 2 6, 4 Q. Soviet Union: 1 @, 2 Q, same data as holotype; 2 9, Turkmenia, Central Karakum, env. Ash- khabad, UTM: 40S FH20, 4.vi.1988, R. Puplesis (ZK VV, RMNH); 1 4, [Derbent (Dagestan), UTM: 39T TG76], 10.vii.[1872], [ Nacht bei Ligt’], Christoph, ‘473’, Zeller Coll., Walsingham collection (BMNH). Excluded from type-series (probably conspecific): 1 8, Krasnoarmeysk (Sarepta), UTM: 38U MX77, Christoph, ‘397’ [red label] (BMNH). Description Male. - Forewing length 2.9-3.7 mm, wingspan 6.2-8.0 mm. Head: frontal tuft pale wish orange to almost white, collar white. Antenna with 46-48 segments; scape white. Forewing and thorax greyish brown, scales with dark tips, except along dorsum; terminal cilia white. Underside of fore- wing with a small pale yellow basal patch to \/s. 215 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Hindwing white, humeral lobe on upperside near frenulum with some yellow special scales, similar to those on forewing underside; costa basally with row of white hair-scales. Female. - Forewing length 3.2-3.3 mm (3.25 + 0.04, 4), wingspan ca 7.3 mm. Antenna with 43-45 segments. Male genitalia (figs. 37-43). — Capsule length 350-410 um. Vinculum wide and short, anteriorly broadly rounded. Uncus with medial process termi- nally truncate or bifurcate (figs. 37, 39). Gnathos symmetrical, central element broadly rounded, an- terior apodemes not evident. Valva length 275-295 um, broad, margins in basal half approximately parallel sided, tip slightly curved, pointed; trans- verse bar of transtilla approximately twice as long as sublateral process. Aedeagus 350-435 um long, with long, terminally fringed, ventral carina and a dorsal lobe at right side, with serrate margin. Vesica with one long, curved, pointed cornutus (155-190 pm), a short conical (65-100 um), a long irregular cornutus, seeming composed of several cornuti and almost hidden amongst the long needle-like cor- nuti; many small cornuti present. Juxta fig. 40. Figs. 37-43. Trifurcula puplesisi, male genitalia. - 37, Capsule, valva outlined, slide 2761; 38, Valva, dorsal aspect, slide 2761; 39, Slightly different uncus of holotype, slide 2760; 40, Juxta, slide 2761; 41, Cornuti, dorsal aspect, slightly squashed, slide BM 25634; 42, Aedeagus, ventral aspect, part of cornuti, protruding from aedeagal tube also shown, slide 2761; 43, Aedeagus, lateral aspect, holotype, slide 2760. Scale 0.1 mm. 216 Female genitalia (figs. 89, 95). - Terminal seg- ments relatively broad; T8 with about 9-11 setae and several spines, broad anal papillae with 28-37 setae. Signa long, 525-630 um. Ductus spermathe- cae with 34 coils. Diagnosis Males resemble other species without black scal- ing and simple yellow patch, in particular austriaca, which is smaller and has fewer antennal segments and /wteola, which is usually much paler and has a more triangular valva. Female recognized by large number of setae on anal papillae and relatively long signa. Biology Hostplant and immature stages unknown. The adults were taken in steppe area, almost desert, in May, June and July. Distribution (fig. 44) Caspian Sea area, just inside Europe and southern Turkmenia. Remarks The specimen from the Christoph collection, la- belled ‘Sarepta’ is tentatively assigned to this spe- cies, but excluded from the type series, since it is much paler and shows slight differences in genita- lia, such as the form of the juxta. The paratype from the Zeller/Walsingham col- lection, also collected by Christoph, has only been labelled with: ‘10/7’ [black ink on green, handwrit- ing] / ‘473 [round label, black ink on green, handw- riting] / ‘Christoph’ [black ink on white, handwrit- ing], and the usual Walsingham collection labels with no. 101450. The locality has been traced from correspondence from Christoph to Zeller, now present in the Entomology Library, BMNH. Only in one letter with lists of specimens, sent by Chris- VAN NIEUKERKEN: Trifurcula subnitidella group Fig. 44. Distribution of Trifurcula puplesisi, mapped on 50 X 50 km UTM squares. toph to Zeller for identification, a number as high as ‘473’ could be found. It is assumed that this refers to the specimen in question. The letter is dated ‘Sarepta Den 12 Marz 1873’ and bears the number 382. Under the number ‘473’ in the list ‘Determi- nanda’ is written ‘Derbent, Nacht b Ligt’. It is also assumed that the specimen was collected in July 1872, since the letter apparently refers to the most recent sending, collected in the previous year. Note further that the locality Derbent, a town on the Caspian Sea, is sometimes confused in lepidopteran literature with the locality ‘Derbend’ in Iran, North of Teheran, where recently various lepidopterists collected. Christoph often travelled via Derbent, when he crossed the Caspian Sea towards Turkmen- istan or northern Iran (Christoph 1877), and as far as I know never collected in the Iranese ‘Derbend’. Etymology A noun in genitive, named in honour of its col- lector, Dr. Rimantas Puplesis, a Lithuanian special- ist of central and east Asian Nepticulidae, who kindly trusted me his material for description. 4. Trifurcula coronillae sp. n. (figs. 3, 10, 18, 19, 29, 45-50, 90, 96, 101, 106) Type material. - Holotype @: Spain, Sierra Blanca, 6 km N Marbella, El Mirador (Malaga), 800 m, UTM: 30S UF34, 5-9.11.1984, Coronilla juncea, from stem-mines, el. 13.vii.1984, EvN no. 84046, E. J. van Nieukerken, Geni- talia slide EVN 2592 (RMNH). - Paratypes: 15 6, 11 9, 30 larvae. Spain: 4 8, 4 9, 7 larvae, data as holotype, e.l. 31.v.1984- 25.viii.1984, (RMNH); 13 larvae, 5 km S Istán, road to Embalse de la Concepción (Málaga), 150 m, UTM: 30S UF2545, 17.1.1988, Coronilla juncea, stem-mines, EvN no. 88057, van Nieukerken & Richter (RMNH); 4 8, El Saler (Valencia), UTM: 30S YJ26, 22.iv.1981, C. Gielis (RMNH, coll. Gielis); 2 larvae, Rodalquilar, 5 km SW Las Negras (Sierra del Cabo de Gata) (Almería), 120 m, UTM: 30S WF8579, 8.1.1988, Coronilla juncea, stem- 2417, TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 45-50. Trifurcula coronillae, male genitalia holotype, slide 2592 (except 48, 49). - 45, Capsule, valva outlined; 46, Valva, dorsal aspect; 47, Juxta; 48, Large cornuti, separately drawn, slide 2593; 49, Aedeagus, ventral aspect, slide 1883; 50, Aedeagus, lateral aspect. Scale 0.1 mm. mines, EvN no. 88005, van Nieukerken & Richter (RMNH); 2 6, 4 9, Sierra Blanca, El Garopala, 3 km NNW Ojén (Malaga), 670 m, UTM: 30S UF3250, 15.1.1988, Coronilla juncea, stem-mines, e.l. 6.vi.1988- 22.v111.1988, EvN no. 88051, van Nieukerken & Richter (RMNH); 4 8, 3 9, 8 larvae, Sierra Blanca, Puerto de Marbella (El Mirador), 3 km W Ojén (Málaga), 900 m, UTM: 30S UF3048, Coronilla juncea, 15.1.1988, el. 30.v.1988-30.vii.1988, EvN no. 88047, van Nieukerken & Richter (RMNH); 1 8, road to Istan (Málaga), 400 m, UTM: 30S UF24, 25.vi.1975, E. Traugott-Olsen (ZMUC). Excluded from type series: 2 & (abdomina lost), data as holotype, el. 1.viii.1984 (RMNH). Description Male (fig. 3). - Forewing length 2.6-3.2 mm (2.88 + 0.17, 18), wingspan 5.6-6.7 mm. Head: frontal tuft yellowish orange to orange , collar slightly paler to almost white. Antenna with 41-48 segments (44.0 + 2.0, 12); scape white, often with scattered 218 brown scales. Forewing and thorax covered with brown tipped scales, distal 4 dark, edge darker, giving the wing a greyish brown impression; white scales sometimes exposed at dorsum, terminal cilia yellowish white, cilia line indistinct. Underside of forewing with large patch, extending from base to %, more or less distinctly divided into three areas with different colour: basally deep yellow, middle part buff yellow, terminally olive brown (figs. 10, 18, 19). Hindwing greyish, basal area with white lamellar scales. Female. - Forewing length 2.4-3.2 mm (2.91 + 0.23, 11), wingspan 5.4-6.4 mm. Antenna with 35- 42 segments (39.8 + 2.2, 10). Male genitalia (figs. 45-50) - Capsule length 340- 355 um. Vinculum anteriorly truncate. Uncus with medial process truncate. Gnathos asymmetrical, with a longitudinal keel, slightly left of middle on ventral surface, posterior margin rounded. Valva length 250-270 um, with almost parallel margins, suddenly narrowed in terminal quarter towards pointed tip, transverse bar of transtilla twice as long as sublateral processes or longer. Aedeagus 360-390 um long, with ventral carina bifurcate; aedeagal tube with spatulate tip ventrally, dorsal lobe not pronounced at right side, but with strong medial spine-like process, curved towards right side, which in normal preparations is almost inseparable from the large cornuti: there appear to be three in stead of two. Vesica with two large spine-like cornuti (one somewhat irregular) and a conical cornutus; long needle-like and many smaller cornuti present. Juxta fig. 47. Female genitalia (figs. 90, 96). Terminal seg- ments broad and rounded; T8 with 6-8 setae and several scales; anal papillae with 24-30 setae. Duc- tus spermathecae with 3% to 3/2 coils. Signa 420- 470 um (ventral) and 460-520 um (dorsal) long. Final instar larva (fig. 101). Yellow, elongate. Headcapsule 325-355 um long (341 + 10.7, 5), 1.06-1.13 (1.09 + 0.02, 5) X as long as wide. Spin- osity: prothorax ventrally with narrow band poste- riorly, meso- and metathorax ventrally with a band along anterior margin and very few spines on calli. Abdominal segments 1-9 with sparse spinosity ven- trally and laterally up to the spiracles, spines near anterior margins smaller; segment 10 with small group of spines anterior of setae. Diagnosis Male easily recognized by the androconial patch with three different colours and absence of black scaling on hindwing. Male genitalia unique by shape of gnathos and cornutus-like process on ae- deagus. Female very similar to josefklimeschi, but separated by the blunt abdominal point, versus the slightly pointed condition in gosefklimeschi. Fe- males of victoris are paler, and have less antennal segments, females of subritidella are darker and have also less antennal segments. Biology Hostplant. - Coronilla juncea L., a shrub up to 1 m, with rush-like stems, slightly resembling a broom. Life history. - The egg is deposited on an inter- nodium of the smooth rush-like stem of the host- plant, and is conspicuous. The larva makes a very conspicuous gallery mine in the green bark, in which the living larva can easily been seen (fig. 106). The mine often starts contorting, frequently encircling the stem; the larva often feeds downward first, but later changes feeding direction one or more times, the mine usually ends in upwards di- rection. The mine is filled with greenish brown frass. There are often numerous mines in a single stem. The brown cocoon is made in soil or on leaf VAN NIEUKERKEN: Trifurcula subnitidella group litter. Larvae have been collected in January and February, but many mines were already vacated in that period, so that larvae probably start mining in autumn. Adults emerged over a very long period, from May to August. The only adults taken in the field were collected in April and June. The species is most likely univoltine. Distribution (fig. 29) Along mediterranean coast of Spain, to be ex- pected elsewhere with its host, which occurs from West Yugoslavia to Portugal (Heywood & Ball 1968). Etymology A noun in genitive case, from Coronilla, the ge- neric name of the hostplant. 5. Trifurcula (Trifurcula) victoris sp. n. (figs. 4, 51-56, 76, 91, 97, 102) Type material. - Holotype @: Spain: El Pozo de los Frailes, 3 km N San José (Almeria), UTM: 30S WF7972, 8.1.1988, Anthyllis cytisoides, stem-mines, e.l. 4.iii.1989, EvN no. 88003, van Nieukerken & Richter (RMNH). — Paratypes: 2 6,6 9, 4 larvae. Spain: 2 8,6 9, 3 larvae, same data as holotype, el. 29.ii.1989-30.v.1989 (RMNH); 1 larva, 3 km NE Nijar, Cerro de Movillas (Almeria), 400 m, UTM: 30S WF7393, 9.1.1988, Anthyllis cytisoides, stem-mines, EvN no. 88007, van Nieukerken & Richter (RMNH). Description Male (fig. 4). - Forewing length 3.0-3.5 mm, wingspan 6.5-7.2 mm. Head: frontal tuft yellowish- orange to ferruginous, collar paler, yellowish white. Antenna with 40-42 segments; scape white, with some darker scales, flagellum brown. Forewing and thorax relatively pale, appearing pale brown-grey: scales with distalmost tip (margin only) brown, few white scales at tornus; terminal cilia white beyond more or less distinct cilia-line. Underside of fore- wing with deep yellow patch of about ‘4 forewing length. Hindwing grey, humeral lobe well deve- loped, on upperside with yellow scales along mar- gin, black scaling absent. Female. - Forewing length 2.9-3.3 mm (3.04 + 0.12, 6), wingspan 6.5-7 mm. Antenna with 32-35 segments (33.5 + 1.3, 4). Ovipositor with short point. Male genitalia (figs. 51-56) — Capsule length 425- 440 um. Vinculum anteriorly truncate or rounded. Uncus with medial process strongly widened, trun- cate. Gnathos very asymmetrical, variable, always with two longitudinal keels of different length, leaving ventrally excavation in which aedeagal tip fits (dashed in fig. 51), anterior apodemes present. Valva length 285-290 um, approximately triangu- lar, dorsally with a row of 4-5 spines along conti- nuation of lateral arm of transtilla, transverse bar 219 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 51-56. Trifurcula victoris, male genitalia. - 51, Capsule, valva and tip of aedeagus outlined, slide 3064; 52, Gnathos of holotype, showing variation, slide 2743; 53, Valva, dorsal aspect, slide 3064; 54, Large cornuti, separately drawn, slide 3064; 55, Aedeagus and juxta, ventral aspect, slide 3064; 56, Aedeagus and juxta, lateral aspect, holotype, slide 2743. Scale 0.1 mm. of transtilla about twice as long as sublateral pro- cesses. Aedeagus 395-435 um long, with ventral carina spatulate, dorsal lobe at right side conspicu- ous, margin serrate; vesica with two large and very similar cornuti, one joined to smaller cone-shaped cornutus; group of spine-like cornuti very much reduced in size and number, only a few present at basis of both large cornuti; numerous small cornuti present. Juxta fig. 55. Female genitalia (figs. 91, 97). - Terminal seg- ments forming slightly pointed abdominal tip; T8 220 with 8-10 setae and some scales; anal papillae with 21-29 setae. Ductus spermathecae with 4 coils. Signa 465-530 um long. Final instar larva (fig. 102). - Yellow, elongate. Head capsule 385-435 um, 1.03-1.05 X as long as wide. Spinosity: prothorax ventrally with narrow band of minute spines posteriorly, meso- and me- tathorax ventrally without spines. Abdominal seg- ments 2-9 with microspines ventrally, hardly vis- ible, segment 10 with small group of spines anterior of setae. Diagnosis Male recognized by uniformly coloured yellow patch, absence of black scaling and relatively pale forewings. T. /uteola is paler, with yellow scale tips instead of brown or grey. Male genitalia character- ized by asymmetrical gnathos with two keels and reduction of spine-like cornuti. Female may be con- fused with coronillae, which is darker; victoris also has more pointed anal papillae. Biology Hostplant. - Anthyllis cytisoides L., a broom-like leafy shrub of 0.5-1.5 m, with velvety hairy stems and leaves. Life-history. - Egg deposited on stem. The larva makes a long gallery in the bark, which is difficult to see, because of the dense indumentum of the stem. It could best be seen because of the swelling and the slight discoloration, especially in the case of old mines. Larva not visible in its mine. The mines were actually collected on stems, heavily infested by gall-forming coccids. Larvae collected in January, adults emerged from February to May. One female emerged from leaf-litter, which had been collected from underneath the hostplants. Distribution (fig. 76) Only known from the extremely dry region in the southeast part of the province of Almeria, Spain. Despite search in other sites along the Spanish southcoast, where the host is common, no more mines could be found. Etymology A noun in genitive case, from latin victor, named after my son Victor Alexander. 6. Trifurcula (Trifurcula) subnitidella (Duponchel) (figs. 5, 9, 11-17, 57, 61-68, 92, 98, 103, 107) Elachista subnitidella Duponchel, [1843]: 326, pl. 77: 8. Lectotype @ [designated by van Nieukerken & Johans- son 1987: 471]: [Austria, Vienna region], Duponchel coll., Genitalia slide EvN 2522 (MNHN) [examined]. Trifurcula griseella Wolff, 1957: 21. Holotype 4: Den- mark, Asserbo, 30.v.1954, N. Wolff, genitalia slide NW 1797 (ZMUC) [Synonymized by van Nieukerken & Johansson 1987: 471] [examined]. Nepticula subnitidella (Duponchel) Zeller 1848: 305 [redescription, mistaken identity], Joannis 1915: 127 [identity], Rebel 1901: 221 [catalogue], Meess 1910: 474 [listed]. Trifurcula subnitidella (Duponchel) van Nieukerken & Johansson 1987: 471 [selection lectotype], 1990: 273 [description, NW Europe]. Trifurcula griseella Wolff, Bradley 1962: 174, fig. 4 Ireland, male genitalia], Bjorn & Pallesen 1971: 111 [Denmark], Bradley et al. 1972: 3 [Britain], Svensson 1974: 171 [Sweden], Emmet 1975: 39-42 [England, Scotland], Karsholt & Nielsen 1976: 18 [Denmark], Emmet 1976: 209 | description, British Isles], Svensson VAN NIEUKERKEN: Trifurcula subnitidella group 1980: 85 [Sweden], Buhl et al. 1983: 120 [Denmark], Svensson 1983: 65 [Sweden], Kasy 1983: 5 [ Austria], Buhl et al. 1984: 3 [Denmark], Svensson 1985: 81 [Sweden], Kasy 1985: 5 [ Austria], Karsholt 1985: 45 [catalogue Denmark], van Nieukerken 1986a: 16 [checklist], Bradley & Fletcher 1986: 2 [list], Prose 1987: 49 [West Germany: Bavaria], Svensson 1987: 3- 3 [catalogue Sweden], Buhlet al. 1987: 100 [Denmark], Buhl et al., 1988: 101 [Denmark]. Description Male (fig. 5). - Forewing length 2.0-2.6 mm (2.37 + 0.17, 44), wingspan 4.2-5.8 mm. Head: frontal tuft yellow-ochre to fuscous, variable; collar similar. Antenna with 31-36 segments (33.4 + 1.3, 40); scape white, sometimes with few brown scales. Forewing dark greyish-brown to fuscous, slightly irrorated by dark tipped scales, no white spots pres- ent; underside with distinct basal patch of deep yellow scales-near costa reaching to % (figs. 9, 11- 13). Hindwing: grey, underside along costa directly behind frenulum with a distinct short row of black lamellar scales (figs. 14, 15, 57). Abdomen dark grey, with yellowish grey tufts. Female. - Forewing length 2.0-2.4 mm (2.22 + 0.16, 9), wingspan 4.4-5.4 mm. Antenna with 27-32 segments (30.4 + 1.7, 9). Male genitalia (figs. 61-67). — Capsule length 330-390 um (349.6 + 14.8, 15). Vinculum ante- riorly rounded. Uncus slightly widened, truncate at tip. Gnathos asymmetrical, central element with an anteriorly curved keel, ending in a pointed process at right side; lateral arms with long and narrow anterior apodemes. Valva length 250-289 um (264.2 + 9.0, 15), basally with almost parallel mar- gins, narrowed in middle towards pointed tip. Ae- deagus 330-365 um (344.0 + 14.0, 15), with ventral carina bifid; aedeagal tube with spatulate tip, dorsal lobe in middle, or slightly on right side, with serrate margin. Vesica with one very long straight, or slightly curved cornutus, with a conical cornutus joined to its basis, and a pointed cornutus with serrations; less then 10 spine-like cornuti and many small cornuti present. Juxta fig. 62. Female genitalia (figs. 92, 98). - Terminal seg- ments rather broad; T8 with 3-6 setae and some scales; anal papillae with 11-20 setae each. Ductus spermathecae with 314 convolutions. Signa 300-355 um (dorsal) and 350-365 um (ventral) long. Final instar larva (fig. 103). - Long and slender, deep yellow. Head capsule 325-345 um, 1.13-1.16 X as long as wide, distinctly narrower than in other three described species. Spinosity: prothorax ven- trally with narrow band posteriorly, meso- and me- tathorax with spines between D1 seta and calli. Abdominal segments 1-9 with sparse spinosity ven- trally and laterally up to the spiracles, spines near anterior margins smaller; segment 10 with small group of spines anterior of setae. (NO) 159] A TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 57-60. Trifurcula males, forewing and hindwing undersides, showing position of black scales, yellow patch and costal fold. - 57, T. subnitidella, Netherlands, Katwijk; 58, T. josefklimeschi, paratype, Austria, Hundsheimer Berg; 59, T. iberica, hindwing only, paratype, Spain, Prullans; 60, T. silviae, paratype, France, Ceillac. Scale 0.3 mm. Diagnosis Male subnitidella differs from other species in the group by the combination of a simple yellow patch and a group of black scales just behind fren- ulum; further it is one of the smaller and darker species, with less antennal segments than most other species (36 maximum). The male genitalia are well characterized by the gnathos process. Fe- males are smaller and darker than the other des- cribed females, and have less antennal segments. Biology Host plant. - Lotus corniculatus L., a common perennial herb of grasslands all over Europe, pos- sibly also on other Lotzs species. Life history. - The egg is conspicuous and dep- osited on the stem of the host, usually at the base 222 of a leaf-stalk (fig. 107), frequently near the stem base. The mine is a long gallery in the bark of the stem (fig. 107): the larva is first mining down for 0.5-2 cm, then going up the stem ina rather straight line, or partly encircling the stem, occasionally go- ing down again in the last part of the mine. The mine may reach a total length of 6 to 9 cm. The mine is at first narrow, reddish brown, with straight edges, but later becomes as wide as the stem, with irregular margins, becoming silvery white in fresh mines. Frass deposited in midline, brown, not al- ways well visible. Larva well visible in mine, ap- pearing as a slight swelling. The stems of the host are not killed by the mining activity. The larva quits the mine through a semicircular slit, and spins a cocoon in the soil or on leaf-liter. VAN NIEUKERKEN: Trifurcula subnitidella group 62 65 Figs. 61-67, Trifurcula subnitidella, male genitalia, slide 2501 (slide 2745 for fig. 62). - 61, Capsule, valva outlined; 62, Juxta; 63, Valva, dorsal aspect; 64, Aedeagus, ventral aspect; 65, Idem, dorsal lobe, dorsal aspect; 66, Large cornuti, separately drawn; 67, Aedeagus, lateral aspect. Scale 0.1 mm. Larvae have been taken in September and Oc- tober, adults fly from May (March in Tunisia) to early September. Reared adults emerged in May. Voltinism not yet clear from these data, clear peaks in flight are not apparent. More data from single localities are needed, and bivoltinism can only be proved by finding larvae and rearing in early summer. In northern Europe T. submitidella is almost ex- clusively found on limestone grasslands (down- land) and coastal dunes, in southern Europe it is found in many habitat types. The species might be vulnerable to habitat loss in more northern parts of its occurrence. Distribution (fig. 68) Widespread in Europe, northward to 60° N, southward to the northern border of the Sahara in Tunisia and eastward to the Crimea and Asia minor. Not yet recorded from Norway, Belgium, Luxem- bourg, Switzerland, Poland, Hungary, Albania and Portugal, nor from any of the large mediterranean islands. The species is here for the first time in detail recorded from Finland, The Netherlands, Germany, Czechoslovakia, France, Spain, Italy, Ru- mania, Bulgaria, Yugoslavia, Greece, Turkey, Soviet Union (Estonia and Ukraine) and Tunisia. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Remarks T. subnitidella is slightly variable in colour, spec- imens from northern Europe tend to be darker than those from southern Europe. The figure of the male genitalia in Wolff's des- cription of T. griseella (1957), has been published as mirror image. In North America, a striking case of convergence has been described by Wagner (1987): his species Microcalyptris lotella was reared from very similar stem-mines in Lotus, and, moreover as adult male it has a yellowish patch on the forewing underside, very similar to that of sabnitidella. Also lotella has been found in a habitat that in Europe could har- bour subritidella: coastal dunes. Yet Microcalyptris Braun (synonymized with Acalyptris Meyrick by van Nieukerken 1986a) is a completely different genus, only distantly related to Trifurcula. Material examined. - 115 8, 14 9, 5 larvae (including type-material cited above). - Austria: 1 8 [genitalia slide MV13111 only], Apetlon, Seewinkel (Burgenland), UTM: 33T XN38, 9.viit.1963, E Kasy (NHMW); 1 5, Fürbach- wiesen, East of Gramatneusiedl (Niederöst.), UTM: 33U XP11, 23.vii.1981, E Kasy (NHMW); 1 6, Hundsheimer Berg S. ( (Niederöst.), UTM: 33U XP43, 8.vil. 1980, F. Kasy (NHMW); 1 4 [photo genitalia slide], Linz, surroundings (Oberöst.), UTM: 33U VP44, 18.vii.1934, J. Klimesch (coll. Klimesch). — Bulgaria: 1 8, Nesebur (Nessebar), UTM: 35T NHS2, 23.viii-4.ix.1962, J. Soffner (DEIC). — Czechoslovakia: 2 &, Detkovice (Moravia), UTM: 33U XQ57, 2.vi and 4.vii.1989, A. LaStuvka (coll. LaStuvka). — Denmark: 1 9, Asserbo, UTM: 33V UC11, 10.vi.1964, N. L. Wolff (ZMUC); 6 8, Laesö, Bovet ( (NE), UTM: 33V PJ25, 8,9 and 13.vii.1982, 30.viand 8.vii.1983, O. Karsholt (ZMUC); 9 6, 5 9, Laesö, Höjsandet (NE]), NINE 33V PJ25, 1 and 6.vii.1983 O. Karsholt (ZMUC); 4 8, Laesö, Nordmarken (NEJ), UTM: 33V PJ25, 5, 6 and 9.vii.1983, O. Karsholt (ZMUC). - Finland: 2 4, Virolahti (EK), UTM: 35V NH30, 27.vi.1989 [netted at seashore over Lotus corniculatus |, J. R. Kaitila (RMNH). - France: 1 4, Cannes (Alp. Mar.), UTM: 32T LP42, iv.1881, Wal- singham (BMNH); 1 8, Col de Soubeyrand (Drôme), 994 m, UTM: 31T FK81, 2.viii.1986, H. W. van der Wolf (coll. van der Wolf); 1 @, La Veuve, Digne, UTM: 32T KP78, 12.viii.1901, Chrétien (MNHN); 1 g, Nesp. [probably Nespouls, near St. Pons] (Hérault), UTM: 31T DJ81, 2.viii.1904, (MNHN). - Germany (West): 2 4, Bamberg: Tütschengereuth (Bayern), UTM: 32U PA22, 4.viii.1978, G. Derra (coll. Derra). - Germany (East): 2 6, Blanken- burg, Muschelkalk UTM: 32U PBS1, 29.v.1965, 10.viii. 1989, H. Steuer (coll. Steuer). - Great Britain: 1 Ó, Mickleham (Surrey), UTM: 30U XB88, 23.vi. 1856, grass, H. T. Stainton (BMNH); 1 6, Mickleham, Headley Lane (Surrey), UTM: 30U XB88, 10.vi.1857, (BMNH). - Greece: 1 g, Kavisos (Evro), 100 m, UTM: 35T ME, 22- 23.viii.1985, A. Moberg (NHRS). - Italy: 2 6, Almese, surroundings (Torino), 350 m, UTM: 32T LQ79, 16 and 20.v.1979, U. Parenti (RMNH, coll. Parenti); 1 8, Asti, Boschi di Valmanera, 124 m, UTM: 32T MQ37, 1.v.1970, G. Baldizzone (GBA); 2 8, Asti, Valmanera (Asti), UTM: 32T MQ37, 28.iv.1968, 26.viii.1968, G. Baldizzone (RMNH, coll. Passerin- d’Entreves); 1 6, Baia Domizia (Caserte), UTM: 33T UF55+, 25.vii.1972, R. Johansson (coll. Johansson): 1 8, Cardona (Alessandria), 300 m, 224 UTM: 32T MQ39, 19.v.1975, G. Baldizzone (GBA); 2 è, Lucotena (Firenze), 500 m, UTM: 32T PP93, 1 l.viii.1982, J. Kuchlein (RMNH, coll. Kuchlein); 1 @, Monti Aurunci, S km N Ieri (Latina), 600 m, UTM: 33T UF77, 24- 30.v. 1969, R. Den (coll. Johansson); 1 8, Poggio di Casasco (Alessandria), 300 m, UTM: 32T NQ06, 3.vi. 1978, G. Baldizzone (GBA), 2 a, Val Susa (Piemonte), Villardora (Torino), 500 m, UTM: 32T LQ79, 26.v.1983, G. Bassi (coll. Bassi). - Netherlands: 1 &, Katwijk, 2 km N: dunes (Zuid-Holland), UTM: 31U ET9686, 16.x.1988, Lotus corniculatus, stem-mines, e.l. 21.v.1989, EvN no. 88175, van Nieukerken & Richter (RMNH); 1 @, Kun- rade, Kunderberg (Limburg), UTM: 31U GS0739, 02.vii.1983, G. R. Langohr (coll. Langohr); 2 @, 2 9, 3 larvae, Kunrade: Kunderberg W. (Limburg), UTM: 31U GS0739, 5.x.1988, Lotus corniculatus, stem-mines, el. 10.v.1989-18.v.1989, EvN no. 88156, van Nieukerken & Richter (RMNH); 1 6, St. Pietersberg, Cannerbos (Lim- burg), UTM: 31U FS8733, 18-19.vii.1950, [22.3-0.3 hrs] (RMNH); 1 6, St. Pietersberg, Zonneberg (Limburg), UTM: 31U FS8934, 20.vii.1950, [21-22 hrs] (RMNH); 1 d, 19,2 larvae, Wijlre, 1 km SE: railway banks (Limburg), UTM: 31U GS0534, 5.x.1988, Lotus corniculatus, stem- mines, e.l. 11.v.1989-15.v.1989, EvN no. 88162, van Nieu- kerken & Richter (RMNH); 1 4, 1 9, Vrakelberg, 2km E. of Wijlre, UTM: 31U GS0537, 22.1x.1989, Lotus corniculatus, stem-mines, el. 1-5.v.1990, E. J. van Nieu- kerken (RMNH); mines, Noord-Bakkum, 3 km N Castri- cum (Noord-Holland), dune meadows, UTM 31U FU1226, 6.x.1989, Lotus corniculatus, E. J. van Nieu- kerken. - Rumania: 1 g, Gusterita near Sibiu (Hammers- dorf near Hermannstadt), UTM: 35T KL87, 12.v.1920, (NHMW); 1 g, Sibiu (Hermannstadt, Pralb.), UTM: 35T KL87, 18.v.1922 (NHMW). - Spain: 1 6, Beuda (Pyr. orient.), UTM: 31T DG77, 14.vii.1967, E. Arenberger (LNKD); 1 4, Biescas (Huesca), UTM: 30T YN12, 1.viii.1989, C. Gielis (coll. Gielis); 3 &, Puerto de Mora (Granada), 1350 m, UTM: 30S VG52, 22.vii.1986, C. Gie- lis (RMNH, coll. Gielis); 1 &, San Miguel de Valero N, 3 km S Linares de Riofrio (Salamanca), 850 m, UTM: 30T TK59, 2.vili.1986, at light ML, Quercus pyrenaica forest & heathland, EvN no. 86091, EJ. v.Nieukerken & S.Richter (RMNH); 1 6, Sierra de Marbella, El Mirandor (Malaga), 700 m, UTM: 30S UF34, 21.vii.1981, E. Trau- gott-Olsen (ETO); 1 8, road Baza-Benamaurel, 15 km from Baza (Granada), UTM: 30S WG25, 16.vii.1987, Bal- dizzone & Traugott-Olsen (GBA); 1 8, road to (Camino de) Ojen (Málaga), 150 m, UTM: 30S UF34, 25.vi.1983, E. Traugott-Olsen (ETO). - Sweden: 2 &, 1 > Byrum Sandvik (Öland), UTM: 33V XD1742, 20.vi.1978, B. Bengtsson (coll. Johansson, ZMUC); 1 © [slide only], Kinnekulle (Vg), UTM: 33V VEO9, 29-30.v.1968, I. Svensson (coll. Svensson); 1 @, Klagshamn (Sk), UTM: 33V UB65, 7.viii.1982, R. Johansson (coll.Johansson). — Soviet Union: 1 8, Dobroe, Krasnolesje [near Simfe- ropol] (Krim, Ukraine), UTM: 36T XQ06, 10.v.1982, Zaguljaev (ZMAS); 1 g, Rakvere (Raustfer) [near Tal- linn] (Estonia), UTM: 35V MF68, 8.vi.1890 (ZMAS); 1 3, Sevastopol, Inkerman (Krim, Ukraine), UTM: 36T WQ44, V. Pliginski (ZMAS). - Tunisia: 1 g, Nefta, UTM: 32S LC94, 14-16.111.1986, Zool. Mus. Copenh. (ZMUC). - Turkey: 2 &, Ankara, 10 km NW Kizilcahaman, 1150- 1250 m, UTM: 36T VK68, 6-7.viii.1989, Fibiger & Esser (ZMUC). - Yugoslavia: 1 6, Herceg Novi, Igalo, UTM: 34T BN90, 1.v.1938, H. G. Amsel (LNKD); 16 6, Krk, Draga Baska (Kroatia), UTM: 33T VK78, 25 and 27.vii.1976,3.viii.1976, l.viii.1977, 3.vitt.1978, 5.v111.1985, 30.vii.1986, 15.viii. 1988, G. Baldizzone (GBA, RMNH); 6 VAN NIEUKERKEN: Trifurcula subnitidella group Fig. 68. Distribution of Trifurcula subnitidella, mapped on 50 X 50 km UTM squares. 8, 2 9, Krk, Misucaynica (Kroatia), UTM: 33T VK78, 18.viii. 1973, 30.vii. 1976, 6.viii.1976, 4 and 9.viii.1986, G. Baldizzone (GBA, RMNH); 1 4, Krk, Paprata/Vinder (Kroatia), UTM: 33T VK79, 16.viii.1978, G. Baldizzone (GBA); 1 8, Krk, Soline (Kroatia), UTM: 33T VK69, 11.viii.1976, G. Baldizzone (GBA); 8 4, Krk, road Krk- Vrbnik (Kroatia), UTM: 33T VK78, 2.viii.1987, 23.v11.1988, 3.vitt.1988, G. Baldizzone (GBA, RMNH) Additional records. — Czechoslovakia: Celechovice (UTM: 33U XQ78), Prostéjov Hamry (33U XQ48), Ko- zarovce (34U CU15) (Z. Laötúvka in litt.). - Denmark: Melby Overdrev (Buhl et al. 1983), Korevle (Buhl et al. 1984), Glatved (32V PH14)(Buhlet al. 1987a), Oster Vra (32V NJ75)(Buhl et al. 1988). - Great Britain: vice coun- a 1, ORTO MEI AS IGLO MG ORS D Pan dell03 (Emmet in litt. 1987). - Sweden: Hall. Vallda Sandö (PJ7674), Sm. Ryningsnäs (WD5647), Ol. Algutsrum (WC9284), Gel. Tingstäde (CK5802), Sdm. Osmo (XF6739), Upl. Fysingen (XG6407) (Svensson in litt. 1987). 7. Trifurcula josefklimeschi sp. n. (figs. 6, 20-22, 58, 69-76, 93, 99, 104, 105) [ Trifurcula orientella Klimesch, 1953: 168, 169 partim: all specimens from Austria. Misidentification. ] Trifurcula nov. spec., van Nieukerken in Kasy 1983: 5. Type material. - Holotype &: Italy, M. Bondia, Villa Faraldi, 3.5 km WNW Andora (Imperia), 550 m, UTM: 32T MP2769, 10.iv.1988, Dorycnium pentaphyllum, stem-mines, e.l. 20.v.1988, EvN no. 88111, E. J. van Nieu- kerken (RMNH). - Paratypes: 43 &, 13 9, 25 larvae. — Austria: 1 @, Deutsch Altenburg, Pfaffenberg (Niede- röst.), UTM: 33U XP43, 24.viii.1935, [Preissecker] (NHMW) [paralectotype orientella Klimesch]; 8 8, Hundsheimer Berg (Niederöst.), UTM: 33U XP43, 17.viii.1973, 28.vi.1976, 9.vi.1977, 3.vii.1977, 3.1x.1977, 22.v.1979, 12.ix.1979, E Kasy (NHMW, RMNH); 1 6, Marchegg, Oberweiden (Niederöst.), UTM: 33U XP44, 6.1x.1931, [Preissecker] (NHMW) [ paralectotype orzen- tella Klimesch]; 1 &, Mödling, Frauenstein (Niederöst.), UTM: 33U WP92, 26.vi.1902, Preissecker (NHMW) 225 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 | paralectotype orzentella Klimesch |. - Czechoslovakia: 2 4, Hrädok n. V. (Slovakia), UTM: 33U YP19, 17.vi. 1988, A. Lastivka; 2 8, Zádiel (Slovakia), UTM: 33U DU88, 7.vi. 1989, A. Lastúvka (coll. Laëtuvka, RMNH).- Greece: 1 8, Kardhamili (Messinias), gorge, UTM: 34S FF1084, 18.11.1990, Dorycnium hirsutum, stem-mines, el. 15.iv.1990, EvN no. 90063, E. J. van Nieukerken (RMNH). - Italy: 1 8, Andora, Testico (Savona), UTM: 32T MP27, 2-5.vii.1983, B. À. Bengtsson (coll. Bengts- son); 3 4, 2 9, Conna (Savona), 300 m, UTM: 32T MP27, 10.ix.1977, 1.1x.1978, 22.vi.1979, G. Baldizzone (GBA, RMNH); 4 8,69, 1 larva, M. Bondia, Villa Faraldi, 3.5 km WNW Andora (Imperia), 550 m, UTM: 32T MP2769, 10.iv.1988, Dorycnium pentaphyllum, stem- mines, e.l. 19.v.1988-4.vi.1988, EvN no. 88111, E. J. van Nieukerken (RMNH); 5 8, Poggio di Casasco, Val Cu- rone (Al), 300 m, UTM: 32T NQ06, 06.vi.1987, G. Bal- dizzone (GBA, RMNH). - Spain: 1 6, 1 9, 6 larvae, 2.5 km SW Beires, along road Beires-Fondón (Almería), 1000 m, UTM: 30S WF1495, 10.1.1988, Dorycnium pen- taphyllum, stem-mines, e.l. 20.v.1988-26.vi.1988, EvN no. 88017, van Nieukerken & Richter (RMNH); 1 4, Collado de Falset (Cataluna), UTM: 30T CF15, 3.vii.1967, E. Arenberger (LNKD); 1 4,29, 18 larvae, Sierra Blanca, Refugio de Juanar, 3 km NW Ojén (Málaga), 840 m, UTM: 30S UF3149, 15.1.1988, Dorycnium hirsutum, stem-mines, e.l. 20.v.1988-8.vi.1988, EVN no. 88048, van Nieukerken & Richter (RMNH); 2 9, Sierra de Marbella, El Mirandor (Málaga), 700 m, UTM: 30S UF34, 21.vii.1987, E. Traugott-Olsen (ETO); 2 8, road Baza- Benamaurel, 15 km from Baza (Granada), UTM: 30S WG25, 16 and 17.vii.1987, Baldizzone & Traugott-Olsen (GBA). - Soviet Union: 1 8, Parkovo, southcoast Crimea (Krim), UTM: 36T WQ(southeast), 26.v.1984, Zaguljaev (ZMAS). - Yugoslavia: 7 @, Krk, Draga Baska (Kroatia), UTM: 33T VK78, 27.vii.1976, 28.viii.1978, 15.viii.1988, G. Baldizzone (GBA, RMNH); 1 8, Krk, Skrpcici (Kroa- tia), UTM: 33T VK68, 04.viii.1977, G. Baldizzone (GBA). Additional material. - Spain: mines, 3 km NE Mar- bella, road to Ojén (Málaga), 200 m, UTM: 30S UF3244, 15.1.1988, Dorycnium hirsutum, EvN no. 88046, van Nieukerken & Richter (RMNH). Additional records. - Czechoslovakia: Cebovce (UTM: 34U CU63), Jablonov (34U DV83), Kiarov (34U CU83), Hustopete (33U XQ22), Zajeti, larvae and adults, 1990, leg. A. and Z. LaStuvka (Z. LaStuvka in litt.). Description Male (fig. 6). - Forewing length 2.3-3.0 mm (2.57 + 0.18, 26), wingspan 4.9-6.5 mm. Head: frontal tuft yellowish white to pale yellowish orange, collar paler. Antenna with 37-43 (40.1 + 2.1, 22) seg- ments; scape white, usually with some brown scales. Forewing and thorax covered with fuscous tipped scales, distal 4 to % of scales dark, edge darker, total impression greyish brown, occasionally few white scales exposed at dorsum; terminal cilia white beyond indistinct cilia-line. Underside of forewing with very large costal fold along basal third, com- pletely covering a pocket with yellow androconial scales of about one quarter wing width (figs. 20, 58). Hindwing with pronounced humeral lobe, along costa with long snow-white hair-scales, inserted on underside, in rest fitting in forewing fold; imme- 226 diately behind frenulum a small group of closely set, short dark fuscous special scales (figs. 21, 22, 58). Female. - Forewing length 2.2-2.8 mm (2.52 + 0.19, 11) wingspan 4.9-6.1 mm. Antenna with 35- 38 (36.7 + 1.3, 10) segments. Male genitalia (figs. 69-75) - Capsule length 340- 385 wm (363.6 + 14.5, 11). Vinculum anteriorly truncate. Uncus with medial process distinctly wi- dened terminally and truncate. Gnathos slightly or not asymmetrical, central element narrowly spatu- late, with a medial keel ventrally, margins often serrate, additional spines may occur on ventral sur- face, but specimens without any spines and smooth margin do occur (figs. 69, 72, 73); anterior apo- demes conspicuous. Valva length 260-290 um (273.6 + 8.3, 11), almost triangular, with inward curved tip; dorsally with a row of several spines along continuation of lateral arm of transtilla, transverse bar of transtilla slightly longer than sub- lateral processes to twice as long. Aedeagus 335-385 um (360.0 + 17.3, 11) long, with ventral carina fringed or bifid, aedeagal tube terminally spatulate, dorsal lobe prominent at right side, with serrate margin; vesica with two large spine-like cornuti, one (155-200 um, 175.5 + 13.5, 10) basically joined to a conical cornutus (50-90 um, 59 + 11.4, 10), the other subterminally bent, partly hidden amongst group of spine-like cornuti (not measurable); small cornuti numerous. Juxta fig. 70. Female genitalia (figs. 93, 99). - Terminalia dis- tinctly pointed. T8 with 6-15 setae and some scales; posterior margin truncate; anal papillae with 9-21 setae. Anterior apophyses laterally widened. Ductus spermathecae with 4/2 to 5 coils. Signa 340-500 um long, almost completely similar in length. Final instar larva (fig. 104) - Yellow, elongate. Headcapsule 375-425 um long, 1.01-1.11 (1.06 + 0.03, 6) X as long as wide (in one specimen 1.19 X). Headcapsule distinctly longer than in subnitidella and coronillae. Spinosity: prothorax ventrally with spinose band posteriorly, meso- and metathorax ventrally with a band along anterior margin and few spines laterally. Abdominal segments 1-9 with spines ventrally, not conspicuous; segment 10 with small group of spines anterior of setae. Diagnosis Male immediately recognized by large costal fold, hiding the yellow patch (N.B. this fold may be difficult to see in well mounted specimens!), the humeral lobe of the hindwing and the black scales near the frenulum. Male genitalia in particular characterized by gnathos with medial keel, but note variability in gnathos! Female similar to other spe- cies in the group, but easily separated by the pointed postabdomen. VAN NIEUKERKEN: Trifurcula subnitidella group Figs. 69-75. Trifurcula josefklimeschi, male genitalia. - 69, Capsule, valva outlined, slide 2590; 70, Juxta, slide 2590; 71, Valva, dorsal aspect, slide 2590; 72, 73, Gnathos, showing variation, respectively slides 3068 and 2744 (holotype); 74, Aedeagus, ventral aspect, holotype, slide 2744; 75, Large cornuti, separately drawn, holotype. Scale 0.1 mm. Biology Hostplants. - Dorycnium hirsutum (L.) Ser. and D. pentaphyllum Scop., both perennial herbs or small shrubs, widely distributed in southern Eu- rope, probably also on the other two European Dorycnium species (D. rectum (L.) Ser. in DC. and D. graecum (L.) Ser. in DC.). Life history. - Egg usually deposited on the un- derside of a leaflet, occasionally on the stem. When in a leaflet, the larva starts with a short and narrow linear mine (fig. 105), leading either in a straight or slightly contorted course towards the petiole, through which it enters the stem (fig. 105). The larva then usually feeds downward first, in a rather straight line, often doubling back once or twice, before the larva quits the mine through a semicir- cular slit. The whole course of the mine often stain- ing reddish or brown, frass hardly visible externally, larva visible as a yellow swelling in the stem. The mine is frequently in the more terminal shoots, and can relatively easily be found because of the stain- ing. When full-grown larvae or empty mines are collected, the leaflet with the early mine has often already been fallen. Larvae have been found in January, February and early April. Adults emerged in mid April (from 227) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 © josefklimeschi V victoris Fig. 76. Distribution of Trifurcula victoris and T. josefklimeschi, mapped on 50 X 50 km UTM squares. Greece), the second half of May and in June (indoor rearing), but were caught from late May until the 12th of September, the largest numbers being taken in June and from late August until early September. This pattern might indicate a bivoltine cycle, but this should be confirmed by finding larvae in summer. Distribution (fig. 76) Widely distributed in southern Europe, from southern Spain eastward to the Crimea, northwards to eastern Austria and Slovakia. Not yet recorded, but to be expected from Portugal, France, Hungary, Rumania, Bulgaria and Turkey. Remarks Material of this species has long been known in collections, but was incorrectly identified as Trifur- cula orientella Klimesch. In fact, all four specimens from Austria, listed amongst the type-material by Klimesch (1953: 169) appeared to belong to this species, which clearly differs from the spec- imen which genitalia were illustrated by Klimesch (1953: fig. 15) and is selected lectotype of orientella below. Etymology A noun in genitive case, named in honour of Dr. Josef Klimesch, who did much pioneer work on the genus Trifurcula and on mediterranean Nepticuli- dae in general. 228 8. Trifurcula iberica sp. n. (figs. 29, 59, 77-82) Type material. - Holotype @: Spain, Sierra Nevada, road to Veleta (Granada), 1700 m, UTM: 30S VG60, 9.vii.1971, E. Arenberger (LNKD). - Paratype: 1 &: Spain, Pyrenees, Prullans, 900 m, UTM: 31T CG99, 2.vii.1980, G. Derra (coll. Derra). Description Male. - Forewing length 2.7-3.2 mm, wingspan 6-7.2 mm. Head: frontal tuft ochreous yellow, collar paler. Antenna with 37-40 segments; scape yellow- ish white, with some brown scales. Forewing and thorax covered with brown tipped scales, total im- pression pale brown; terminal cilia white beyond more or less distinct cilia-line. Underside of fore- wing with basal patch of yellow androconial scales extending to 4. Hindwing with white hair-scales instead of costal bristles; along costa a row of short but conspicuous black special scales, running from 4 to middle of hindwing (fig. 59). Female unknown. Male genitalia (figs. 77-82) - Capsule length 345- 355 um. Vinculum anteriorly truncate, rather nar- row. Uncus with medial process pointed, some lat- eral setae present. Gnathos almost symmetrical or slightly asymmetrical, central element truncate to rounded, posterior margin distinctly serrate; ante- rior apodemes present. Valva length 275 um, nar- row triangular, gradually narrowed towards tip, dorsal excavation less than half length; transverse bar of transtilla up to three times as long as sublat- VAN NIEUKERKEN: Trifurcula subnitidella group Figs. 77-82. Trifurcula iberica, male genitalia, slide 2741 (77-80) and 1928 (holotype: figs. 80, 82). - 77, Capsule, valva outlined; 78, Juxta; 79, Valva, dorsal aspect; 80, Large cornuti, separately drawn; 81, Aedeagus, ventral aspect; 82, Aedeagus, lateral aspect, position of spinelike cornutus shown. Scale 0.1 mm. Diagnosis The male of zberica is easily distinguished from other species in the group by the long row of black scales on the hindwing costa. The genitalia are characterized by the almost symmetrical gnathos with serrate margin, but resemble those of s¢/vzae, see there. Biology Hostplant and immature stages unknown. Adults taken in mountains, at medial (900 m) and high altitude (1700 m), both in July. Distribution (fig. 29) Only known from two specimens from the Span- ish Pyrenees and Sierra Nevada. Etymology A latin adjective, from Ibericus (= Hibericus), meaning Spanish, referring to the single localities in the Spanish mountains. 229 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 9. Trifurcula silviae sp. n. (figs. 7, 29, 60, 83-88, 94, 100) Type material. - Holotype 4: France, 1 km NW Ceillac, + 10 km S Chateau-Queyras (Htes Alpes), 1800 m, UTM: 32T LQ2449, 24.vii.1987, alpine meadow on S. slope, netted at dusk, EVN no. 87139, van Nieukerken & Richter (RMNH). - Paratypes: 6 6, 1 9. France: 1 8, data as holotype (RMNH); 1 6, Chaner. [not traced, ?near Digne], 22.viii.1903, Chrétien (MNHN); 2 @, Digne (Alp. Ht. Prov.), UTM: 32T KP78, viii.1903, (MNHN); 1 4, Puy Vacher [near La Grave] (Htes Alpes), [2000 m], UTM: 32T KQ88, 29.vi.1898, (MNHN); 1 6, Viens (Vau- cluse), UTM: 31T GJ0664, 21.viii.1974, R. Buvat (coll. Buvat); 1 9, same data, but 8.viii.1975 (RMNH). Description Male (fig. 7). — Forewing length 2.6-3.1 mm (2.80 + 0.18, 6), wingspan 5.8-7.0 mm. Head: frontal tuft pale yellow, mixed with white, collar yellowish white. Antenna with 35-39 segments (36.5 + 1.5, 6) ; scape white. Forewing and thorax relatively pale ochreous yellow, scales with ochreous brown tips mixed with completely white scales; terminal cilia white, cilia-line obsolete or absent. Underside of forewing without patch of androconial scales, narrow costal fold relatively well developed, edged with row of brown scales (fig. 60). Hindwing al- most white, costa of humeral lobe with row of short dark brown scales, running to % (fig. 60). Female. - Forewing length 2.6 mm, wingspan 5.8 mm. Antenna with 32 segments. Male genitalia (figs. 83-88) - Capsule length 300- 330 um. Vinculum anteriorly truncate. Uncus with medial process truncate, not widened, with several lateral setae. Gnathos slightly or not asymmetrical, central element narrow, posterior margin distinctly serrate; anterior apodemes hardly visible. Valva length 215-240 um, narrow triangular, tip short pointed, slightly curved, transverse bar of transtilla about 1.5 times as long as sublateral processes. Ae- deagus 310-340 wm long, with ventral carina fringed, aedeagal tube apically pointed, dorsal lobe at right side prominent, margin serrate; vesica with long pointed cornutus (140-210 um), basally joined to conical cornutus (75 um), a long cornutus with serrate tip hidden amongst group of long needle- like cornuti, many small cornuti present. Juxta fig. 84. Female genitalia (figs. 94, 100). - Terminal seg- ments broadly rounded. T8 with about 6 setae and few scales; anal papillae with 23-25 setae. Ductus spermathecae with 4 coils. Signa 335-365 um long. Diagnosis Male differs from all other species in the subni- tidella group by the absence of the yellow patch, it is also paler than most species, except /uteola. T. silviae male differs from other pale Trifurcula spe- cies of similar size by the distinct rows of brown 230 scales along costal fold of forewing and along hindw- ing costa. Male genitalia differ from similar iberica by relatively longer sublateral processes and slightly broader valva. Female differs from other species described here by paler colour and blunt ovipositor. Biology Hostplant and immature stages unknown. The holotype and one paratype were collected in an alpine meadow on a steep southern slope, where the Fabaceae Lotus corniculatus L., Anthyllis vulne- raria L. and Onobrychis montana DC. are the most likely candidates to be its host. The species appar- ently occurs over a wide range of habitats, from almost lowland mediterranean localities (Viens, Digne), to high alpine country (Ceillac: 1800 m, Puy Vacher: ca 2000 m.). Adults have been caught from June to August. Distribution (fig. 29) Only known from a relatively small area in sou- theastern France, in the Alps and pre-alps. Remarks The specimens in the Chrétien collection were found amongst material of T. immundella, together with a number of other misidentified species. The single female is considered to belong to si/viae on the basis of its occurrence with one male and no other related species, the external characters and the genitalia, which clearly belong to a species in the subnitidella group. Etymology A noun in genitive case, named in honour of my wife Silvia Richter, who not only collected both holotype and one paratype, during an alpine moun- tain hike, but contributed much to the collecting during many joint field trips. PHYLOGENY OF THE TRIFURCULA SUBNITIDELLA GROUP The phylogeny of the genus Trifurcula and its three subgenera has been dealt with by Scoble (1980) (with the present subgenera as genera) and van Nieukerken (1986b). In these studies it has been shown that the subgenus Glaucolepis Braun (=Fedalmia Beirne) is the sistergroup of [Levarchama Beirne + Trifurcula s.str.| and that the latter two subgenera are sistergroups to each other. Both the monophyly of the genus and subge- neric clades has been amply demonstrated by apo- morphies, to which little is to be added. Only one more apomorphy for Levarchama has been menti- oned by van Nieukerken & Johansson (1990): the costal hair-pencil on the hindwing underside. The apomorphy for Trifurcula s. str: uncus divided dor- soventrally needs to be re-examined: in my opinion VAN NIEUKERKEN: Trifurcula subnitidella group Figs. 83-88. Trifurcula silviae, male genitalia. - 83, Capsule, valva outlined, holotype, slide 2742; 84, Juxta, slide 2604; 85, Valva, dorsal aspect, holotype; 86, Large cornuti, separately drawn, slide 2606; 87, Aedeagus, ventral aspect, slide 2606; 88, Aedeagus, lateral aspect, holotype. Scale 0.1 mm. the ventral less sclerotized part, interpreted as part of the uncus by Scoble (1980), might well be a somewhat better sclerotized anal tube. This needs to be confirmed by detailed examination of whole mounts of complete genitalia, and might still be an apomorphy for the subgenus. In any case, a new interpretation of this character does in no way jeo- pardize the monophyly of Trifurcula s. str. The phylogeny within this subgenus has not been discussed before, except the position of the two South African species, discussed by Scoble (1980), who could not solve the question whether they together form the sistergroup of |Levarchama + Trifurcula] or the sistergroup of Palaearctic Trifur- cula s. str. I will come back to that question below, and first concentrate on the Palaearctic species. For an evaluation of the polarity of characters, Levarchama has been used as outgroup. In this study Trifurcula s. str. has been divided in two species groups, which are believed to be monophy- letic sistergroups. The phylogeny of the swbnitidella group is presented in the cladogram in fig. 108. Monophyly of the szbnitidella group is actually based on one character only: 1. Male forewing with patch of yellowish androco- nial scales on underside, near wingbase. Such andro- conial scales are absent in Levarchama and in the pallidella group, and not completely similar scales are only found in a subgroup of Glaucolepis, so that 2511 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 89-94. Trifurcula spp., female postabdomen, dorsal aspect. - 89, T. puplesisi, slide 2763; 90, T. coronillae, slide 2750; 91, T. victoris, slide 2751; 92, T. subnitidella, slide 2746; 93, T. josefklimeschi, slide 2748; 95, T. silviae, slide 1818. Scale 0.1 mm. 232 VAN NIEUKERKEN: Trifurcula subnitidella group 95 100 Figs. 95-100. Trifurcula spp., female genitalia. - 95, T. puplesisi, slide 2763; 96, T. coronillae, slide 2750; 97, T. victoris, slide 2752; 98, T. subnitidella, slide 2746; 99, T. josefklimeschi, slide 2748; 100, T. silviae, slide 1818. 255 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 101 103 102 Figs. 101-104. Trifurcula spp., final instar larva, headcapsule. - 101, T. coronillae, Spain, Istan; 102, T. victoris, Spain, San José; 103, T. subnitidella, Netherlands, Kunrade; 104, T. josefklimeschi, Spain, Sierra Blanca. the presence of these scales is regarded as an evo- lutionary novelty. They are secondarily lost in s2/- viae, which can be regarded to belong to the szbnz- tidella group on the basis of other characters. The pallidella group is believed to be monophy- letic on the basis of at least the following apo- morphy: 2. Hostplant belonging to the Genisteae (‘brooms’). The hostplants of Levarchama and the subnitidella group belong to other tribes of Fabaceae: Loteae and Coronilleae, thus on the basis of the outgroup rule, a hostplant belonging to Loteae or Coronilleae is the plesiomorphous condition, and hence Genis- teae apomorphous. No other Nepticulidae are known to feed on brooms. Although only few spe- cies have actually been reared, evidence is accumu- lating that indeed all species are associated with brooms. 234 3. A doubtful apomorphy is the form of the larger cornuti: two are curved and one is spine-like in virtually all species. The current position of the first species of the subnitidella group, T. austriaca, is tenuous. It is included here solely on the basis of the yellow patch and the general external resemblance to other spe- cies in the group. The genitalia, in particular the aedeagus, resemble more the pallidella group, al- though they show some peculiarities, not noticed in other species, such as the curved carinal lobes. If the aedeagal characters (shape of cornuti, dorsal lobe) belong to the groundplan of the subgenus, and aus- triaca has no broom species as host, it still belongs to the subnitidella group, and possibly is the sister- species of all other species together. On the other hand, if it is found to have a species of broom as VAN NIEUKERKEN: Trifurcula subnitidella group Figs. 105-107. Larval stem-mines of Trifurcula species. - 105, T. josefklimeschi, mine in Dorycnium hirsutum, Spain, Sierra de Marbella; 106, T. coronillae, mines in Coronilla juncea, type locality; 107, T. subnitidella, mines in Lotus corniculatus, Netherlands, Kunrade. Scale 1 cm. 295 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 AN No S d 9 9 9 ov A) A AV . è n ot a 0 NI oÙ . LN x av R nv CC o N N A N° N Fig. 108. Cladogram of Trifurcula subnitidella group, numbers refer to apomorphies, explained in text. host, it is possibly misplaced here meaning that the androconial patch has apparently evolved twice in- dependently, or that it should be regarded as an underlying apomorphy. Awaiting further informa- tion of this species, austriaca is tentatively treated here in the szbnitidella group. The remaining eight species share the following characters as apomorphies: 4. Aedeagus terminally narrowed, dorsal lobe well developed, at right side or in middle; often with serrate margin. 5. Pattern of the three large cornuti: one spine-like, basally joined with a conical, and a long irregular (often spine-like) cornutus hidden amongst the group of needle-like cornuti. The first branching in this clade has not been resolved, and is provisionally presented as a tricho- tomy. The species luteola and puplesisi resemble each other closely, but the similarities seem to be mere plesiomorphies, so that their phylogenetic relationship cannot be demonstrated. The remain- ing six species form a better defined monophyletic entity on the basis of the apomorphy: 6. Gnathos highly atypical for Nepticulidae, with keels and/or serrations, often highly asymmetrical. The gnathos of the previous species resembles that of the pallidella group, it is symmetrical and never with keels, and thus forms the plesiomorphic con- dition within the subgenus. Also in the next clade the first branching is pres- ented as a trichotomy, since the monophyly of [coronillae + victoris] could not be demonstrated. The monophyly of the final four species is based on the following unique character: 7. Hindwing costa in male with characteristic black or dark brown androconial scales T. subnitidella and josefklimeschi are tentatively regarded as sister-species on the basis of their sim- ilarity and one possible apomorphy: 236 8. Black scales along hindwing costa confined to small group close to frenulum. Finally, T. zberica and silviae most likely are sis- ter-species, also on the basis of a general similarity in habitat, externals and genitalia of which the following character might be regarded as apo- morphy: 9. Gnathos without keels, but with serrated margin. The phylogeny of the szbnitidella group is still based on relatively few characters, more detail of females, immature stages and hostplants would be most welcome in order to refine the cladogram. One final point to discuss here, is the phyloge- netic position of the two South African species, described by Scoble (1980). Only males are known from these species, so that we have no biological data to support our theories. In my opinion both species definitely belong to Trifurcula s. str. on the basis of the group of needle- like cornuti, an apomorphy for the subgenus, and the absence of a split uncus (in ventral view), a character for Levarchama. However, the hair-pen- cil of T. barbertonensis Scoble (fig. 12 in Scoble), closely resembles that of Levarchama, which might suggest another solution. Since no biological data are available, and since it is unknown whether these species have a patch of androconial scales on the forewing underside, it is yet impossible to assign them to one of the Palaearctic species groups. The aedeagus as figured by Scoble (1980) seems to be different from both species groups, so that it is still possible that the South African species form the sistergroup of the Palaearctic Trifurcula s. str. APPENDIX Trifurcula orientella Klimesch Trifurcula orientella Klimesch, 1953: 168, 169, fig. 15. Lectotype & (here designated): Yugoslavia, Dalmatia mer., Umg. v. Gravosa, 15-31.v.1939, J. Klimesch, Ho- lotypus’, genitalia slide Kl. 511 (coll. Klimesch) [examined]. Trifurcula orientella was described on the basis of six male specimens from Gravosa, Yugoslavia and four male specimens from Austria. When I found the austrian specimens in the Vienna museum, they appeared to have genitalia differing from the figure, presented by Klimesch, and actually appear to be- long to the species here described as josefklimeschi. Since Klimesch’s figure clearly has been the basis for the identity of orientella, it seems most approp- riate to select the specimen used for this figure as lectotype. This is also the specimen labelled by Klimesch ‘Holotypus’. Unfortunately, two other paralectotypes from the type locality, examined by me, also appear to belong to another and larger species, either T. immundella Zeller or a closely related species, which has been reared by me from Calycotome. The remaining three paralectotypes have not been examined by me. T. orientella is a relatively small species, clearly belonging to the pallidella group, which is known from the Dalmatian coast, the isle of Krk (leg. Baldizzone) and from Conna on the Italian Riviera. From the latter locality I have reared this species from stem-mines in the broom Genista germanica IE, ACKNOWLEDGEMENTS The author is indebted for the loan of material and information on distribution to the following persons: G. Baldizzone (Asti, Italy), G. Bassi (To- rino, Italy), B. A. Bengtsson (Löttorp, Sweden), R. Buvat (Marseille, France), G. Derra (Bamberg, Ger- many), A. M. Emmet (Saffron Walden, UK), R. Gaedike (DEIC, Eberswalde, Germany), C. Gielis (Lexmond, Netherlands), B. Gustafsson (NHRS, Stockholm, Sweden), R. Johansson (Växjö, Sweden), O. Karsholt (ZMUC, Copenhagen, Den- mark), F. Kasy (NHMW, Vienna, Austria), S. Kerp- pola (Helsinki, Finland), J. Klimesch (Linz, Aus- tria), J. H. Kuchlein (Wageningen, Netherlands), G.R. Langohr (Simpelveld, Netherlands), A. and Z. Lastúvka (Brno, Czechoslovakia), G. Luquet (MNHN, Paris, France), U. Parenti (Torino, Italy), P. Passerin d’Entreves (Torino, Italy), R. Puplesis (ZKVV, Vilnius, Lithuania), U. Roesler (LNKD, Karlsruhe, Germany), H. Steuer (Bad Blankenburg, Germany), I. Svensson ((Kristianstad, Sweden), E. Traugott-Olsen (Marbella, Spain), K. Tuck (BMNH, London, UK) and H. W. van der Wolf (Nuenen, Netherlands). Ole Karsholt and Jan van Tol are acknowledged for critical remarks on the manuscript. Mrs. Ingrid Henneke assisted with photographic work. REFERENCES Arnett, R. H. & G. A. Samuelson, 1986. The insect and spider collections of the world. 220 pp. - EJ. Brill/Flora & Fauna publications, Gainesville. Bjorn, & G. Pallesen, 1971. Fund af smäsommerfugle fra Danmark i 1970. - Flora og Fauna, Arhus 77: 107-111. Bradley, J. D., 1962. Lepidoptera in Ireland, May, 1961. 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Anmärkningsvärda fynd av Microlepi- doptera i Sverige 1979. - Entomologisk Tidskrift 101: 75-86. Svensson, I., 1983. Anmärkningsvärda fynd av Microlepi- doptera i Sverige 1982. - Entomologisk Tidskrift 104: 59-65. Svensson, I., 1985. Anmärkningsvärda fynd av Microlepi- doptera i Sverige 1984. - Entomologisk Tidskrift 106: 71-82. Svensson, I., H. Elmquist, B. Gustafsson, H. Hellberg, L. Imby & G. Palmqvist, 1987. Catalogus Lepidopterorum Sueciae. Kodlista LI. 11 + 1 +9 + 96 + 28 + 3 + 27 + 6 pp. - Naturhistoriska Riksmuseet, Stockholm. Wagner, D. L., 1987. A new Microcalyptris species from California (Lepidoptera: Nepticulidae). - Pan-Pacific Entomologist 63: 278-283. Wolff, N. L., 1957. Trifurcula griseella nov. spec. (Lepi- doptera, Nepticulidae).- Entomologiske Meddelelser 28: 19-22. Zeller, P. C., 1848. Die Gattungen der mit Augendeckeln versehenen blattminirenden Schaben. - Linnaea ento- mologica 3: 248-344. Received: 26 September 1990. Accepted: 1 October 1990. ERIK J. van NIEUKERKEN National Museum of Natural History, Leiden STIGMELLA ROLANDI SP. N.: A WIDESPREAD SOUTHERN EUROPEAN SPECIES ON ROSA (LEPIDOPTERA: NEPTICULIDAE) Nieukerken, E. J. van, 1990. Stigmella rolandi sp. n.: a widespread southern European species on Rosa (Lepidoptera: Nepticulidae) — Tijdschrift voor Entomologie 133: 239-243, figs. 1-10. [ISSN 0040-7496]. Published 14 December 1990. Stigmella rolandi sp. n., belonging to the Stigmella sanguisorbae group, is described from southern Europe. It has previously been misidentified as S. spinosissimae Waters, a western European species. The species is characterized by a costal hair pencil on the male hindwing. The distribution is mapped, and the biology described: the larva feeds on Rosa and Sangut- sorba. E. J. van Nieukerken, Rijksmuseum van Natuurlijke Historie, Postbus 9517, 2300 RA Leiden, Netherlands. Key-words. - Nepticulidae, leaf-miners, taxonomy, Palaearctic, Rosaceae. The aim of this paper is to name a widespread species of Stigmella, which has been known for almost 40 years, but until recently was misidentified as Stigmella spinosissimae (Waters). This misiden- tification followed Klimesch (1951), who described the genitalia and biology of the present species, which he identified as S. spinosissimae, on the basis of the same hostplant (Rosa pimpinellifolia L.) and the description of external features by Waters (1928). Study of type material of Nepticula spino- sissimae Waters, however, showed that this is a species in the anomalella species group, whereas spinosissimae sensu Klimesch belongs to the san- guisorbae species group (van Nieukerken 1986, Jo- hansson & Nielsen 1990). The European species of Stigmella Schrank are relatively well known: Johansson & Nielsen (1990) treated the 76 species of Northwest Europe in de- tail. Only 20 additional species from southern Eu- rope and the mediterranean region were listed by van Nieukerken (1986). Most of these have been satisfactorily described, including figures of male genitalia, and in some cases female genitalia, by Johansson (1971) and in papers by Klimesch (ref- erences in Johansson & Nielsen 1990). To date only about six more undescribed European species of the genus are known in collections, and not many more are expected to be found. Therefore, identification of European Stzgmella species can be achieved with a fairly high degree of certainty. In this light, des- cription of a single widespread new species seems justified and will facilitate identification. The other undescribed species belong to different species groups, mainly the ruficapitella and malella species groups, and will be described in due time. The Stigmella sanguisorbae group counts four species (van Nieukerken 1986): of these, S. sangui- sorbae (Wocke) and S. thuringiaca (Petry) have been described and illustrated in detail by Johansson & Nielsen (1990) and S. muricatella Klimesch in the original description (Klimesch 1978). The fourth species is (re)described and named below. The methods and abbreviations are largely the same as in the previous paper (van Nieukerken 1990), but genitalia measurements were taken at 400 X. Stigmella rolandi sp. n. (figs. 1-10) [Nepticula spinosissimae Waters; Klimesch 1951: 4, 1958: 95, 1961: 751, Sz6cs 1955: 170, 1956: 390, 1963: 108, 1965: 59, 1973: 452, 1977: 92, 1978: 267, 1981: 213. Misidentifications] Nepticula ‘spec. nov.’; Zimmermann 1944: 63. Stigmella spinosissimae sensu Klimesch; van Nieukerken 1986: 9. Type material. - Holotype @: Greece, Parnassós Oros, 5-6 km S. Polidhroson (Fokis), Abies-Pinus W., 1000-1200 m, 38.36N-22.33E [UTM: 34S FH37], 28.ix.1980, st. 59, Rosa el. 25/29.iv.1981, VU no 80690KE, S. B. J. Menken & E. J. van Nieukerken, Genitalia slide EvN 2780 (RMNH). - Paratypes: 81 4, 2 2. - Austria: 1 g, Dürn- stein, UTM: 33U WP46, el 8.iv.1936, J. Klimesch 239 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 1,2. Stigmella rolandi, male. - 1 (top), habitus hol- otype; 2 (bottom), Underside wings, showing special scales and hair-pencil, Austria. (RMNH). - Czechoslovakia: 1 8, Détkovice (Moravia), UTM: 33U X057, 30.viii. 1988, el iii.1989, Rosa; 18,19, Mikulov (Moravia), UTM: 33U XQ20, 29.ix.1989, el 11.1990, Sanguisorba minor; 2 6, SI. Kras-Zádiel, UTM: Z. Laötúvka) — France: 1 4, La Penne-sur-l'Ouvèze (Drôme), UTM: 31T FK70, 25-27.vi.1986 (coll. H. van der Wolf); 4 8, Viens (Vaucluse), UTM: 31T GJ06, 22.vii. 1964, 24. viii.197 1, 8.viii.1975, 18.viii.1976, R. Buvat (RMNH, coll. Buvat). — Italy: 1 @, Abruzzo, Sulmona (l'Aquila), UTM: 34T EL88, 7.viii.1982 (coll. J. H. Kuch- lein); 10 @, Monti Aurunci (Latina), 4 km NW Castel- forte, UTM: 33T VF07, 400 m, 22-23.vi, 1.vii.1969; 2 &, Monti Aurunci, 6 km N Itri, UTM: 33T UF77, 15.viii.1972 (coll. R. Johansson, RMNH); Sardegna, Mt. Istiddi, UTM: 31T NK12, 1.ix.1978 (coll. G. Derra); 1 6, Sardegna, Belvi (Nuoro), UTM: 31T:NK12, 650 m, 20.vii.1984; 1 6, Sar- degna, Fontana Raminosa (Nuoro), UTM: 31T NKI11, 900 m, 4.viii.1984; 3 8, Sardegna, Sarcidano (Nuoro), UTM: 31T NK00, 720 m, 1.vii.1984 (all. leg. and coll. J. H. Kuchlein). - Spain: 3 4, Cadalso de los Vidrios (Mad- rid), UTM: 30T UK8062, 7.viii. 1986, E. J. van Nieukerken (RMNH); 1 6,8 km E of Orcera (Jaen), 1150 m, UTM: 30S WH24, 19.vii.1986 (coll. Gielis); 5 &, San Roque (Cadiz), UTM: 30S TF81, 26.vii.1986, C. Gielis (RMNH, coll. Gielis); 2 &, Vega del Codorno (Cuenca), UTM: 30T WK97, 1350 m, 23.vii. 1985 (coll. J. H. Kuchlein, RMNH). — Yugoslavia: 1 g, Drenovo, Kavadarci (Macedonia), UTM: 34T EL88, 1-10.vi.1957, FE Kasy (NHMW); 31 &, 1 9, Krk, Misucaynica, road Krk-Vrbnik (Kroatia), UTM: 33T VK78, 20.vii-11.viii, 1986-1988, G. Baldizzone (RMNH, coll. Baldizzone); 3 &, Krk, Draga Baska (Kroa- tia), UTM: 33T VK78, 15.viii.1988 (coll. Baldizzone). — USSR, Ukraine: 1 8, Krim, Kara-Dag, 20 km W Feodosia, 240 UTM: 36T XQ78, 14.vii.1977, Reznik (ZMAS); 4 À, same locality, 5-22.vii.1987, R. Puplesis (ZKVV, RMNH). Description Male (fig. 1).- Forewing length 1.6-2.1 mm (1.88 + 0.11, 56), wingspan 3.7-4.7 mm. Head: frontal tuft pale yellowish orange to ferruginous, collar yellowish white. Antenna fuscous, with 25-29 seg- ments (26.6 + 1.1, 44); scape yellowish white. Fore- wing and thorax dark fuscous to fuscous black, scale bases often paler greyish; terminal cilia dark grey, occasionally separated by a more or less distinct cilia line. Underside of forewing (fig. 2) with an elon- gated andraconial patch, extending from base to %, on costal side of fold, with fuscous grey special scales, leaving a narrow furrow in middle; all scales outwards oblique towards furrow. Hindwing grey, first three to four costal bristles normal strong and short, followed by group of more than 20 hairlike costal bristles of Yo wing length, forming a hair- pencil (fig. 3), which in rest is inserted in furrow in forewing andraconial patch. Abdomen fuscous, with distinct yellowish grey anal tufts. Female. - Forewing length 1.8-1.9 mm (N=2), wingspan 4.1-4.2 mm. Antenna with 20-22 seg- ments. Forewing without special scales, hindwing with costal bristles of normal length. No anal tufts. Male genitalia (figs. 4-6) - Capsule length (from tip of tegumen to central part of anterior margin of vinculum) 163-210 um (188.4 + 12.5, 19). Vincu- lum with anterior margin emarginate. Tegumen hood-shaped. Uncus distinctly bilobed, lobes separ- ate, each with some setae on prominent sockets. Gnathos with posterior horns widely separate, transverse bar anteriorly slightly protruding at corners, forming indistinct anterior processes. Valva length 148-180 um (164.2 + 8.4, 17), widest beyond middle, suddenly tapering towards long curved distal process; transverse bar of transtilla long, sublateral processes small. Aedeagus 104-185 um (146.5 + 16.7, 17) long, tube broad, but slightly variable in dimensions. Vesica with relatively few small cornuti, some being a little larger. Female genitalia (figs. 7, 8) — T8 with three longitudinal bands of setae and scales; anal papillae without setae. Posterior and anterior apophyses long and narrow, approximately of same length. Bursa globular, densely covered with pectinations, no signum apparent. Accessory sac small, no retic- ulate field visible. Ductus spermathecae without distinct coils. Diagnosis The male can be recognized from all other uni- formly coloured Stigmella species by the coastal hairpencil and androconial scales on the forewing underside. From above S. roland: resembles S. san- guisorbae most, and females cannot be reliably se- parated. S. thuringiaca has paler olive-brown to grey-brown and more shining forewings, is some- what larger, and usually has a darker head. Not likely to be confused with species of the anomallela group, that feed on the same hosts: S. anomalella (Goeze) has distinct purplish wing tips, S. spinosis- simae (Waters) has a fuscous head and bronze fore- wings with copper reflections and S. centifoliella (Zeller) has a postmedial fascia. The male genitalia resemble also sanguisorbae most, but roland: has much less cornuti. S thurin- giaca has still more cornuti, plus pectinations on the vesica and pectinate hairs on the dorsal face of the valvae. Female genitalia with smaller accessory sac than sangwisorbae or thuringiaca and without sig- num. See illustrations in Johansson & Nielsen (1990). Biology Hostplants. - Rosa spp., including R. pimpinel- lifolia L. (= R. spinosissima L.) and Sanguisorba minor Scop. Mine (fig. 9). - A gallery mine. Egg deposited on under- or upperside, often near a vein. Mine often following the serrations of leaf-margin. Frass in midline, leaving very narrow white margins in early mine, but filling only about half mine width later. Larva yellow. Mines are difficult or not to separate from those of the anomalella group. Life history. - Probably bivoltine. Larvae have been found in late August and September, adults reared from February to April. Early summer larvae have not yet been found, but adults are found from early June until early September, thus at least partly from a probable second generation. Adults usually taken at light. Distribution (fig. 10) Widespread in southern and southern central Europe: eastern Austria, eastern Czechoslovakia, southern France, Spain, Italy, Sardinia, Yugoslavia, Greece and Soviet Union: Ukraine. Literature records: Hungary: Budapest region (Sz6cs 1955, 1956, 1963, 1981), Bakony mountains VAN NIEUKERKEN: Stigmella rolandi sp. n. Fig. 3. Stigmella rolandi, male hindwing: costal bris- tles and costal hair-pencil (Italy: Monti Aurunci). Scale: 0.2 mm. (Szöcs 1973), Matra mountains (Szöcs 1977) and Pilis mountains (Szöcs 1978). Etymology A noun in genitive singular. Named in honour of my friend Roland Johansson, specialist of Nepticu- lidae, and outstanding painter of these moths. ACKNOWLEDGEMENTS The author would like to thank the following persons for the loan and/or gift of material: G. Baldizzone (Asti, Italy), R. Buvat (Marseille, France), G. Derra (Bamberg, Germany), C. Gielis (Lexmond, Netherlands), R. Johansson (Växjö, Sweden), the late F Kasy (Wien, Austria), J. H. Kuchlein (Wageningen, Netherlands), A. and Z. Lastúvka (Brno, Czechoslovakia), R. Puplesis (Vil- nius, Lithuania) and H. W. van der Wolf (Nuenen, Netherlands). The author is particularly indebted to Roland Johansson, for the continuous stream of information and the many discussions about nepti- culid taxonomy, which amongst others led to the present paper. REFERENCES Johansson, R., 1971. Notes on the Nepticulidae (Lepidop- tera) I. A revision of the Nepticula ruficapitella group. — Entomologica scandinavica 2: 241-262. Johansson, R. & E. S. Nielsen, 1990. Tribus Nepticulini. - In: Johansson, R. et al. The Nepticulidae and Opos- tegidae (Lepidoptera) of NW Europe. — Fauna ento- mologica scandinavica 23: 111-238, pls. Klimesch, J., 1951. Zur Kenntnis der Genitalmorphologie einiger Nepticula-Arten (Lep., Nepticulidae). - Zeitschrift der Wiener Entomologischen Gesellschaft 36: 4-9. Klimesch, J., 1958. Beiträge zur Kenntnis der Lepidop- teren-Fauna der Wachau in Niederösterreich (Microle- pidoptera). - Zeitschrift der Wiener Entomologischen Gesellschaft 43: 17-22, 43-44, 76-77, 91-97. Klimesch, J., 1961. Ordnung Lepidoptera. I. Teil. Pyralid- ina, Tortricina, Tineina, Eriocraniina und Micropte- rygina. - In. H. Franz (ed.). Die Nordost-Alpen im 241 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 4-9. Stigmella rolandi, genitalia and leafmine. - 4, Capsule male genitalia, slide 2563 (Spain: Cadalso); 5, Valva, inner aspect, slide 2563; 6, Aedeagus, holotype, slide 2780; 7, Female terminal segments, dorso-lateral view, slide 2783 (Yugoslavia: Krk); 8, Bursa copulatrix, detail showing enlarged pectinations, slide 2783; 9, leaf-mine on Rosa sp. from type-locality, one of three mines from which holotype was reared. Scales: 0.1 mm (figs. 4-8), 5 mm (fig. 9); 4-6 and detail of 8 on same scale. 242 VAN NIEUKERKEN: Stigmella rolandi sp. n. Fig. 10. Distribution of Stigmella rolandi, mapped on 50 X 50 km UTM squares. Spiegel ihrer Landtierwelt, 2: 481-789. Universitätsverlag Wagner, Innsbruck. Klimesch, J., 1978. Beitrag zur Kenntnis der Nepticuli- denfauna von Anatolien und der Insel Rhodos (Lepi- doptera, Nepticulidae). — Tijdschrift voor Entomolo- gie 121: 239-278. Nieukerken, E. J. van, 1986. A provisional phylogenetic check-list of the western palaearctic Nepticulidae, with data on hostplants (Lepidoptera). - Entomologica scandinavica 17:1-27. Nieukerken, E. J. van, 1990. The Trifurcula subnitidella group (Lepidoptera: Nepticulidae): taxonomy, distri- bution and biology. — Tijdschrift voor Entomologie 133: 000-000. Szócs, J., 1955. A budapesti Mártonhegy lepke-faunája. — Folia entomologica Hungarica, s. n. 8: 157-171. Szócs, J., 1956. Magyarország Nepticulidái (Lepidopt.) (Die in Ungarn vorkommenden Nepticula-Arten (Le- pidopt.)) — Folia entomologica Hungarica, s. n. 9: 381- 394. Szöcs, J., 1963. A lepkehernyoók természetes tápnövényei. [Die natiirlichen Futterpflanzen der Schmetterlings- raupen.]- Folia entomologica Hungarica, s. n. 16: 83- 120. Szöcs, J., 1965. Microlepidoptera I, Nepticulidae. - Fauna Hungarica 76: 48-104. Szócs, J., 1973. Adatok a Bakony aknázómoly faunájához. Angaben zur Minierfliegen-fauna [sic!] des Bakony- Gebirges. - A Veszprém megyei múzeomok Közlemé- nyei 12: 451-455. Szócs, J., 1977. Adatok a Mátra-Hegység Aknázómoly- faunájához. (Data to the mining moth fauna found in the Mts. Matra.) - Folia Historia naturalis Musei Ma- trensis 4: 91-99. Szócs, J., 1978. Adatok a Pili-hegység Aknázómoly fauná- jához. (Data to the mining moths fauna from the Pilis Mts.) — Folia entomologica Hungarica, s. n. 31: 265- Di, Sz6cs, J., 1981. Angaben über minierenden Motten aus Budapest und Umgebung. - Folia entomologica Hun- garica, s. n. 42: 209-220. Waters, E. G. R., 1928. A new Nepticula from North Wales. - Entomologist's monthly Magazine 64: 105- 106. Zimmermann, F., 1944. Zur Kenntnis der Verbreitung der Nepticuliden in den Reichsgauen Wien und Niederdo- nau (Lepidopt.). - Zeitschrift der Wiener Entomolo- gischen Gesellschaft 29: 3-6, 60-64, 78-91, 107-122. Received: 18 October 1990 Accepted: 18 October 1990 243 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 244 TILA M. PEREZ! & WARREN T. ATYEO? Laboratorio de Acarologia, Universidad Nacional Autonoma de México “Department of Entomology, University of Georgia NEW TAXA OF FEATHER MITES (ACARINA, PTEROLICHIDAE) FROM MEGAPODES (AVES, MEGAPODIIDAE) Pérez, T. M. & W. T. Atyeo, 1990. New taxa of feather mites (Acarina, Pterolichidae) from Megapodes (Aves, Megapodiidae). - Tijdschrift voor Entomologie 133: 245-249, figs. 1-10, table 1. [ISSN 0040-7496]. Published 14 December 1990. Ascetolichus gen. n. (Pterolichoidea, Pterolichidae) is established for the type species Pterolichus (Pseudalloptes) palmiger Trouessart and two new species, A. microthrix and A. ruidus. All hosts are Megapodiidae. Correspondence: W. T. Atyeo, Department of Entomology, University of Georgia, Athens, Georgia 30602, U.S.A. Key words. — Feather mites, Pterolichidae, Ascetolichus, Megapodiidae parasites. In the late 1800s and early 1900s E. L. Trouessart, often in collaboration with other authors, named many taxa of feather mites taken primarily from study skins in ornithological collections in Paris and Angers. His collections included mites from three species of Megapodiidae: ‘… les Talegalles (Talegallus Cuvieri et Aepipodius Bruijnini) de Nouvelle-Guinée’ and ‘... le Mégapode de Jobi (Megapodius Jobiensis) de île Jobi (Nouvelle-Gui- née). From these hosts, Trouessart (1887) and Trouessart & Neumann (1888) described seven mite species, however, the location of most of the types is unknown. By examining megapode study skins we have recovered the seven species of Trouessart plus many new species. From this rich and diverse acarofauna, we will establish herein a new genus for one of seven Trouessart species and two new species. MATERIALS AND METHODS We have examined 54 collections of feather mites from 95 museum study skins of megapodes at the American Museum of Natural History (see Atyeo & Braasch 1966 for collecting technique). These mites are from 13 of the 19 megapode species as recognized by White & Bruce (1986). Each collec- tion contains all mite specimens taken from an individual skin, but the specimens may include only a portion of the species known from the host spe- cies (see Méjia-Gonzalez & Pérez 1988 for limita- tions of the collecting technique). The megapode species examined followed by the number of sam- ples taken and the number of skins examined are given in table 1. Names denoted by asterisks are hosts from which species of the new genus were collected. All taxa being described, regardless of the sample size, are believed to be from the Megapodiidae as we have never encountered similar forms in any of our extensive collections from the birds of the world. Signatures for idiosomal setae follow the Grand- jean system as modified by Griffiths et al. (1990). Table 1. The number of feather mite collections (left number) obtained from museum study skins examined (right number) of megapodes. Asterisks indicate hosts of Ascetolichus species. Megapodius freycinet Gaimard, 1823 11 of 15 M. eremita Hartlaub, 1867 3 of 4 M. affinis A. B. Meyer, 1874 (= decollatus Oustalet, 1878) 1 of 3 (= jobiensis Oustalet, 1881) 3 of 5 M. laperouse Gaimard, 1823 9 of 14 M. pritchardii G. KR. Gray, 1864 0 of 10 M. (= Eulipoa) wallacei G. R. Gray, 1860 5 of 7 Leipoa ocellata ocellata Gould, 1840 1 of 1 L. ocellata rosinae Mathews, 1912 0 of 2 Alectura lathami J. E. Gray, 1831* 2 of 11 Talegalla cuvieri Lesson, 1828 3 of 3 T. jobiensis jobiensis A. E. Meyer, 1874* 4 of 5 T. fuscirostris Salvadori, 1877 4 of 5 Aepypodius arfakianus (Salvadori, 1877)* 3 of 4 A. bruijnii (Oustalet, 1880)* 2 of 2 Macrocephalon maleo S. Müller, 1846 3 of 4 245 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 1-2. Ascetolichus palmiger (Trouessart). Ventral and dorsal aspects of male. Setal signatures follow Griffiths et al. (1990). N 200 um Figs. 3-4. Ascetolichus palmiger (Trouessart). Ventral and dorsal aspects of female. Setal signatures follow Griffichs et al. (1990). 246 Measurements, in micrometres, are the mean + standard error (when N>10), followed in paren- theses by the observed limits and number of obser- vations. The SEM micrographs were taken with a Philips 505 using dehydrated specimens collected from museum study skins. Holotypes of new species will be deposited in the American Museum of Natural History. Abbrevia- tions for other type repositories and accession numbers of bird and mite collections are: AMNH, American Museum of Natural History, New York; TRT, Trouessart Collection, Paris; and UGA, Uni- versity of Georgia, Athens. Ascetolichus Pérez & Atyeo, gen. n. Diagnosis. - Small pterolichine mites with quad- rate idiosoma and gnathosoma; well-developed dorsal shields; posterolateral angles of propodo- soma extended into points; all idiosomal and leg setae present; setae vi expanded, minutely branched; setae c2, cp large, branched; setae c3 minute, ventrolateral; cupules ia conspicuous; epi- merites I free; legs subequal; legs III, IV ventral; pretarsi symmetrical, dentate; tarsi I and II shorter than corresponding tibiae; genua I, II, tibiae I, II with dorsal crests or pebbled surfaces; bases of sole- nidia 0, p respectively elevated above tibiae, tarsi; genua I, II with setae mG bi- or trifurcate. Male idiosoma with shallow terminal cleft on which is inserted leaflike psl and branched hl; setae e2 ovate, fringed; adanal discs small, about 8 in diame- ter, edentate; tarsus IV with subterminal claw. Fe- male with idiosoma parallel-sided, more or less truncated posteriorly; external spermduct present; various terminal setae elaborated (as in fig. 4); oviporus and associated structures anterior of seju- gal furrow. Type species. - Pterolichus (Pseudalloptes) pal- miger Trouessatt. Etymology. - Contraction of asketos (Gr., cur- iously wrought, ornament) and Pterolichus, mascu- line. Ascetolichus palmiger (Trouessart) comb. n. (figs. 1-6, 8) Pterolichus (Pseudalloptes) palmiger Trouessart, 1887: 116-7; Canestrini & Kramer 1899: 61. Males. — Dorsal shields with small pits unequally distributed; genua I, II with ridges parallel to axis; genua I, II with setae mG, cG coarsely branched; setae cp long, bifurcate, unequal branches; measure- ments: length, including gnathosoma 279 + 2 (262- 290, 19), width 170 = 1 (162-177, 19), sce:sce 86.5 + 0.8 (78.4-92.1, 19), c2 length 46.2 + 0.7 (39.2- 49.0, 17), cp length 58.4 + 0.7 (50.9-63.7, 19), hl length 8.3 + 0.2 (7.8-9.8, 18), ps1 19.6 + 0.3 (17.6- PEREZ & ATYEO: Feather mites of Megapodes 21.6, 18), gnathosomal length 45.2 + 0.4 (43.1-49.0, 19), gnathosomal width 46.9 + 0.3 (45.1-49.0, 17). Females. - Proterosoma as in male; dorsal hys- terosoma with small pits, especially along margins; measurements: length, including gnathosoma 435 (424-463, 7), external spermduct about 13.7; width 230 (220-254, 7), sce:sce 120.4 (113.7-129.4, 7), c2 length 59.1 (54.9-66.6, 6), c3 length 87.1 (82.3-92.1, 6), e2 length 51.9 (50.9-54.9, 6), f2 length 22.7 (19.6-23.5, 7), psl 14.8 (11.8-17.6, 7), ps2 23.3 (19.6-27.4, 5), gnathosomal length 65.8 (64.7-68.7, 7), gnathosomal width 69.2 (62.7-76.4, 7). Type data. - From Aepypodius bruns: New Guinea, | Waigeu Island], lectotype 6, paralectotype Q (TRT slide 37 A 1). i Material examined. — The type series (see Re- marks) and other specimens from Aepypodius bruijnü: Indonesia: West Irian: Waigeu Island, 15 6, 4 9, circa 1894, A. A. Bruijn (AMNH 539411, UGA 5339; AMNH 539410, UGA 5340). From A. arfakianus; Indonesia: West Irian: Idenburg River, 6 km SW Bernhard Camp (39° 15’E, 3°30'S), 1 6, 3 9, 21 February 1939, R. Archbold (AMNH 338429, UGA 5336); Snow Mountains, 3 @ , 1 9, 24 September 1910, A. S. Meek (AMNH 539393, UGA 5337). From Talegallus cuvieri: New Guinea: no other data, 2 Q (TRT slides nos. 37 A 2, 22 H 16). Remarks. - The number of observations for setae and gnathosomal measurements are small as many setae are lacking from the specimens and many gnathosomata have been deformed during speci- men preparation. The measurements for the gnath- osoma are included in this description only; they are similar for the other species for which there are few specimens. Trouessart (1887) stated that the hosts for this species were T. cuvieri and A. bruijnii, New Guinea. Our collections from A. bruijnii are conspecific with Trouessart’s specimens from the listed hosts (see above). We assume that Trouessart’s two females from T. cuvieri represent accidental associations. Ascetolichus species (fig. 7) One of the males used for SEM micrographs is an undescribed species (fig. 7) from the study skin of Aepypodius arfakianus from the Bernhard Camp (listed above). All males from the Aepypodius col- lections have been mounted and additional speci- mens were not found. Although the host associa- tion must be tentative until additional specimens are collected, it is possible that two species of Asce- tolichus could occur on A. arfakianus. Two or three congeners on a single host, with each congener occuping a different microhabitat has been well documented (Atyeo & Pérez 1988, Méjia-González & Pérez 1988). The species is related to Ascetolichus palmiger as 247 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 sog Figs. 5-8. SEMs of dorsal aspects of Ascetolichus species. — 5, Ascetolichus palmiger, male: arrow = setae e2; 6, A. palmiger, male propodosoma: upper arrow = setae cG, lower arrow = seta mG, genual ridges between setae; 7, Ascetolichus species, male: arrow = setae psl; 8, A. palmiger, female. Scale bars: 5, 7,8 = 100 um; 6 = 50 um. 200 um (3) 2 oe 099 Dos 09 & gue 060% — O 5 06 Ooo 0) O) OL HOR > 000000 AO ar Q oO oc Ô 3700 20 S 0 (è) 900 9990 ER ) 6) Oo do lo) ie) So © 0° È ANS Toy Figs. 9-10. Dorsal aspects of Ascetolichus species. - 9, A. rwidus sp. n., female; 10, A. microthrix sp. n., male. G = position of male genital organ. 248 shown by the ridges on the anterior legs, but the dorsal ornamentation is different, setae c2 and cp are smaller, and setae psl are bifurcated. Ascetolichus ruidus Pérez & Atyeo, sp. n. (fig. 9) Female (holotype). - Dorsal idiosoma with large, deep pits; genua I, II, femur II with antaxial surfaces pebbled; tibiae I, II with setae mG, cG thin, bi- or trifurcate; setae cp long, bifurcate, unequal branches; measurements: length, including gnath- osoma 424, width 197, external spermduct 43.1, sce:sce 11.7, c2 66:6, cp 84.3, e2 3912, f2 19.6, ps] DB 1982 197% Male. - Unknown. Type data. - The holotype female was collected from Alectura lathami: Australia: Queensland: (?) Barron River, 19 July 1884, G. M. Mathews (AMNH 539322, UGA 5324). The holotype is dep- osited in AMNH. Etymology. - From rwidus (Gr., rough) to refer to the rugose tegument of legs I and II. Remarks - We have only two collections from A. lathami and in one, the unique holotype was found. Until validated by additional material, the host as- sociation is considered tentative. Ascetolichus microthrix Pérez and Atyeo, sp. n. (fig. 10) Males (N=2). - Dorsal idiosoma with large pits; legs I, II without pebbled tegument or ridges paral- lel to axis; genua I, II with setae mG simple or with small basal branch; setae cp bifurcate, with few branches; measurements: length, including gnath- osoma 280 (278-282), width 164 (162-166), sce:sce 95.1 (94.1-96.0), c2 length 29.4 (27.4-31.4), cp length 53.9 (49.0-58.8), hl length 11.8 (11.8), psl 27.9 (27.4-28.4). Female (N=1). - Proterosoma and dorsal idio- soma similar to male; measurements: length, in- cluding gnathosoma 416, width 208, sce:sce 113.7, external spermduct 29.4, c2 31.4, cp 74.5, e2 23.5, PAG ps 137, \ps2 bles: Type data. - From Talegalla j. jobiensis: Indone- sia: West Irian: Humboldt Bay, Hollandia, holotype 6, 1 ©, 1 Q paratypes, 5 August 1928, E. Mayr (AMNH 539377, UGA 5332). The holotype is dep- osited in AMNH, paratypes in UGA. PEREZ & ATYEO: Feather mites of Megapodes Etymology. - From mikros (Gr., small) and thrix (Gr. seta) to refer to the small setae c2. Remarks. - The two males in this collection had setae e2 missing; they have been represented with dotted lines as structures similar to those of A. palmiger. The females of the two new species, A. rwidus and A. microthrix differ from those of A. palmiger by having long external spermducts. The female of A. ruidus differs from A. microthrix in having the pits of the hysterosomal shield approximately equal in size, a longer spermduct, and longer setae (espe- cially c2, e2, ps1, compare measurements). ACKNOWLEDGEMENT This research was supported by the National Science Foundation (BSR 89-08301). REFERENCES Atyeo, W. T. & N. L. Braasch, 1966. The feather mite genus Proctophyllodes (Sarcoptiformes: Proctophyllo- didae). - Bulletin of the University of Nebraska State Museum 5: 1-354. Atyeo, W. T. & T. M. Pérez, 1988. Species in the genus Rhytidelasma Gaud (Acarina: Pterolichidae) from the Green Conure, Aratinga holochlora (Sclater) (Aves: Psittacidae). — Systematic Parasitology 11: 85-96. Canestrini, G. & P. Kramer, 1899. Demodicidae und Sar- coptidae. - Das Tierreich 7: 1-193. Griffiths, D. A., W. T. Atyeo, R. A. Norton & C. A. Lynch, 1990. The idiosomal chaetotaxy of astigmatid mites. — Journal of Zoology, London 220: 1-32. Méjia-González, E. & T. M. Pérez, 1988. Three new spe- cies of Fainalges Gaud and Berla (Analgoidea: Xolal- gidae) with descriptions of their developmental series. — Acarologia 29: 73-86. Trouessart, E. L. (1886) 1887. Diagnoses d'espèces nou- velles de Sarcoptides plumicoles (Analgesinae). — Bul- letin de la Société d'études scientifiques d'Angers 15: 85-156. Trouessart, E. L. & G. Neumann, 1888. Diagnoses d’espe- ces nouvelles de Sarcoptides plumicoles (Analgesinae). - Bulletin scientifique de la France et de la Belgique 19: 325-380. White, C. M. N. & M. D. Bruce, 1986. The birds of Wal- lacea. - British Ornithologists’ Union Check-list No. 7, 524 p. Received: 1 May 1990 Revised version accepted: 30 July 1990 LS) Se Ne) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 250 D. L. J. QUICKE! & C. van ACHTERBERG? "Department of Animal and Plant Sciences, University of Sheffield *Nationaal Natuurbistorisch Museum, Leiden THE TYPE SPECIMENS OF ENDERLEIN'S BRACONINAE (HYMENOPTERA: BRACONIDAE) HOUSED IN WARSAW Quicke, D. L. J. & C. van Achterberg, 1990. The type specimens of Enderlein's Braconinae (Hymenoptera: Braconidae) housed in Warsaw. — Tijdschrift voor Entomologie 133: 251- 264. [ISSN 0040-7496]. Published 14 December 1990. The type specimens of 147 species of Braconinae described by Enderlein in the collection of the Polish Academy of Science, Institute of Zoology, Warsaw have been examined. Eighty- one new combinations are reported, the systematic positions of 39 species are confirmed and lectotypes are designated for 62 species. The genera Diolcia Enderlein and Monolcia Enderlein are synonymized with Campyloneurus Szépligeti; Diamblomerina Enderlein is synonymized with Nedinoschiza Cameron; Diamblomera Enderlein is removed from syn- onymy with Odontoscapus Kriechbaumer and provisionally treated as a valid genus. Cya- nopterus solox Enderlein is treated as a new junior synonym of Exurobracon triplagiata Cameron. Correspondence: D. L. J. Quicke, Department of Animal and Plant Sciences, University of Sheffield, Sheffield, U.K. Key words. - Hymenoptera, Braconidae, Braconinae, Enderlein, reclassification, lectotypes. Günter Enderlein (1905a, 1905b, 1920) described a large number of new taxa of Braconidae including 153 new species and 17 new genera of Braconinae, the vast majority being dealt with in the last of his papers. The primary types of 147 taxa are listed in this paper because they are deposited in the Instytut Zoologii of the Polish Academy of Sciences in War- saw (PAN). Enderlein worked at the Museum fiir Naturkunde, Stettin (now Szczecin in north-west Poland) between the years 1906 and 1919. The principally tropical Braconidae dealt with in his papers were mainly from the collection of the former curator of the Stettin Museum, Dr. H. Dohrn and most of them were collected in Sumatra by Mr. M. Ude. The specimens in the collection formerly belonging to the Museum fiir Naturkunde in Berlin are duplicates retained by Enderlein. As a consequence of the Second World War, the collec- tion finally ended up in the Polish Academy of Sciences. It is now housed at a field station/ museum belonging to the PAN at Lomna, approximately 30 km north-west of Warsaw, and the main part of the material is in very good condition. Although it had been possible to study some of this material in the past by means of small postal loans, damage that accrued to a considerable part of such loaned material due to bad postal handling has meant that Dr Eusgeniusz Kierych, the curator of entomology at the PAN, was reluctant to continue such loans. Thus in order to examine the entirity of the Enderlein collection so that the identities of many of his species could be re-assessed, the authors have recently visited the PAN facility at Lomna and the findings are reported below. Enderlein’s species concept seems to have been reasonable, although he described the same species on several occasions twice in the same paper; only rarely are his type series heterogeneous. However, his generic concept was, judging from his work on the Braconinae, very poor indeed and he would often describe two new genera based on one or two closely related species. For example, his genera An- tidiolcus and Plagiozina (both of which have now been synonymized with Nesaulax Roman; Quicke 1984a) are based on what are in fact two sibling species. Similarly, his genera Diolcia and Monolcia are based on another extremely closely related pair of species, both of which are transferred below to Campyloneurus Szépligeti. On other occasions he split the members of a single genus between as many as four genera, for example, species of Hemz- bracon Szépligeti were described under Ipobracon Thomson, Antiolcus Enderlein and Udamolx En- derlein as well as in Hemibracon itself. Enderlein’s inconsistencies and the errors in the generic level classification of his braconine species 251] TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 1. Overview of the current taxonomic status of the Braconinae described by G. Enderlein and deposited in PAN. Current genus Antiolcia Enderlein Archibracon Saussure Bathyaulax Szépligeti Blastomorpha Szépligeti Campyloneurus Szépligeti Chaoilta Cameron Chelonogastra Ashmead Compsobracon Ashmead Cordibracon van Achterb. Craspedolcus Enderlein Cratobracon Szépligeti Curriea Ashmead Diamblomera Enderlein Digonogastra Viereck Species mitelligera Enderlein, 1920: 116-117 atricauda Enderlein, 1920: 97 gutta Enderlein, 1920: 97-98 flavipera Enderlein, 1920: 126-127 gutta Enderlein, 1920: 122-123 laeviventris Enderlein, 1920: 126 latiangulata Enderlein, 1920: 125-126 nigriceps Enderlein, 1920: 125 pectinatus Enderlein, 1920: 123 rugiventris Enderlein, 1920: 124-125 celebensis Enderlein, 1920: 55-56 angulosus Enderlein, 1920: 115 basalis Enderlein, 1920: 105-106 bicarinatus Enderlein, 1920: 112 cingulicauda Enderlein, 1920: 105 flavicosta Enderlein, 1920: 115-116 impressimargo Enderlein, 1920: 109 latispeculum Enderlein, 1920: 103 limbaticauda Enderlein, 1920: 112 marginiventris Enderlein, 1920: 109- 110 nigricosta Enderlein, 1920: 108 punctativentris Enderlein, 1920: 110 reticulatus Enderlein, 1920: 107 serenans Enderlein, 1920: 104-105 speculiger Enderlein, 1920: 109 tibialis Enderlein, 1920: 108 tricarinatus Enderlein, 1920: 113 trispeculatus Enderlein, 1920: 107-108 carinicornis Enderlein, 1920: 55 compta Enderlein, 1920: 56 disciventris Enderlein, 1920: 103 thodeanus Enderlein, 1920: 57 laqueatus Enderlein, 1920: 120 fraternus Enderlein, 1920: 92 maculicosta Enderlein, 1920: 71-72 obscuriventris Enderlein, 1920: 93 trisulcatus Enderlein, 1920: 92 gibber Enderlein, 1920: 65-66 grata Enderlein, 1920: 117 gratiosus Enderlein, 1920: 53 guttifer Enderlein, 1920: 53 acuticellus Enderlein, 1920: 100-101 grisescens Enderlein, 1920: 101 acuticampa Enderlein, 1920: 86 alboniger Enderlein, 1920: 78 aterimma Enderlein, 1920: 98 aureopilosa Enderlein, 1920: 80 biareata Enderlein, 1920: 92 bicristata Enderlein, 1920: 88 bicuneata Enderlein, 1920: 87-88 bicunea Enderlein, 1920: 83-84 brevicapula Enderlein, 1920: 91 brevicunea Enderlein, 1920: 78-79 camerom Enderlein, 1920: 82 chilensis Enderlein, 1920: 58 columbiana Enderlein, 1920: 82 crenulata Enderlein, 1920: 79 digitata Enderlein, 1920: 84 duploareata Enderlein, 1920: 82-83 ecuadorensis Enderlein, 1920: 85 fenestrata Enderlein, 1920: 89 252 Original genus Antiolcia Udamolx Udamolx Euryacria Gontobracon Bathyaulax Bathyaulax Bathyaulax Gontobracon Bathyaulax Platybracon Monolcia Campyloneurus Diolcia Campyloneurus Monolcia Campyloneurus Campyloneurus Diolcia Campyloneurus Campyloneurus Campyloneurus Campyloneurus Campyloneurus Campyloneurus Campyloneurus Monolcia Campyloneurus Platybracon Platybracon Campyloneurus Cyanopterus Udamolcia Craspedolcus Ipobracon Craspedolcus Craspedolcus Merinotus Antiolcia Aphrastobracon Aphrastobracon Diamblomera Diamblomera Ipobracon Ipobracon Udamolx Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Cyanopterus Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Type-locality Cameroons: Barombi S. Africa: Natal S. Africa: Natal Malagasy: Amber Mountains S. Africa: Natal Malagasy: Amber Mountains Malagasy: Tamatave S. Africa: Transvaal: Zout- pansberg S. Africa: Transvaal: Zout- pansberg Tanzania Indonesia: N. Sulawesi Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Taiwan: Taihorin Indonesia: Java: Sukabumi Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Java: Sukabumi Sukaranda Sukaranda Sukaranda Sumatra: Sumatra: Sumatra: Indonesia: Indonesia: Indonesia: Ceylon Indonesia: Indonesia: Indonesia: Indonesia: Indonesia: Indonesia: Indonesia: Sumatra: Liangagas Indonesia: Sumatra: Sukaranda Brazil: Santa Catharina Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Java: Sukabumi Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Tanzania: Nyembe Bulungwa Bioka (= Fernando Poo) Cameroons: Barombi Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Colombia: Rio Magdalena Colombia Brazil Brazil: Obodos Peru Colombia Brazil: Obidos Ecuador: Curaray Guyana: Demerara Peru: Chanchamayo: Rio Toro Mexico: Chiapas Chile Colombia Brazil (? Guyana): Demerara Brazil: Santa Catharina Brazil: Santa Catharina Ecuador: Balzapampa Brazil: Pebas Sukaranda Sukaranda Sukaranda Sukaranda Sukaranda Sukaranda Sumatra: Sumatra: Sumatra: Sumatra: Sumatra: Sumatra: Table 1. (continued). Current genus Ectemnoplax Enderlein Euwrobracon Ashmead Gammabracon Quicke Gronaulax Cameron Hemibracon Szépligeti Hybogaster Szépligeti Iphiaulax Foerster Ischnobracon Baltazar Isomecus Kriechbaumer Megabracon Szépligeti Monocoila Roman Nedinoschiza Cameron Nesaulax Roman Pachybracon Cameron, -group !) Physaraia Shenefelt Plaxopsis van Achterberg Plesiobracon Cameron Rbadinobracon Szépligeti Rhamnura Enderlein Serraulax Quicke Shelfordia Cameron Spathulibracon Quicke Stenobracon Szépligeti Syntomernus Enderlein Trigastrotheca Cameron Trispinaria Quicke QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae Species flavicaligata Enderlein, 1920: 80 latecrenulata Enderlein, 1920: 81-82 latefasciata Enderlein, 1920: 90 laticampa Enderlein, 1920: 90-91 laticunea Enderlein, 1920: 79 longicapula Enderlein, 1920: 91 macella Enderlein, 1920: 86 mediofusca Enderlein, 1920: 88-89 nigripecta Enderlein, 1920: 83 obtusicampa Enderlein, 1920: 85-86 ochripes Enderlein, 1920: 86-87 penniseta Enderlein, 1920: 87 rectivena Enderlein, 1920: 81 sigillata Enderlein, 1920: 79 speculata Enderlein, 1920: 89-90 subfracta Enderlein, 1920: 96-97 tuberculata Enderlein, 1920: 84-85 peruliventris Enderlein, 1920: 110-111 merinotoides Enderlein, 1920: 128-129 solox Enderlein, 1920: 58 nigriseta Enderlein, 1920: 68-69 laticampus Enderlein, 1920: 69 apicalis Enderlein, 1920: 96 catharinensis Enderlein, 1920: 98-99 crassecrenulatus Enderlein, 1920: 81 nigripalpalis Enderlein, 1920: 90 politus Enderlein, 1920: 99 vaucristatus Enderlein, 1920: 99-100 w-impressus Enderlein, 1920: 63 crista Enderlein, 1920: 75-76 breviseta Enderlein, 1920: 60 sautert Enderlein, 1920: 128 udet Enderlein, 1920: 127-128 rhyssides Enderlein, 1920: 61-62 quadrirugulosus Enderlein, 1920: 94 filiseta Enderlein, 1920: 57-58 lurida Enderlein, 1920: 111 speciosus Enderlein, 1920: 121-122 excisus Enderlein, 1920: 95-96 gracilis Enderlein, 1920: 67-68 pravivena Enderlein, 1920: 94-95 albescens Enderlein, 1920: 117-118 albicans Enderlein, 1920: 102-103 apicalis Enderlein, 1920: 114 gibbiventris Enderlein, 1920: 104 insolita Enderlein, 1920: 118 sumatrana Enderlein, 1905b: 236 magnificus Enderlein, 1920: 77-78 cincticauda Enderlein, 1920: 104 ruficauda Enderlein, 1920: 71 capillicauda Enderlein, 1905a: 196 filicauda Enderlein, 1905a: 195 denticornis Enderlein, 1920: 123 bispeculum Enderlein, 1920: 69 criniseta Enderlein, 1920: 68 flagriseta Enderlein, 1920: 64-65 ingentiseta Enderlein, 1920: 70-71 quadricarinata Enderlein, 1920: 66-67 rimicunea Enderlein, 1920: 67 rugiventris Enderlein, 1920: 77 elegantulus Enderlein, 1920: 62-63 pusillus Enderlein, 1920: 121 tridentata Enderlein, 1920: 60-61 sannio Enderlein, 1920: 54 Original genus Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Ipobracon Antiolcus Ipobracon Ectemnoplax Bracon Cyanopterus Merinotus Merinotus Antiolcus Udamolx Ipobracon Ipobracon Udamolx Udamolx Hemibracon Ipobracon Megagonia Iphiaulax Iphiaulax Rhadinobracon Craspedolcus Cyanopterus Ectemnoplax Diamblomerina Antidiolcus Merinotus Plagiozina Udamolcia Campyloneurus Monolcia Campyloneurus Udamolcia Gastrotheca Ipobracon Campyloneurus Meronotus Rhamnura Rhamnura Goniobracon Merinotus Merinotus Merinotus Merinotus Merinotus Merinotus Ipobracon Hemibracon Syntomernus Odontopygia Pseudospinaria Type-locality Guyana: Demarara Brazil: Santa Catharina Brazil: Obidos Guyana: Demerara Peru: Chanchamayo: Rio Toro Guyana: Demerara Peru: Chanchamayo Brazil: Obidos Argentine: Mendoza Colombia Ecuador: Guayaquil Brazil: Obidos Brazil: Santa Catharina Peru Colombia: Rio Magdalena Brazil: Obidos Peru: Chanchamayo: Rio Toro Taiwan: Takao Indonesia: Sumatra: Sukaranda India: Sikkim Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Costa Rica Brazil: Santa Catharina Brazil: Santa Catharina Ecuador: Balzapamba Peru: Chanchamayo Guyana: Demerara Brazil: Sao Paulo de Olivenca Indonesia: Sumatra: Sukaranda Indonesia: Sumba Taiwan: Takao Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Ecuador: Bucay Peru: Chanchamayo: Rio Toro S. Africa Indonesia: Sumatra: Indonesia: Sumatra: Indonesia: Sumatra: Indonesia: Sumatra: Indonesia: Sumatra: Indonesia: Sumatra: Indonesia: Sumatra: India Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Tanzania: Usambara: Bulwa Indonesia: Sumatra: Liangagas Namibia Togo: Bismarckburg Cameroons: Joh.-Albrechtshöhe Cameroons: Kribi Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Indonesia: Java: Sukabumi Indonesia: Sumatra: Sukaranda Taiwan: Takao Indonesia: Sumatra: Sukaranda Indonesia: Sumatra: Sukaranda Sukaranda Sukaranda Sukaranda Sukaranda Sukaranda Sukaranda Sukaranda 255 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Table 1. (continued). Current genus Undabracon Quicke Zaglyptogastra Ashmead Species jucundus Enderlein, 1920: 119 griseiseta Enderlein, 1920: 59 helvimacula Enderlein, 1920: 123-124 plumiseta Enderlein, 1920: 59-60 Original genus Type-locality Udamolcia Indonesia: Sumatra: Sukaranda Megagonia Indonesia: Sumatra: Sukaranda Goniobracon Ethiopia: Eritrea: Asmara Megagonia Indonesia: Sumatra: Sukaranda Unplaced species (these taxa will be treated in a forthcoming paper by the junior author) albimanus Enderlein, 1920: 118-119 angustisulca Enderlein, 1920: 73-74 basispeculum Enderlein, 1920: 73 consimilis Enderlein, 1920: 113-114 denticornis Enderlein, 1920: 120-121 flavimarginatus Enderlein, 1920: 114 gracilis Enderlein, 1920: 118 laevibasis Enderlein, 1920: 75 laeviventris Enderlein, 1920: 76-77 latisulca Enderlein, 1920: 74 maculistigma Enderlein, 1920: 107 parvispeculum Enderlein, 1920: 73 serenimanus Enderlein, 1920: 114-115 sexrugosus Enderlein, 1920: 72-73 strigidorsum Enderlein, 1920: 93 trirugosus Enderlein, 1920: 72 trispeculum Enderlein, 1920: 70 undicuneus Enderlein, 1920: 106-107 Udamolcia Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Monolcia Indonesia: Sumatra: Sukaranda Udamolcia Indonesia: Sumatra: Sukaranda Monolcia Indonesia: Java: Sukabumi Udamolcia Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Campyloneurus Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Monolcia Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Craspedolcus Indonesia: Sumatra: Sukaranda Ipobracon Indonesia: Sumatra: Sukaranda Merinotus Indonesia: Sumatra: Sukaranda Campyloneurus Indonesia: Sumatra: Sukaranda 1. Including Udamolcia Enderlein, 1920; the group is currently being revised by the junior author. are almost entirely attributable to his giving an inordinate amount of weight to the number of metasomal tergites that have transverse sub-poste- rior grooves. Only rarely does this character have any significance at the genus level in the Braconi- nae, but Enderlein based several of his new genera solely on its distribution. It is perhaps not too sur- prising therefore that only a few of his genera have stood the test of time. ANNOTATED CATALOGUE The following is a list of all the species of Bra- coninae described by Enderlein (1905a, 1905b, 1920) whose types are in the PAN collection. Nearly all of these species were described in Ender- lein's 1920 paper, however, the five species of Rhamnura Enderlein were described in Enderlein (1905a) and one species of Gastrotheca Guérin- Méneville (not Fitzinger) (= Physaraia Shenefelt) were described in Enderlein (1905b). The type specimens of all of these are in the PAN except for those of three of the species of Rhamnura and paralectotypes of the remaining two, which are in the Berlin Museum. Enderlein’s original publications generally give considerable detail concerning the type series and the type localities. Where the type series includes members of only one sex he usually placed a "Typus’ label on one individual and ‘Co-Typus’ labels on the 254 remainder thus giving a clear indication of which specimen he wished the species concept to be based upon; in nearly all such cases we have selected his specimen bearing the ‘Type’ label as lectotype. However, where the type series contains both males and females he usually placed a ‘Type’ label on one female and on one male; in these cases we have generally selected the female as the lectotype as this is the sex upon which most species descrip- tions in the Braconidae are based. Because of Ender- lein’s detailed notes on the type material, only a summary is provided below; the type locality of the primary types is given in table 1. It should be noted that Shenefelt (1978) also gives some type locality details corresponding to that of the holotype or those specimens bearing ‘Type’ labels. The species are listed below under their original generic combinations in alphabetical order. Where appropriate, notes are provided on the conditions of the specimens, homogeneity of the type series and previous taxonomic changes. The correct ge- neric placement of most of the species studied is given after the heading ‘Identity’; however, for some it has not been possible to make a definitive statement about their generic placement and in these cases a few notes are usually given indicating possible affinities. Their placement will be dis- cussed in a forthcoming paper on the Indo-Austral- ian genera of Braconinae by the junior author. Antidiolcus excisus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type not found, but should be present according to the original description. Notes. - This is the type species of Antidiolcus Enderlein which was synonymized with Nesaulax Roman by Quicke (1984a). Identity. - Nesaulax excisus: Quicke (1984a). Antiolcia grata Type material. - Female holotype. Notes. - On Enderlein’s type label this is labelled as ‘Antiolcus gratus. Identity. - Curriea grata comb. nov. Antiolcia mitelligera Type material. - One female with ‘Type’ label hereby designated as lectotype and so labelled; one male paralectotype also with ‘Type’ label. Notes. — This is the type species of Antiolcia Enderlein which was synonymized with Odontosca- pus Kriechbaumer by Quicke (1981). Because this taxon is aberrant in Odontoscapus it seems best to retain it in Antiolcia till its position will be dis- cussed in the forthcoming paper by the junior au- thor. Both specimens are rather badly eaten-out but the male is more badly damaged than the female. The specimens are labelled by Enderlein as ‘Antiolcus mitelliger’. Identity. - Anziolcia mitelligera: Quicke (1981). Antiolcus apicalis Type material. - Female holotype. Notes. - This is the type species of Antiolcus Enderlein which was synonymized with Hemz- bracon Szépligeti by Quicke (1989c). Identity. - Hemibracon apicalis: Quicke (1989c). Antiolcus subfractus Type material. - Female holotype. Identity. - Digonogastra subfracta comb. nov. Aphrastobracon gratiosus Type material. - Male holotype. Notes. - Placed by Enderlein in the Aphrastobra- coninae which is now considered to be a tribe of Braconinae. Identity. - Curriea gratiosa comb. nov. Aphrastobracon guttifer Type material. - Male holotype. Notes. - See A. gratiosus (above). Identity. - Curriea guttifer comb. nov. Bathyaulax laeviventris Type material. - Female holotype. Identity. - Bathyaulax laeviventris. QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae Bathyaulax latiangulata Type material. - Female holotype. Notes. - The generic identity of this species is hereby confirmed but no opinion is being given concerning the validity or otherwise of the specific synonymy proposed by Granger (1949). Identity. — Bathyaulax foveiventris (Roman) (synonymized by Granger, 1949). Bathyaulax nigriceps Type material. - Male holotype. Identity. - Bathyaulax nigriceps. Bathyaulax rugiventris Type material. - Male holotype. Identity. - Bathyaulax rugiventris. Bracon merinotoides Type material. - Female holotype. Notes. - This species was transferred to Euwro- bracon Ashmead by Fahringer (1927) and subse- quently synonymized with E. forticornis (Cameron) by Quicke (1989b). Identity. - Euwrobracon forticornis (Cameron). Campyloneurus albicans Type material. - One female with "Typus’ label hereby designated as lectotype; 1 ‘male’ paralecto- type with ‘Typus’ label; 2 female and 2 male para- lectotypes with ‘Co-Typus’ labels. Notes. - The specimen labelled by Enderlein as a male and bearing a ‘Typus’ label is in fact a female. Identity. - Belongs to the Pachybracon Cameron- group. Campyloneurus basalis Enderlein (not Szépligeti) Type material. - One female with ‘Typus’ label hereby designated as lectotype; 1 male paralecto- type with ‘Typus’ label; 11 female paralectotypes with ‘Co-Typus’ labels. Identity. - Campyloneurus enderleini Fahringer, 1931; replacement name. Campyloneurus cincticauda Type material. - Female holotype. Identity. — Plesiobracon cincticauda comb. nov. Campyloneurus cingulicauda Type material. - Female holotype. Identity. - Campyloneurus cingulicauda. Campyloneurus disciventris Type material. - One female with ‘Typus label hereby designated as lectotype; 1 male paralecto- type with ‘Typus’ label; 2 female and 6 male para- lectotypes with ‘Co-Typus’ labels. Identity. — Chelonogastra disciventris comb. nov. 255 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Campyloneurus gibbiventris Type material. - Female holotype. Identity. - Belongs to the Pachybracon Cameron- group. Campyloneurus impressimargo Type material. - One female with "Typus’ label hereby designated as lectotype; 1 female paralecto- type ‘Co-Typus' label. Identity. - Campyloneurus impressimargo. Campyloneurus latispeculum Type material. - Female holotype. Identity. - Campyloneurus latispeculum. Campyloneurus maculistigma Type material. - One female with “Typus’ label hereby designated as lectotype; 1 female paralecto- type ‘Co-Typus’ label, and belonging to Chaoilta. Identity. - Uncertain. Campyloneurus marginiventris Type material. - Female holotype. Identity. - Campyloneurus marginiventris. Campyloneurus nigricosta Type material. - One female with ‘Co-Typus’ label hereby designated as lectotype; 3 female para- lectotypes with ‘Co-Typus’ labels and conspecific, agreeing well with the original description. One female has the basal half of the pterostigma yellow and is not conspecific. Because this character was not indicated in the original description this spec- imen cannot be the lectotype, despite its ‘Typus’ label. Identity. - Campyloneurus nigricosta. Campyloneurus punctativentris Type material. - Male holotype. Identity. — Campyloneurus punctativentris; a provisional identification because it is a male. Campyloneurus reticulatus Type material. - Female holotype. Identity. - Campyloneurus reticulatus Campyloneurus serenans Type material. - Female holotype. Notes. - The location of the type is not indicated by Shenefelt (1978); it is in the PAN collection. Identity. - Campyloneurus serenans. Campyloneurus speculiger Type material. - One female with ‘Type’ label hereby designated as lectotype; 4 female paralecto- types with ‘Co-Typus’ labels. Identity. - Campyloneurus speculiger. 256 Campyloneurus tibialis Type material. - One female with ‘Type’ label hereby designated as lectotype; 2 female paralecto- types with ‘Co-Typus’ labels. Identity. - Campyloneurus tibialis. Campyloneurus trispeculatus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 male paralecto- type with ‘Type’ label; 1 female paralectotype with ‘Co-Typus’ label. Identity. - Campyloneurus trispeculatus. Campyloneurus undicuneus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 male paralecto- type with Type’ label; 14 female and 5 male para- lectotypes with ‘Co-Typus’ labels. Identity. - Uncertain. Craspedolcus fraternus Type material. - One female with ‘Co-Typus’ label hereby designated as lectotype; 1 female para- lectotype with ‘Type’ label. The lectotype is not conspecific with the paralectotype and is the only specimen agreeing with the original description ( 1. Tergit poliert glatt‘). The paralectotype be- longs to Craspedolcus trisulcatus. Identity. - Craspedolcus fraternus. Craspedolcus obscuriventris Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 male paralecto- type with ‘Type’ label and one of the other males from Sukaranda must be also a paralectotype. The other 3 are obviously added later. Identity. - Craspedolcus obscuriventris. Craspedolcus quadrirugulosus Type material. - Female holotype. Identity. - Isomecus quadrirugulosus comb. nov. Craspedolcus strigidorsum Type material. - Female holotype. Identity. - Uncertain. Craspedolcus trisulcatus Type material. - One female with ‘Type’ label is missing the ovipositor sheath, both left wings and the right fore wing. Therefore we select and hereby designate the other specimen with 'Co-Typus’ label as lectotype. One female paralectotype with ‘Type’ label. Enderlein indicated in his description that he had a larger and a smaller specimen, so his state- ment at the end of the description that he had only one female is obviously false. Identity. - Craspedolcus trisulcatus. Cyanopterus chilensis Type material. - Male holotype. Notes. — The holotype has the head severely damaged, making its generic placement less certain at present. Identity. - Digonogastra chilensis; provisional identification. Cyanopterus filiseta Type material. - Female holotype. Identity. - Megabracon filiseta comb. nov. Cyanopterus solox Type material. - Female holotype. Notes. - This species was not mentioned in the recent revision of Exzrobracon Ashmead by Quicke (1989b). Identity. — Euurobracon triplagiata (Cameron) syn. nov. Cyanopterus thodeanus Type material. - Female holotype. Identity. - Compsobracon thodeanus comb. nov. Diamblomera acuticella Type material. - Female holotype. Notes. - This is the type species of Diamblomera Enderlein which was synonymised with Odontosca- pus Kriechbaumer by Quicke (1981); however, we now consider the two genera not to be synonymous and therefore we treat Diamblomera provisionally as a valid genus. Identity. - Diamblomera acuticella. Diamblomera grisescens Type material. - Female holotype. Identity. - Diamblomera grisescens. Diamblomerina speciosa Type material. - One female with ‘Type’ label hereby designated as lectotype; 2 female paralecto- types with ‘Co-Typus’ labels. Notes. - This is the type species of Diamblome- rina Enderlein which is treated here as a new junior synonym of Nedinoschiza Cameron. Identity. - Nedinoschiza speciosa comb. nov. Diolcia bicarinata Type material. - One female with ‘Type’ label hereby designated as lectotype; 4 female paralecto- types with ‘Co-Typus’ labels. Notes. - This is the type species of Diolcıa En- derlein which is treated here as a new junior syn- onym of Campyloneurus Szépligeti. Identity. — Campyloneurus bicarinatus comb. nov. QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae Diolcia limbaticauda Type material. - Female holotype. Identity. - Campyloneurus limbaticauda comb. nov. Ectemnoplax peruliventris Type material. - Female lectotype (hereby desig- nated; previously selected and labelled by C. van Achterberg); 1 male paralectotype and a long series of paralectotypes labelled as ‘Co-Typus’ by Ender- lein. Identity. - Ectemnoplax peruliventris. Ectemnoplax lurida Type material. - Female holotype. Identity. - Monocoila lurida comb. nov. Euryacria flavipera Type material. - Female holotype. Notes. - This is the type species of Euryacria Enderlein which was synonymized with Bathyaulax Szépligeti by Quicke (1981). Identity. - Bathyaulax flavipera: Quicke (1981). Gastrotheca sumatrana Type material. - One female with ‘Type’ label hereby designated as lectotype; the female paralec- totype is missing. Notes. - Only the lectotype was examined by Donaldson (1989). Identity. - Physarata sumatrana: (1978). Shenefelt Goniobracon denticornis Type material. - Holotype female. Identity. - Serraulax denticornis comb. nov. Goniobracon gutta Type material. - Female lectotype (designated by Shenefelt 1978: 1684); 1 male paralectotype. Notes. - Goniobracon Szepligeti is a junior syn- onym of Bathyaulax Szépligeti (see Quicke 1981); however, so many mistakes in the interpretation of these genera have occurred in the past that it is considered best to reclassify all species separately. Identity. - Bathyaulax gutta comb. nov. Goniobracon helvimacula Type material. - Male holotype. Identity. — Zaglyptogastra helvimacula comb. nov. Goniobracon pectinatus Type material. - Female holotype. Identity. - Bathyaulax pectinatus comb. nov. 257 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Hemibracon elegantulus Type material. - One female from Sumatra with Typus’ label hereby designated as lectotype; 10 male and 6 female paralectotypes are in PAN and additional paralectotypes from Taiwan are in the Deutsche Entomologisches Institut at Eberswalde. Notes. - All the specimens in the type series appear to be conspecific with one another. The generic placement of this species is hereby con- firmed but no statement is being made concerning its synonymy with Stenobracon trifasciatus Szé- pligeti, 1908 as proposed by Watanabe (1932) and through that, with S. niceville: (Bingham, 1901). Identity. - Stenobracon nicevillei (Bingham). Hemibracon w-impressum Type material. - Female holotype. Identity. - Hemibracon w-impressus. Iphiaulax sauteri Type material. - Female lectotype with ‘Type! label hereby designated; 1 female and 4 males pa- ralectotypes with only lectotype label. Identity. - Iphiaulax sauteri. Iphiaulax udei Type material. - Female holotype. Identity. - [phiaulax udei. Ipobracon acuticampus Type material. - Female holotype; the head, the major part of the mesosoma and part of the met- asoma (including the ovipositor) missing because of damage by dermestids. Identity. - Digonogastra acuticampa comb. nov. Ipobracon alboniger Type material. - Female holotype. Identity. - Digonogastra alboniger comb. nov. Ipobracon angustisulca Type material. - One female with ‘Type’ label hereby designated as lectotype; 15 female paralec- totypes with ‘Co-Typus’ labels. Identity. - Uncertain, close to I. laevibasis Ender- lein (q.v.). Ipobracon aureopilosus Type material. - Female holotype. Identity. - Digonogastra aureopilosa comb. nov. Ipobracon basispeculum Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type with ‘Co-Typus’ label. Notes. - Probably conspecific with I. sexrugosus Enderlein (see below). Identity. - Uncertain. 258 Ipobracon biareatus Type material. - One female from Peru with ‘Type’ label hereby designated as lectotype; 1 female paralectotype with ‘Co-Typus’ label is not conspe- cific. Identity. - Digonogastra biareata comb. nov. Ipobracon bicristatus Type material. - Female holotype. Identity. - Digonogastra bicristata comb. nov. Ipobracon bicuneatus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type without ‘Co-Typus’ label. Identity. - Digonogastra bicuneata comb. nov. Ipobracon bicuneus Type material. - Female holotype. Identity. - Digonogastra bicunea comb. nov. Ipobracon brevicapulus Type material. - Female holotype. Identity. - Digonogastra brevicapula comb. nov. Ipobracon brevicuneus Type material. - One female with ‘Type’ label hereby designated as lectotype; 3 female paralecto- types with ‘Co-Typus’ labels and a further three females without ‘Co-Typus’ labels but clearly in- cluded in the type series by Enderlein (1920). Identity. - Digonogastra brevicunea comb. nov. Ipobracon cameroni Type material. - Female holotype. Identity. - Digonogastra cameroni comb. nov. Ipobracon columbianus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type with ‘Co-Typus’ label. Identity. - Digonogastra columbiana comb. nov. Ipobracon crassecrenulatus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type with ‘Co-Typus’ label. Identity. - Hemibracon crassecrenulatus comb. nov. Ipobracon crenulatus Enderlein (not Szépligeti) Type material. - Female holotype. Notes. - Given the replacement name crenulato- rius by Shenefelt (1978: 1814). Identity. - Digonogastra crenulatoria (Shenefelt) comb. nov. Ipobracon crista Type material. - One female with "Typus’ label hereby designated as lectotype; 1 male paralecto- type with ‘Typus’ label; 38 female and 14 male paralectotypes with 'Co-Typus’ labels. Identity. - Hybogaster crista comb. nov. Ipobracon digitatus Type material. - Female holotype. Identity. - Digonogastra digitata comb. nov. Ipobracon duploareatus Type material. - Female holotype. Identity. - Digonogastra duploareata comb. nov. Ipobracon ecuadorensis Type material. - Female holotype. Identity. - Digonogastra ecuadorensis comb. nov. Ipobracon fenestratus Type material. - Female holotype. Identity. - Digonogastra fenestrata comb. nov. Ipobracon flavicaligatus Type material. - Female holotype. Identity. - Digonogastra flavicaligata comb. nov. Ipobracon laevibasis Type material. - Female holotype. Notes. - Appears to be closest to Digonogastra Ashmead but may warrant placement in a separate genus when that group is revised. Identity. - Uncertain. Ipobracon laeviventris Type material. - Female holotype. Notes. - The head is missing and therefore we cannot be certain about the generic placement of this species at present. Identity. - Uncertain. Ipobracon latecrenulatus Type material. - Female holotype. Identity. — Digonogastra latecrenulata comb. nov. Ipobracon latefasciatus Type material. - Female holotype. Identity. - Digonogastra latefasciata comb. nov. Ipobracon laticampus Type material. - Female holotype. Identity. - Digonogastra laticampa comb. nov. Ipobracon laticuneus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type with ‘Co-Typus’ label. Identity. - Digonogastra laticunea comb. nov. QUICKE & VAN ACHTERBERG: Enderlein's Braconinae Ipobracon latisulca Type material. - One female with “Typus’ label hereby designated as lectotype; 1 male paralecto- type with “Typus label; 49 female and 3 male pa- ralectotypes with ‘Co-Typus’ labels. Identity. - Uncertain, probably close to Craspe- dolcus Enderlein. Ipobracon longicapulus Type material. - Female holotype. Identity. - Digonogastra longicapula comb. nov. Ipobracon macellus Type material. - Male holotype. Identity. - Digonogastra macella comb. nov. Ipobracon maculicosta Type material. - One female from Java with ‘Type’ label hereby designated as lectotype and has been so labelled; 1 male paralectotype also with ‘Type’ label; 5 female and 9 male paralectotypes with ‘Co-Typus’ labels. Notes. - The male with the ‘Type’ label may not be conspecific with the lectotype female because it has forewing vein cu-a more or less interstitial, hindwing vein 1r-m shorter than vein SC+R1 and a posteriorly simple propodeum. The lectotype be- longs to the Craspedolcus group having the poste- rior margin of the propodeum ‘pinched-up’ in the middle and forewing vein cu-a distinctly postfurcal (see Quicke 1984b). Identity. - Craspedolcus maculicosta comb. nov. Ipobracon magnificus Type material. - Female holotype. Notes. - Transferred to Lasiophorus by Fahrin- ger (1931); however, Plaxopsts, the genus to which it belongs, is distinct from Lasiophorus despite the fact that both have a strong facial protuberance (van Achterberg 1983). Identity. - Plaxopsis magnificus comb. nov. Ipobracon mediofuscus Type material. - Female holotype. Identity. - Digonogastra mediofusca comb. nov. Ipobracon nigripalpalis Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female paralecto- type with ‘Co-Typus’ label. Notes. — This species was transferred to Atany- colus Foerster by Shenefelt (1978); Hemibracon Szépligeti is closely related to Atanycolus. Identity. - Hemibracon nigripalpalis comb. nov. 259 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Ipobracon nigripectus Type material. - One female with ‘Type’ label hereby designated as lectotype; one paralectotype female with ‘Co-Typus’ label. Identity. - Digonogastra nigripecta comb. nov. Ipobracon obtusicampus Type material. - Female holotype. Notes. - The head, propleuron and ovipositor are missing. Identity. — Digonogastra obtusicampa comb. nov. Ipobracon ochripes Type material. - Female holotype. Identity. - Digonogastra ochripes comb. nov. Ipobracon parvispeculum Type material. - One female with ‘Type’ label hereby designated as lectotype; 2 paralectotype fe- males with ‘Co-Typus’ labels. Identity. - Uncertain, appears to be closely re- lated to I. sexrugosus (q.v.). Ipobracon penniseta Type material. - Female holotype. Identity. - Digonogastra penniseta comb. nov. Ipobracon rectivena Type material. - Female holotype. Identity. - Digonogastra rectivena comb. nov. Ipobracon rugiventris Type material. - One female with ‘Type’ label hereby designated as lectotype; one paralectotype female with ‘Co-Typus’ label. Identity. — Spathulibracon rugiventris comb. nov. Ipobracon sexrugosus Type material. - Male holotype. Identity. - Uncertain. Ipobracon sigillatus Type material. - Female holotype. Identity. - Digonogastra sigillata comb. nov. Ipobracon speculatus Type material. - Female holotype. Identity. - Digonogastra speculata comb. nov. Ipobracon trirugosus Type material. - Female holotype. Notes. - The location of the type specimen is not indicated in Shenefelt (1978); it is in the PAN collection. Identity. - Uncertain. Ipobracon tuberculatus Type material. - Female holotype. Identity. - Digonogastra tuberculata comb. nov. Megagonia breviseta Type material. - One female with "Typus’ label hereby designated as lectotype; 1 male paralecto- type with "Typus’ label; 1 female paralectotype with ‘Co-Typus’ label. Identity. - Iphiaulax breviseta comb. nov. Megagonia griseiseta Type material. - Female lectotype hereby desig- nated; one female paralectotype. Identity. - Zaglyptogastra griseiseta comb. nov. Megagonia plumiseta Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 paralectotype fe- male with ‘Co-Typus’ labels. Notes. - The lectotype is in poor condition with no head or anterior metasoma. The other specimen with ‘Co-Typus’ label was not selected, since, al- though its head is present, the mesosoma is dam- aged. Identity. - Zaglyptogastra plumiseta comb. nov. Merinotus bispeculum Type material. - One female with ‘Type’ label hereby designated as lectotype; four paralectotype males, one with ‘Type’ and three with ‘Co-Typus’ labels. Identity. - Shelfordia bispeculum comb. nov. Merinotus criniseta Type material. - One female with ‘Typus’ label hereby selected as lectotype and so labelled; one male paralectotype with ‘Type’ label; 3 female pa- ralectotypes with ‘Co-Typus’ labels. Identity. - Shelfordia criniseta comb. nov. Merinotus flagriseta Type material. - One female with ‘Type’ label hereby designated as lectotype; sixteen paralecto- type females with ‘Co-Typus’ labels. Additional material. — Five males det. as fla- griseta by Enderlein but without ‘Co-Typus’ labels. Notes. - No opinion is being given here as to whether the males that were tentatively identified by Enderlein are conspecific with the females of the type series. Identity. - Shelfordia flagriseta comb. nov. Merinotus gibber Type material. - One female with ‘Type’ label hereby designated as lectotype; 106 paralectotype females with 'Co-Typus’ labels. Notes. — All the specimens in the type series appear to be conspecific; the type series is currently split between two cabinet drawers. Identity. - Cratobracon gibber comb. nov. Merinotus gracilis Type material. - Holotype. Notes. - The metasoma is missing but Ender- lein’s label and original description indicate that it is a female. Identity. - Nesaulax gracilis comb. nov. Merinotus ingentiseta Type material. - One female with ‘Type’ label hereby selected as lectotype; one female paralecto- type with 'Co-Typus’ label. Identity. - Shelfordia ingentiseta comb. nov. Merinotus laticampus Type material. - One female with ‘Type’ label hereby selected as lectotype and so labelled; one male paralectotype with ‘Type’ label; fourteen fe- male paralectotypes with ‘Co-Typus’ labels. Notes. - The lectotype and the paralectotype male are not congeneric; the paralectotype male belongs to the genus Shelfordia Cameron. Identity. - Gronaulax laticampus comb. nov. Merinotus nigriseta Type material. - One female with ‘Type’ label hereby selected as lectotype and so labelled; eight female paralectotypes with ‘Co-Typus’ labels. Identity. - Gammabracon nigriseta comb. nov. Merinotus quadricarinatus Type material. - One female with ‘Type’ label hereby selected as lectotype; 1 ‘male’ paralectotype with a ‘Type’ label; 89 female paralectotypes with ‘Co-Typus’ labels. Notes. - The ‘male’ paralectotype is in fact a female with the ovipositor broken off near its base. Identity. - Shelfordia quadricarinata comb. nov. Merinotus rimicuneus Type material. - One female with ‘Type’ label hereby selected as lectotype and so labelled; one male paralectotype with ‘Type’ label; 2 female and 1 male paralectotypes with ‘Co-Typus’ labels. Identity. - Shelfordia rimicunea comb. nov. Merinotus ruficauda Type material. - Female holotype. Identity. - Rhadinobracon ruficauda comb. nov. QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae Merinotus trispeculum Type material. - One female with ‘Type’ label hereby designated as lectotype; four paralectotype males, one with ‘Type’ and three with ‘Co-Typus’ labels. Identity. - Uncertain. Monolcia angulosa Type material. - One female with ‘Type’ label hereby designated as lectotype; 5 female paralecto- types with ‘Co-Typus’ labels. Identity. - Campyloneurus angulosus comb. nov. Monolcia apicalis Type material. - One female with ‘Type’ label hereby designated as lectotype and so labelled; one male paralectotype with ‘Type’ label; three female paralectotypes with ‘Co-Typus’ labels. Identity. — Belongs to Pachybracon Cameron- group. Monolcia consimilis Type material. - 1 female with ‘Type’ label here- by designated as lectotype; 1 female paralectotype with 'Co-Typus’ label. Identity. - Uncertain. Monolcia flavicosta Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 paralectotype fe- male with ‘Co-Typus’ label. Notes. - The lectotype has the metasoma glued on to the mesosoma. Identity. — Provisionally included in Campylo- neurus despite its apically truncate scapus. Campy- loneurus flavicosta comb. nov. Monolcia flavimarginata Type material. - Female holotype. Identity. - Uncertain. Monolcia serenimanus Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 paralectotype fe- male with ‘Co-Typus’ label. Notes. - The lectotype and paralectotype do not appear to be conspecific, the paralectotype having a preapical dorsal nodus and apico-ventral serra- tions on the ovipositor, both of which are absent in the lectotype. Identity. - Uncertain. Monolcia tricarinata Type material. - Female holotype. Notes. — This is the type species of Monolcia Enderlein which is hereby synonymized with Cam- pyloneurus Szépligeti. Identity. — Campyloneurus tricarinatus comb. nov. 261 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Odontopygia tridentata Type material. - One female with "Typus’ label hereby designated as lectotype; 2 female paralecto- types with ‘Co-Typus’ labels. Notes. - This is the type species of Odontopygia Enderlein which was synonymized with Trigas- trotheca Cameron by Quicke (1987). Identity. — Trigastrotheca tridentata: Quicke (1987). Plagiozina pravivena Type material. - One female with ‘Type’ label hereby designated as lectotype; one male paralecto- type with ‘Type’ label; eight female and one male paralectotypes with ‘Co-Typus’ labels. Notes. - This is the type species of Plagiozina Enderlein which was synonymized with Nesazlax Roman by Quicke (1984a) and is most likely a synonym of N. flagellaris Roman, 1913. Identity. - Nesaulax pravivena: Quicke (1984a) Platybracon carinicornis Type material. - One female with ‘Typus’ label hereby designated as lectotype; 5 female paralecto- types with ‘Co-Typus’ labels. Notes. - Platybracon Szépligeti is a junior syn- onym of Chaoilta Cameron (see Roman 1913, Quicke 1987); however, the two were dealt with separately in Shenefelt's (1978) catalogue. Identity. - Chaoılta carinicornis comb. nov. Platybracon celebensis Type material. - Female holotype. Notes. - See P. carinicornis (above). Belongs to the decorata group of species having a horn-like projection above the plate-like facial projection. Identity. - Blastomorpha celebensis comb. nov. Platybracon comptus Type material. - One female with Typus’ label hereby designated as lectotype; 1 female paralecto- type with 'Co-Typus’ label. Notes. - See P. carinicornis (above). Identity. - Chaoılta compta comb. nov. Pseudospinaria sannio Type material. - Female holotype. Notes. - Erroneously included in the Exothecinae by Enderlein (1920: 54); Pseudospinaria Enderlein is also a member of the Braconinae. Identity. — Trispinaria sannio comb. nov. Rhadinobracon rhyssides Type material. - Female holotype. Identity. - Ischnobracon rhyssides comb. nov. Rhamnura capillicauda Type material. - Lectotype female (designated by Shenefelt 1978: 1719); 4 female and 1 male para- lectotype are in the Museum fiir Naturkunde in Berlin. Notes. - Redescribed and illustrated by van Ach- terberg (1981). Identity. - Rhamnura capillicauda. Rhamnura filicauda Type material. - Lectotype female (designated by Shenefelt 1978: 1719); 2 paralectotype females are in the Museum fiir Naturkunde in Berlin. Identity. - Rhamnura filicauda. Syntomernus pusillus Type material. - One female with ‘Type’ label hereby designated as lectotype and so labelled; one ‘male’ paralectotype with ‘Type’ label; 5 female pa- ralectotypes with ‘Co-Typus’ labels. Notes. - The ‘male’ paralectotype has no meta- soma and therefore its sex cannot be verified. Identity. - Syntomernus pusillus. Udamolcia albescens Type material. - One female with “Typus’ label is damaged, therefore we designate hereby one of the two females with ‘Co-Typus’ label as lectotype; 2 female paralectotypes one with ‘Type’and one with ‘Co-Typus’ label. Notes. - This is the type species of Udamolcia Enderlein which was synonymized with Pachy- bracon Cameron by Quicke (1984c). Identity. - Belongs to the Pachybracon-group. Udamolcia albimanus Type material. - Male holotype. In the collection under Campyloneurus. Identity. - Uncertain. Udamolcia denticornis Type material. - One female with “Typus label hereby designated as lectotype; 22 female paralec- totypes with ‘Co-Typus’ labels. Identity. - Uncertain; a member of the Atanyco- lus Foerster group (see Quicke 1987). Udamolcia gracilis Type material. - Male holotype. In the collection under Campyloneurus. Identity. - Uncertain. Belongs to the Campylo- neurus-group. Udamolcia insolita Type material. - Female holotype. In the collec- tion under Campyloneurus. Identity. — Belongs to the Pachybracon-group. Udamolcia jucunda Type material. - Female holotype. Identity. - Undabracon jucundus comb. nov. Udamolcia laqueata Type material. - One female with ‘Typus’ label hereby designated as lectotype; 1 male paralecto- type with “Typus’ label; 3 female and 1 male para- lectotypes with ‘Co-Typus’ labels. Identity. - Cordibracon laqueatus comb. nov. Udamolx aterimma Type material. - Female holotype. Identity. - Digonogastra aterimma comb. nov. Udamolx atricauda Type material. - Male holotype. Identity. - Archibracon atricauda: Quicke (1989a). Udamolx catharinensis Type material. - One female with ‘Type’ label hereby designated as lectotype; 1 female and 1 male paralectotypes with ‘Co-Typus’ labels and 1 male paralectotype without 'Co-Typus’ label. Identity. — Hemibracon catharinensis comb. nov. Udamolx gutta Type material. - Female holotype. Notes. - This is the type species of Udamolx Enderlein which was synonymized with Archi- bracon Saussure by Quicke (1983). Identity. — Archibracon cameroni Brues (syn- onymized by Quicke 1989a). Udamolx polita Type material. - Female holotype. Identity. - Hemibracon politus comb. nov. Udamolx vaucristatum Type material. - 1 female with ‘Type’ label is the holotype. The male (with only Enderlein’s identi- fication label) was excluded from the type series by Enderlein (1920: 100). Identity. - Hemibracon vaucristatus comb. nov. ACKNOWLEDGEMENTS We would like to thank Professor Dr B. Pisarski, Dr E. Kierych, Mr P. Marczak (PAN) and Magda Kierych for their help and hospitality during our visits to Warsaw and Lomna. Partly this research was made possible by a Royal Society short term exchange awarded to the senior author. REFERENCES Achterberg, C. van, 1981. The position of the genus Rhamnura Enderlein among the Braconidae, Hyme- QUICKE & VAN ACHTERBERG: Enderlein's Braconinae noptera). — Entomologische Berichten, Amsterdam 41: 88-91. Achterberg, C. van, 1983. Six new genera of Braconinae from the Afrotropical Region (Hymenoptera, Braco- nidae). — Tijdschrift voor Entomologie 126: 175-202. Donaldson, J. S., 1988. Revision of the braconid genus Physaraia (Hymenoptera, Ichneumonoidea). — Syste- matic Entomology 14: 15- 39. Enderlein, G., 1905a. Rhamnura, eine neue Braconiden- gattung mit ausserordentlich langem Legrohr des Weibchens. - Zoologischer Anzeiger 29: 195-198. Enderlein, G., 1905b. Neue Braconiden aus dem indischen und afrikanischen Gebiet. — Stettiner Entomologische Zeitung 66: 227-236. Enderlein, G., 1920. Zur Kenntnis aussereuropäischer Braconiden. - Archiv für Naturgeschichte 84(A) (11): 51-224. Fahringer, 1931. Opuscula braconologica. 2. Aethiopi- schen Region. Lieferung 5: 305-384. Granger, Ch., 1949. Braconides de Madagascar. — Mé- moires de l'Institut Scientifique de Madagascar, A) 2: 1-428. Quicke, D. L. J., 1981. A reclassification of some Oriental and Ethiopean species of Braconinae (Hymenoptera: Braconidae). - Oriental Insects: 14: 493-498. Quicke, D. L. J., 1983. The Afrotropical genus Archibracon Saussure (Hymenoptera: Braconidae: Braconinae): Characteristics and new generic synonymy. - Entomol- ogist’s monthly magazine 119: 147-150. Quicke, D. L. J., 1984a. Two new genera of Afrotropical Braconinae (Hymenoptera: Braconidae) with a partial review of those genera with ‘merinotoid’ metasomas. — Entomologist's Monthly Magazine 120: 37-45. Quicke, D. L. J., 1984b. Redescription of Craspedolcus with notes on included species and related genera (Hymenoptera: Braconidae). — Oriental Insects 18: 354-360. Quicke, D. L. J., 1984c. Further reclassification of Afro- tropical and Indo-Australian Braconinae (Hymenop- tera: Braconidae). - Oriental Insects 18: 339-353. Quicke, D. L. J., 1987. The Old World genera of Braconine wasps (Hymenoptera: Braconidae). — Journal of Nat- ural History 21: 43-157. Quicke, D. L. J., 1989a. Parasitic braconine wasps of the genus Archibracon (Hymenoptera: Braconidae). — Journal of Natural History 23: 29-70. Quicke, D. L. J., 1989b. The Indo-Australian and E. Pa- laearctic braconine genus Ezurobracon (Hymenop- tera: Braconidae: Braconinae). — Journal of Natural History 23: 775-802. Quicke, D. L. J., 1989c. Reclassification of some New World species of Braconinae (Hym., Braconidae). — Entomologist's monthly magazine 125: 119-121. Roman, A., 1913. Philippinische Schlupfwespen aus dem schwedischen Reichsmuseum. I. - Arkiv för Zoologi 8: 7-59. Shenefelt, R. D., 1978. Hymenopterorum Catalogus (nov. ed.). Part 15. Braconidae, vol. 10. - The Hague, Junk, pp 1425-1865. Watanabe, C., 1932. A revision of Braconid species para- sitic in the injurious insects of rice plant and sugar cane in Japan and Formosa. — Transactions of the Sapporo Natural History Society 12: 63-72. Received: 14 September 1990 Revised version accepted: 25 October 1990 263 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 INDEX TO SPECIES GROUP NAMES 258 257 262 255 262 258 acuticampus acuticella albescens albicans albimanus alboniger angulosa angustisulca apicalis (Antiolcus) apicalis (Monolcia) aterrima atricauda aureopilosus basalis basispeculum biareatus bicarinata bicristatus bicuneatus bicuneus bispeculum brevicapulus brevicuneus breviseta cameroni capillicauda carinicornis cariniventris catharinensis celebensis chilensis cincticauda cingulicauda columbianus comptus consimilis crassecrenulatus crenulatus criniseta crista denticornis (Goniobracon) denticornis (Udamolcia) digitatus disciventris duploareatus ecuadorensis elegantulus EXCISUS fenestratus filicauda 257 260 259 261 261 257 256 261 256 261 262 255 255 260 257 257 263 255 DI 256 261 262 263 259 255 259 263 259 259 255 259 261 259) 256 259 257 259 ZO] 259 259 256 259 256 259 255 255 255 256 259 260 filiseta flagriseta flavicaligatus flavicosta flavimarginata flavipera fraternus gibber gibbiventris gracilis (Merinotus) gracilis (Udamolcia) grata gratiosus griseiseta grisescens gutta (Goniobracon) gutta (Udamolx) guttifer helvimacula impressimargo ingentiseta insolita jucunda laevibasis laeviventris (Bathyaulax) laeviventris (Ipobracon) laqueata latecrenulatus latefasciatus latiangulata laticampus (Ipobracon) laticampus (Merinotus) laticuneus latispeculum latisulca limbaticauda longicapulus lurida macellus maculicosta maculistigma magnificus marginiventris mediofuscus merinotoides mitelligera nigriceps nigricosta nigripalpalis nigripectus nigriseta obscuriventris obtusicampus ochripes parvispeculum pectinatus penniseta peruliventris plumiseta polita pravivena punctativentris pusillus quadricarinatus quadrirugulosus rectivena reticulatus rimicuneus rhyssides ruficauda rugiventris (Bathyaulax) rugiventris (Ipobracon) sannio sauteri serenans serenimanus sexrugosus sigillatus solox speciosa speculatus speculiger strigidorsum subfractus sumatrana tibialis thodeanus tricarinata tridentata trirugosus trispeculatus trispeculum trisulcatus tuberculatus udei undicuneus vaucristatum w-impressum R. T. A. SCHOUTEN Museon, Den Haag A NEW SPECIES OF EUCHROMIUS GUENEE, 1845 (LEPIDOPTERA: PYRALIDAE: CRAMBINAE) FROM AUSTRALIA Schouten, R. T. A., 1990. A new species of Ewchromius Guenée, 1845 (Lepidoptera: Pyra- lidae: Crambinae) from Australia. - Tijdschrift voor Entomologie 133: 265-267, figs. 1-5. [ISSN 0040-7496]. Published 14 December 1990. Euchromius cornus sp. n. is described from Australia. The external characters, male and female genitalia are described and figured in detail. R. T. A. Schouten, Museon, Dept. of Biology, Stadhouderslaan 41, 2517 HV Den Haag, The Netherlands. Key words. - Pyralidae; Crambinae; Euchromius, Australia; new species. Presumably in 1991, a checklist of the Lepido- ptera of Australia will be published. The only spe- cies of Ewchromius occurring in Australia is des- cribed in this article to have the name available for this checklist. A revision of the other species will be published shortly. RTAS refers to author's collection, other abbre- viations (codens) used in the list of type material follow Arnett & Samuelson (1986). The terminol- ogy used in the descriptions follows Schouten (1988). Euchromius cornus sp. n. (figs. 1-5) Type material. - Holotype: 8, ‘Sherlock R. W. Australia. E. Clements. 98-188’, genitalia slide 17616 BM (BMNH). - Paratypes: 1 8, Alexandria (BMNH); 1 8, Comrav (BMNH); 2 6, 4 9, Broken Hill (RTAS, SAMA); 1 8, to Alhannau above Column (SAMA); 1 d, Stevenson River (SAMA); 2 8, 1 9, 3 km. NW by W. of Millstream HS. (ANIC); 1 4, 2 9, 1 km. NE of Millstream HS. (ANIC, RTAS); 1 g, 1 9, 1 km. N of Millstream HS. (ANIC, RTAS); 1 9, 1/2 km. WNW of Millstream HS. (ANIC); 2 6, 1km NNE of Millstream WA (ANIC, RTAS); 1 4, Camooweal (ANIC); 1 9, 31 km NNW Longreach, Q. (ANIC); 1 6, Kimberley Research Stn Via Wyndham N. W. Aust. (ANIC); 2 6, Ivanhoe. W. A (ANIC, RTAS); 1 8, Moora W Q (ANIC); 1 9, Limestone Creek, 42 km NNW of Boulia, Q. (ANIC); 1 9, JimJim Creek 19 km WSW of Mt. Cahill, N. T. (ANIC); 1 @, Charleville Q. (ANIC); 2 6,1 9, Injune Q. (ANIC, RTAS); 1 9, Cunnam- ula Q. (ANIC); 1 @, 1 9, Adaville Q. (ANIC); 1 9, Mil- merran Q. (ANIC). Diagnosis. - Externally almost indistinguishable from Euchromius species with a double medial fas- cia and frons with one point. Differs in male geni- talia from all species in having the uncus armed with a short, double, dorsal projection. Differs in female genitalia from E. geminus, E. californicalis, E. limaellus, E. ocelleus, E. saltalis, E. mythus and E. tanalis in having a short ductus bursae without clear sclerotizations. External characters (fig. 1). - Wingspan 16-22 mm. Frons produced forward, conical with clear point, creamy white, no ventral ridge; vertex creamy white; labial palp two to two and a half, sides creamy white at base, becoming light brown, creamy white from above and below; maxillary palp creamy white to light brown, dark ringed at base of last segment; antenna creamy white. Thorax creamy white to light brown; patagia creamy white; tegulae creamy white, evenly mottled. Forewing, Figs. 1-2. Euchromius cornus. 1, Head, lateral aspect, scales removed, scale 1 mm; 2, Male, sclerotization of tergite VIII, scale 1 mm. 265 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Fig. 3. Euchromius cornus, male genitalia, holotype; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect, slide BM 17616, scale 0.5 mm. groundcolour creamy white densely suffused with ochreous brown to brown scales; medial fascia dou- ble, arched to nearly straight, running to one-sixth to one-fifth of the dorsum; subterminal line ochre- ous brown, inconspicuous, slightly closer to termi- nation of posterior area than to terminal dots, sometimes about halfway between terminal dots and termination of posterior area; area adjacent to terminal dots white; eight or nine black terminal dots, formula 2-2-3-1 or 1-2-3-2 or 2-2-3-2; fringes shiny, creamy white with two to three brown lines. Hindwing creamy white, subterminal fascia absent, termen darkly bordered; fringes white with creamy white line. 266 Tergite VIII (fig. 2). - Tergite normally sclerot- ized, posterior part triangular, stalk only partly vis- ible, anterior pattern not visible. Male genitalia (fig. 3). - Uncus with a short, double, dorsal projection; gnathos longer, with dor- sal dagger-shaped projections at each side of base, two dorsal thorns relatively large, terminal part long; tegumen without appendix angularis; saccu- lus normal, processus of sacculus absent, processus basalis long strongly sclerotized, two processi infe- riores valvae, cucullus club-shaped, bent upward; juxta triangular, armed with two dorsal projections; vinculum small; aedeagus normal sized, one dentate cornutus anterior of anellus connection, second, Fig. 4. Euchromius cornus, female genitalia; ventral as- pect, slide BM 17469, scale 0.5 mm. SCHOUTEN: New Euchromius ------I- AL A >< si \ ayy) © U \ \ t Fig. 5. Distribution of Euchromius cornus. fainter group of cornuti more posterior. Female genitalia (fig. 4). - Papillae anales nor- mal; membrane of tergite VIII with cord-like scle- rotization, edges of tergite VIII not connected; os- tium lip-shaped; ductus bursae short, broad under ostium; ductus seminalis narrow; bursa copulatrix oblong, two long signa, about equal in size. Ecology. - Unknown. Two generations, the first starts in February and ends at the end of May, most specimens were caught in April and May. The se- cond generation flies from August to the end of November, most specimens in September and Oc- tober. Distribution (fig. 5). - Restricted to Australia, where it can be found in most provinces. REFERENCES Arnett, R. H. & G. A. Samuelson, 1986. The insect and spider collections of the world. 220 pp. - EJ. Brill/Flora & Fauna publications, Gainesville. Haworth, A. H., 1811. Lepidoptera Britannica, 3: 377-511. — London. Schouten, R. T. A., 1988. Revision of the species of the genus Euchromius Guenée, 1845 (Lepidoptera: Pyra- lidae: Crambinae) occurring in the Afrotropical region. — Zoologische Verhandelingen Leiden 244: 1-64. Received: 2 October 1990. Accepted: 11 October 1990. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 BOOK ANNOUNCEMENTS AND REVIEWS Larsen, T.B., 1990. The butterflies of Egypt. - Apollo Books, Lundbyvej 36, DK-5700 Svendborg, Denmark: 1-112, figs. 1-7, tables 1-4, plates 1-8. [ISBN 87 88757 14 5, price 240 Danish Crowns]. The current knowledge of the butterflies of Egypt is summarized for the first time since 1948. All 58 species are depicted in full colour, together with some photo's of the most characteristic hab- itats. For each species the distribution inside and outside Egypt is discussed together with their tax- onomical status, ecology and habitat preferences. Separate chapters deal with the ecological zones of Egypt and its characteristic species, the biogeo- graphy, migrant species, economic importance and species expected to be found in Egypt in the near future. This book is recommended warmly to eve- ryone working on Mediterranean butterflies or in- tending to visit Egypt. [R. T. A. Schouten] Merzheevskaya, O. I., 1989. Larvae of owlet moths (Noc- tuidae). Biology, morphology, and classification. — Amerind Publishing Co, New Delhi (outside India exclusively marketed by E. J. Brill, Leiden etc): i-xx + 1-419, figs. 1-97. ISBN 90-04-08804-0. Price 115.- Dutch guilders (appr. US $57.50). This book is a translation of ‘Gusenitsy Sovok (Noctuidae), ikh Biologiya i Morfologiya (Oprede- litel’)’, which was published in 1967. It provides information on the biology and morphology of owlet moth larvae and keys to their subfamilies, genera, and species. Morphological characters, body coloration, and dermal patterns are detailed for each species. The larvae of 144 species of owlet moths are described and data on developmental changes in instars given for 91 of them. This book should be helpful to entomologists, agricultural entomologists, plant-protection spe- cialists, and teachers and students of biology, agri- culture, and forestry. [Information from the publisher] Oman, P. W., W.J. Knight & M. W. Nielson, 1990. Leaf- hoppers (Cicadellidae): A bibliography, generic check- list and index to the world literature 1956-1985. - CAB International Institute of Entomology, Wallingford: 1- 368. ISBN 0 85198 690 0. Price not given. The book consists of a bibliography of more than 7000 titles for the period 1956-1985, an index to the bibliography to guide users to specified subject- matter fields and a complete check-list of generic and family-group names used in the Cicadellidae. Other important features are a review of cicadellid literature for the period 1942-1955, a compilation showing usage of family-group names in the taxo- nomic literature during the years 1956-1985 and an alphabetized list and grouping of subfamily and tribal names within the family. The present bibli- ography supplements Metcalf’s Bibliography of the Cicadelloidea. [From the introduction of this book]. Williams, D. J. & G. W. Watson, 1990. The scale insects of the Tropical South Pacific region. Part 3. The Soft Scales (Coccidae) and other families. - CAB Interna- tional Institute of Entomology, Wallingford, UK: 1- 267, figs. 1-71. ISBN 0 85198 659 5. Price not given. A continuation of the series started by the same authors in 1988 (cf. Tijdschrift voor Entomologie 132: 114). The families discussed in the present volume are Margarodidae, Ortheziidae, Eriococci- dae, Dactylopiidae, Coccidae, Cerococcidae, Lecano- diaspididae, Kerriidae, Asterolecaniidae, Conchas- pididae and Halimococcidae. Keys are provided to the families and genera, and many new species are described and illustrated. [From the abstract |. JAN H. STOCK Institute of Taxonomic Zoology, University of Amsterdam A NEW SPECIES OF AUSTRODECUS (PYCNOGONIDA) FROM NEW SOUTH WALES, AUSTRALIA Stock, Jan H., 1990. A new species of Austrodecus (Pycnogonida) from New South Wales, Australia. — Tijdschrift voor Entomologie 133: 269-272, figs. 1-9. [ISSN 0040-7496]. Pub- lished 14 December 1990. Austrodecus staplesi sp.n. is the first member of the genus described from continental Australia (New South Wales). The genus is mainly Gondwanian in its distribution, but for three species recorded from the western belt of the Pacific Ocean. Dr. J. H. Stock, Institute of Taxonomic Zoology, University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, The Netherlands. Key words. - Austrodecus, Pycnogonida; New Species; New South Wales; biogeography. The genus Awstrodecus s.str. counts actually 24 named and 5 unnamed species (Stock 1957, 1968, Hedgpeth & McCain 1971, Clark 1972a, b, Child 1983, 1988, Pushkin 1977). The genus was briefly mentioned, without specific identification, in a pop- ular paper by Staples (1977: 123) from ‘southern Australian waters ... [where it] may be associated with the hydroid Halicorniopsis elegans’. With the exception of one dubious record of A. glaciale Hogdson, 1907, off Tasmania (see Gordon 1944: 6, Stock 1957: 46), there are no other Australian re- cords, and as far as I know Staples’ Australian mate- rial never has been formally described. Through the courtesy of Dr. Harry A. ten Hove, Amsterdam, I received a single specimen, fortu- nately a male, of an Austrodecus collected on the coasts of New South Wales. It represents a new species, described in this paper. Family AUSTRODECIDAE Stock, 1954 Genus Austrodecus Hodgson, 1907 Austrodecus staplesi sp.n. (figs. 1-8) Type material. - 1 8 (holotype), Australia, New South Wales, Split Solitary Island, near Coff's Harbour; depth 12-14 m; rocky area with small caves, some corals, algae and some sand; 26 Apr. 1986; leg. H. A. ten Hove, P. Hutchings & R. Phipps (Zoölogisch Museum Amsterdam Ba. 3322): Description Holotype 8. - The new species belongs to the gordonae-section (Stock 1957) of the genus, char- acterized by a 4-segmented oviger and the absence of auxiliary claws. Trunk (figs. 1-2) with 4 tall mid-dorsal spurs, one on each trunk segment. Ocular tubercle taller than trunk spurs, but rather plump; eyes well-pig- mented. Less tall dorsal spurs on coxa 1 of legs 1 and 4 (1 spur), coxa 1 of legs 2 and 3 (2 spurs), and coxa 3 of legs 1 to 4 (1 spur). Abdomen (fig. 1) tuber- culate, slightly overreaching coxa 3 of leg 4. Palp (fig. 3) 5-segmented, segment 5 indistinctly subdivided, bifid (fig. 4). Oviger 4-segmented, 1 spine on segment 3, 2 median and 4 distal spines on segment 4 (fig. 5). Legs (fig. 6) with a long dorsodistal seta on femur and tibiae 1 and 2. Femur longest segment. Femoral cement gland ventral, situated slightly before mid- dle of segment; discharging through long duct (fig. 7). Other glands open through slightly raised pores (fig. 8) on dorsal surface of tibia 1 (4 pores) and tibia 2 (2 pores). Propodus strongly curved; sole with c. 6 spinules; no auxiliary claws. As far as I can ascer- tain, male genital pores situated on ventral surface of coxa 3 (!) of all (!) legs. Measurements of holotype (in um). - Length trunk (tip ocular tubercle to tip abdomen) 1556: length cephalic segment 653; width across 2nd lat- eral processes 539; length abdomen 406; length proboscis 1099. Third leg: first coxa 144; second coxa 144; third coxa 101; femur 376; first tibia 264; second tibia 241; tarsus 49; propodus 269; claw 120. Remarks Within the gordonae-section, the new species 269 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Figs. 1-8. Austrodecus staplesi n.sp., & holotype. 1, body, dorsal; 2, body, from the left; 3, palp; 4, distal part of palp; 5, oviger; 6, third leg; 7, femoral cement gland; 8, gland apertures on dorsal surface of tibia 2 of leg 3. differs from A. frigorifugum Stock, 1954, A. stocki Child, 1988, and A. oblongum Pushkin, 1977 (ge- neric status of the latter uncertain because of aber- rant shape of proboscis) in the absence of dorso- distal femoral spurs. A. gordonae Stock, 1954, has low mid-dorsal trunk tubercles, an almost straight and shorter propodus, and lacks a femoral cement gland tube. A. palazense Child, 1983 has a longer distal tubercle on palp segment 2, as well as tuber- culate and widely separated lateral trunk processes; the oviger of this species is unknown. As point of fact, only three species of Austrodecus share with the new species the pres- ence of a tubiform cement gland aperture located in the middle part of the ventral surface of femur. Two of these, A. aconae (Hedgpeth & McCain, 1971), originally described as a species of Pantopipetta, 270 and A. kelpi Pushin, 1977 differ from the new spe- cies in the absence or poor development of mid- dorsal trunk spurs, and in the axial implantation of the distal palp segment. The cther species, A. tubiferum Stock, 1957, shows no doubt the greatest resemblance to the new species. A. tubiferum is known from Sagami Bay (Stock 1954, 1957) and Okinawa (Child 1988), and has a similar pattern of spurs on trunk and legs. The new species differs from A. tubiferum in having longer mid-dorsal trunk spurs, a more strongly curved pro- podus, a slightly different configuration of the distal palp segments, and less slender legs (especially on the level of tibia 2). The propodus of A. tubiferum is shorter than tibia 2, that of A. staples: longer than tibia 2. The new species differs from all other members NS SKK | (A Ui AT mas LO la FR as Si V © = ur SS MS STOCK: A new species of Austrodecus iL È wi. Fig. 9. Global reconstruction at 100 Ma B.P., after Smith & Briden, 1977. Dots indicate the actual distribution of the 24 named and 5 unnamed species of the genus Austrodecus (some species known from more than one locality, some localities with more than one species). With the exception of the Japanese, Palau and Kermadec records, the distribution is essentially Gondwanian. of the genus in the combination of 4-segmented ovigers and lacking auxiliary claws. Etymology This species is dedicated to Mr. David Staples, of Melbourne, in recognition of his works on Austral- ian Pycnogonida. Biogeography Child (1983: 699, 1988: 55) has suggested a ‘west- ern Indian Ocean corridor’ and a ‘western Pacific corridor’ to explain the presence of Austrodecus species on Aldabra Atoll (Indian Ocean), and in warm or warm-temperate waters in the western Pacific (Kermadec Islands, Palau, Japan). Stock (1957: 24, fig. 10) presumed a relationship to the plate tectonics of the southern hemisphere land masses, long before plate drift was a fashionable subject. Fig. 9 shows that (when the distribution of Austrodecus is plotted on a late Mesozoic map of the continents), the genus is essentially Gondwan- lan, with extensions along the western margin of the Pacific Ocean. REFERENCES Child, C. A., 1983. Pycnogonida of the western Pacific islands, II Guam and the Palau islands. - Proceedings of the biological Society of Washington 96: 698-714. Child, C. A., 1988. Pycnogonida from Aldabra atoll. — Bul- letin of the biological Society of Washington 8: 45-78. Clark, W. C., 1972a. Pycnogonida of the Snares Islands. — New Zealand Journal of marine and freshwater Re- search 5: 329-341. Clark, W. C., 1972b. Pycnogonida of the Antipodes Islands. — New Zealand Journal of marine and freshwater Re- search 5: 427-452. Gordon, I., 1944. Pycnogonida. - Report of the B.A.N.Z. Antarctic Research Expedition (B) 5 (1): 1-72. Hedgpeth, J. W. & J. C. McCain, 1971. A review of the pycnogonid genus Pantopipetta (family Austrodeci- dae, emended) with the description of a new species. — Antarctic Research Series 17: 217-229. Pushkin, A. F., 1977. New species of Austrodecidae from the Antarctic. - Exploration of the Fauna of the Seas, Academy of Sciences of the U.S.S.R. 21 (29): 54-59. Smith, A. G., & J. C. Briden, 1977. Mesozoic and Cenozoic paleocontinental maps: 1-63 (Cambridge University Press). Staples, D., 1977. Pycnogonida, the "nobodies”. - Austra- lia’s Oceans (Journal of the Oceans Society of Austra- lia) 1: 122-125. Dj TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Stock, J. H., 1954. Pycnogonida from Indo-West Pacific, Australian and New Zealand waters. - Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kjabenhavn 116: 1-168. Stock, J. H., 1957. The pycnogonid family Austrodecidae. — Beaufortia 6: 1-81. 272 Stock, J. H., 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific Oceans. - Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kjobenhavn 131: 7-65. J. van TOL National Museum of Natural History, Leiden ZOOLOGICAL EXPEDITIONS TO THE KRAKATAU ISLANDS, 1984 AND 1985: ODONATA Descriptions and records of Malesian Odonata, 2 Tol, J. van, 1990. Zoological expeditions to the Krakatau Islands, 1984 and 1985: Odonara. Descriptions and records of Malesian Odonata, 2. - Tijdschrift voor Entomologie 133: 273- 279, figs. 1-11, tab. 1 [ISSN 0040-7496]. Published 14 December 1990. A list of the Odonata collected by the members of the Krakatau Expeditions (1984 and 1985), organized by the La Trobe University (Melbourne) and LIPI (Jakarta). Most of the species mentioned are not from the Krakatau Islands, but from westernmost Java (Jawa) or southernmost Sumatra (Sumatera). Agriocnemis pygmaea (Rambur) and Orthetrum t. testaceum (Burmeister) are recorded from the Krakatau Islands for the first time. J. van Tol, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Key words. - Odonata; Malesia; Krakatau; Java; Sumatra. Members of most entomological expeditions to the Krakataus have made collections of dragonflies (Odonata). Yukawa & Yamane (1985) recently summarized the present knowledge. Their list of references should be completed with the work of Ris (1912), since that publication provides the iden- tifications of the specimens collected by Jacobson in May 1908, and only referred to as ‘spec. 1° and ‘spec. 2’ by Jacobson (1909). Thornton & New (1988b) gave some preliminary results of the present mate- rial and some sight observations. The present collection of Odonata includes con- siderably more species, mainly due to sampling ac- tivities in the southernmost part of Sumatra and the westernmost part of Java. A detailed account of the general results of these expeditions were given by Thornton and Rosengren (1988), while Thornton & New (1988a) deal with the freshwater commun- ities. Presently, the only known relatively perman- ent water bodies on the Krakatau Islands are con- crete-lined ponds, one on Sertung and two on northern Panjang. Before 1940 there were several pools and a brackish-water lake with breeding pop- ulations of eurytopic Odonata, e.g. Pseudagrion mi- crocephalum (Rambur), Ischnura senegalensis (Rambur), Diplacodes trivialis (Rambur) and Pan- tala flavescens (Fabricius) (Dammerman 1948). The following systematic account lists all species collected during the expeditions of 1984 and 1985. Species collected on the Krakatau islands Rakata, Sertung, Anak Krakatau or Panjang are marked with an asterisk. All records are given in full, but data on longitude-latitude are summarized in the next paragraph. LOCALITIES Sumatra. - Liwa (1), sweeping, 5°04’S 104° 03’E; Liwa (2), secondary forest, sweeping, 700 m, 5° 04'S 104° 03’E; Liwa (4), 5°04’S 104° 03’E. Krakatau Islands. — Rakata, Owl Bay, 6°09'S 105°28'E; Panjang, 6°05’S 105°28'E; Anak Kraka- tau, sweeping, 6°06'S 105°26'E; Sertung, forest, 6°05’S 105°23’E; Sertung, Spit, 6°04’S 105°24’- DONE: Java. - Ujung Kulon, Pulau Peucang, 6°45’S 105° 15’E; Ujung Kulon, Cideon, 6°46'S 105° 15’E; Ujung Kulon, Cibunar, 6° 48'S 105° 17’E; Hills be- hind Carita. The present and former names of the islands are: Rakata, also known as Rakata Besar and Krakatau; Panjang, also known as Rakata Kecil and Lang eiland; Sertung, also known as Verlaten eiland. Compilation of information on topography and the history of events are to be found in Dammerman (1948), and the more recently published works of Simkin & Fiske (1983) and Thornton & Rosengren (1988). SYSTEMATIC LIST The systematic list follows Davies & Tobin (1984, 1985). Records for each species are arranged in chronological order. Short comments on distri- bution and abundance of the species are given, us- 275 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 ually based on Lieftinck's publications (Lieftinck 1934, 1935, 1953 and 1954) and the RMNH collec- tion, Leiden. Special attention was paid to notes giving details about occurrence in southern Sumatra and western Java. Especially Lieftinck’s records of ‘Mt Tanggamoes’ [= Gunung Tanggamus] are of great interest for the interpretation of the observa- tions from Liwa. The status of Odonata before 1940 is well established through many collecting trips of entomologists working for the Bogor museum (for- merly Buitenzorg). Abbreviations used: RMNH = National Mu- seum of Natural History, Leiden. Family Amphipterygidae Devadatta argyoides (Selys, 1859) Sumatra, Liwa (2), 191 BB, 1 Sep 1984, 1 female. Although generally uncommon, it was very abundant in the Gn Tanggamus area in December 1934 (Lieftinck 1935). Family Calopterygidae Neurobasis chinensis chinensis (Linnaeus, 1758) (fig. 1) Sumatra, Liwa (2), 191 BD, 1 Sep 1984, 1 male; Idem, 191-BA, 1 female. The nominal subspecies is common on Sumatra, including the southernmost part. Javanese popula- tions are ranked as a separate subspecies, N. chinen- sis florida Hagen. Vestalis luctuosa (Burmeister, 1839) Sumatra, Liwa (2), 191 BC, 1 male. Java, Ujung Kulon, Cidaon, 190 GA and 190 HC, 15 Sep 1984, 1 male, 1 female; Idem, 190 ND, 16 Sep 1984, 1 male. Vestalis luctuosa is a common species on Java, but on Sumatra it is confined to the southernmost part. V. amoena Selys, a common species of this genus on Sumatra, was not represented. Females of Newrobasis chinensis and the re- gional Vestalis species are superficially very similar, and are not easy to separate in the field, especially so, since in many specimens of N. chinensis the pseudopterostigma is absent. Both genera are most easily distinguishable based on their wing venation. The Arculus is oblique, but not angulated in Vestalis, while it is conspicuously angulated in New- robasis (figs. 1 and 2) (Fraser 1933). Besides, Neu- robasis has cross-veins in the median (basal) space. Both Vestalis species can be identified on the col- oration of the labium (cf. Schmidt 1934: figs. 19a- b). Vestalis lugens Selys Sumatra, Liwa (2), 190 BE, 1 Sep 1984, 2 females; Idem, Liwa (1), 191 AA, 6 Sep 1984, 1 female; Idem, 274 Figs. 1-2. Wing base in Calopterygidae. - 1 (above), Ves- talis amoena; 2, (below) Neurobasis chinensis. Liwa (2), 191 AB, 7 Sep 1984, 1 female. See remarks under V. /uctuosa. Family Chlorocyphidae Rhinocypha angusta (Selys, 1853) Sumatra, Liwa (2), 191 BD, 1 Sep 1984, 1 male. Common on Sumatra, but absent from Java. Rhinocypha selysi Kriiger, 1898 Sumatra, Liwa (2), 191 BC, 1 Sep 1984, 1 male. Species confined to Sumatra (contra Lieftinck 1934, cf. Lieftinck 1954). Generally a rare species, except in southern Sumatra. In the RMNH collec- tion 18 males and 13 females are represented. Most specimens are from the Gn Tanggamus area and Benkulen; the other sites are (from north to south): Deli, Bolimbingan, 600 m (leg. Straatman) [locality not found]; Emmahaven, Padang [1°00'S 100° 22’E] (leg. Van der Starre); Sandaran Agung (Kurintji) [= probably Sanggaran agung, Kerintji; 2°08'S 101°31’E] (leg. Jacobson). Family Euphaeidae Euphaea variegata (Rambur, 1842) Sumatra, Liwa (2), 191 BA, 1 Sep 1984, 1 male. Java, Ujung Kulon, Cidaon, 190 HC, 15 Sep 1984, 1 female. Widespread and common on Sumatra and Java; recorded as ‘very common’ from the S. Lampoeng Residency by Lieftinck (1935). It was also found on Panaitan Island (Lieftinck 1953). Family Coenagrionidae Agriocnemis femina (Brauer, 1868) (figs. 9-11) Sumatra, Liwa (1), 191 AA, 6 Sep 1984, 1 female. Eurytopic species, very common and widespread from Bengal, Assam (Fraser 1933) eastwards to northern Australia. See further notes under A. pyg- maed. * Agriocnemis pygmaea (Rambur, 1842) (figs. 5-8) Krakatau Is. - Anak Krakatau, 215.DE, 21 Aug 1985, 1 female [short-lobed form, headless |. New to the Krakataus. Habitat: all kinds of stag- nant waters. Presumably a wind-borne species (Lieftinck 1954). Very widespread , and usually very common in the afrotropical, oriental and australian region. VAN Tor: Odonata Krakataus 1984-1985 The females of Agriocnemis are not easy to dis- tinguish, since both A. femina and A. pygmaea are polytypic. In this area also another species, A. min- ima Selys (figs. 3-4), has to be taken into account. I have now provisionally examined material avail- able in the RMNH collection (figs. 3-11). It seems that at least two forms of both species based on the shape of the prothorax, can be recognized. Females of A. pygmaea may have a strongly developed hind- lobe with a median depression, or the hindlobe is nearly completely lacking (figs. 5-8). Apart from these structural forms, there also exist two colour forms, an orange-red and a greenish-brown form. Therefore, as already stressed by Ris (1916), four forms can be recognized, since differently shaped hindlobes of the pronotum occur in both colour forms. According to Ris (1916), Agriocnemis femina 10 11 Figs. 3-11. Prothorax in females of Agriocnemis, left lateral and dorsal view. — 3-4, A. minima, Java; 5-6, A. pygmaea long-lobed form from Sulawesi; 7-8, À. pygmaea, short-lobed form from Ceylon; 9, A. femina, form with medially depressed hind lobe, northern Borneo; 10-11, A. femina, form with upright hind lobe, Sumatra. 275 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 falls apart in at least four structurally different forms. Also the males of this species show geogra- phical variation. Females of A. femina always have a well-developed hindlobe, but it is strongly upright in specimens from Ceylon, or it is conspicuously depressed in the middle in more eastern popula- tions (figs. 9-11). According to Lieftinck (1954: 72 footnote) two well definable subspecies occur on Sumatra. Family Platycnemididae Coeliccia membranipes membranipes (Rambur, 1842) Sumatra, Liwa (1), 191 AA, 6 Sep 1984, 1 female. Common species on Java and Sumatra in slowly flowing waters. Family Aeshnidae Gynacantha basiguttata Selys, 1882 Java, Ujung Kulon, 190 AG, 17 Sep 1984, 1 male. Considered as ‘rare’ on Java by Lieftinck (1934); more common on Sumatra, and also known from Billiton, Simalur, Enggano and western Borneo (collection RMNH). Apparently not rare on Panai- tan Ísland (Lieftinck 1953). Breeds in forest pools. The adults fly only at dusk, as all Gynacantha species. Family Libellulidae Agrionoptera insignis insignis (Rambur, 1842) Java, Ujung Kulon, Pulau Peucang, 15 July 1984, 190 AD, 1 male; Idem, 190 AF, AK, AP, AR, Sep 1984, 1 male 3 females; Idem, 190 AC, 20 Sep 1984, 1 female; Java, Ujung Kulon, Cibunar, 190 GB, Sep 1984, 1 male; Java, Ujung Kulon, P. Peucang, 24.1, 22 Sep 1984, 1 female. Confined to the coastal area of a significant part of the oriental region. Locally abundant in e.g. man- grove forests. It breeds in shady pools in forests (Lieftinck 1954). Although it was once ‘one of the commonest, and certainly the most striking’ (Lief- tinck 1934) dragonfly of the Krakataus, it was not collected there during this expedition, nor by Yu- kawa & Yamane (1985). The collection made in 1908 by Jacobson (Ris 1912) consisted of this spe- cies and Diplacodes trivialis (Rambur, 1842) only. Cratilla lineata assidua (Lieftinck, 1953) Java, Ujung Kulon, Pulau Peucang, 11 Sep 1984, 1 male. A woodland species, obviously uncommon on Java. The nominotypical subspecies occurs widely on Sumatra and was once collected on the Krakatau islands, while subspecies assidua Lieftinck was re- corded from Panaitan Island (Lieftinck 1953). 276 Lathrecista asiatica (Fabricius, 1798) Java, Ujung Kulon, Pulau Peucang, 190 AJ, 13 Sep 1984, 1 male. Although widespread in the oriental region, Lief- tinck (1954) considered it uncommon on Java. Lyriothemis magnificata (Selys, 1878) Java, Ujung Kulon, Pulau Peucang, 190 AQ, 3 Sep 1984, 1 female. A rare species from Sumatra, Billiton and Java. Unknown from western Java according to Lieftinck (1934), but in the RMNH collection several spec- imens from this area are available, e.g. Udjung Kulon, Tjibunar [river], 28 Nov 1958, 1 female. Orthetrum sabina (Drury, 1770) Sumatra, Liwa (1), white light, 190 AC, 31 Aug 1984, 1 female; Idem, Liwa (2), 191 BE, 1 Sep 1984, 1 male. Java, Ujung Kulon, Pulau Peucang, 190 AN, 17 Sep 1984, 1 male; Idem, Carita, 310 A, 11 Aug 1985, 1 female. Widespread and common species; abund- ant in cultivated areas. Collected on the Krakataus by Yukawa & Yamane (1985). * Orthetrum testaceum testaceum (Burmeister, 1839) Sumatra, Liwa (2), 191 BA and 191 BC, 1 Sep 1984, 2 males; Sumatra, Liwa (4), 191 DA, 7 Sep 1984, 1 male. Krakatau Is., Penjang, 3 Sep 1984, 192 BA, 1 male. New to the Krakatau Islands. Common in the lowlands of Java and Sumatra, and also known from Panaitan Island (Lieftinck 1953). Breeding in all kind of waters (Lieftinck 1934). * Diplacodes trivialis (Rambur, 1842) Sumatra, Liwa (1), 191 AA, 6 Sep 1984, 1 female. Krakatau Is, Anak Krakatau, 192 AA, 13 Sep 1984, 1 female; Krakatau Is., Anak Krakatau, 310B; 15 Aug 1985, 1 female; Idem, 310 C, 17 Aug 1985, 1 male; Krakatau Is., Rakata, Owl Bay, 310 F, G and H, 26 Aug 1985, 1 male 2 females. Very common and widespread species. Recorded from the Krakataus since 1908. Neurothemis fluctuans (Fabricius, 1793) Sumatra, Liwa (4), 191 DB, 7 Sep 1984, 3 males 1 female; Idem, 191 DC, same date, 2 males. See remarks under the following species. Neurothemis fluctuans (Fabricius, 1793) / N. ramburii (Brauer, 1866) Java, Ujung Kulon, Pulau Peucang, 190 AQ, Sep 1984, 1 male; Idem, 190 AH, Sep 1984, 1 female (homeochrome); Idem, (no number), 11 Sep 1984, 1 female (homeochrome); Idem, 190 AI, 11 Sep 1984, 1 female (homeochrome); Idem, 190 AL, 13 Sep 1984, 1 male; Idem, 190 AE, 15 Sep 1984, 1 female (heterochrome); Idem, 24.1, 22 Sep 1984, 1 female (heterochrome); Ujung Kulon, 190 GA, 20 Sep 1984, 1 female (heterochrome). According to Lieftinck (1954) Newrothemis fluc- tuans is rare and extremely local on Java, while Neurothemis ramburt is widespread but very local there. I am unable to identify the specimens from Ujung Kulon; the specimens from Sumatra seem to belong to N. fluctuans. It is well known that iden- tification of the genus Newrothemis meets many problems in this part of their range. Lieftinck (1954: 151 footnote) mentions hybridization be- tween N. fluctuans, N. ramburi and N. terminata, three species frequently flying on the same site. Lieftinck (1953) records all three species from Pa- naitan Island, and according to Lieftinck (1954) they can usually be kept apart fairly easily. I have examined large series in the RMNH collection, all identified by Lieftinck, but relevant characters seem to show significant variation between populations. A further study of this complex is needed to judge on the status of these taxa. Also Ris (1911: 567) has expressed his doubts about the specific status of these taxa. An extensive study of the variation of the characters involved (e. g. size of wing marking, number of cubito-anal cross-veins) could contribute Table 1. Odonata on the Krakatau islands. Rakata Besar Species Before Now Coenagrionidae Pseudagrion microcephalum Ischnura senegalensis e Agriocnemis femina Agriocnemis pymaea Libellulidae Agrionoptera insignis e Cratilla lineata e Raphismia bispina Acisoma panorpoides La Orthetrum sabina Orthetrum glaucum Orthetrum testaceum Diplacodes trivialis Crocothemis servilia Neurothemis terminata Neurothemis tullia Pantala flavescens Rhyothemis phyllis Tramea limbata euryale Macrodiplax cora Aeshnidae Anax guttatus Total number of species Rakata Kecil Before Fi VAN Tor: Odonata Krakataus 1984-1985 to our understanding of the historical biogeography of this species complex. Neurothemis terminata Ris, 1911 Java, Ujung Kulon, hills behind Carita, 11 Aug 1985, 1 male; Idem, Cidaon, 190 HB, 15 Sep 1984, 1 male. A common species on Java. See also remarks under N. fluctuans / N. ramburii. * Neurothemis tullia feralis (Burmeister, 1839) Krakatau Is., Anak Krakatau, outer rim, Malaise trap 21-24 Aug 1985, 213 EA, 1 male. Second species known from Anak Krakatau, and only the second record of this species from the Krakataus (first record: Rakata, 23 Oct 1923). It is rather rare and local on Java (Lieftinck 1934); there are no reliable records from Sumatra. It inhabits marshlands and weedy ponds. * Rhyothemis phyllis phyllis (Sulzer, 1776) Krakatau Is., Panjang, 192 BB, 3 Sep 1984, 1 male. Rather common species on Java, probably also breeding in oligohaline waters. The present record is the first of this species from Panjang, and only the second record for the Krakatau islands (first record: Rakata, 1921). Anak Krakatau Sertung Now Before Now Before Now No — QW — — Ww ZT TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 Camacinia gigantea (Brauer, 1867) Java, Ujung Kulon, Pulau Peucang, 190 AM, 18 Sep 1984, 1 female. Widespread from Burma and Assam eastwards to Eastern New Guinea. Usually rather uncommon, restricted to low altitudes in non-cultivated areas. * Pantala flavescens (Fabricius, 1798) Krakatau Is., Sertung Spit, at camp, 310 D, 18 Aug 1985, 1 male; Idem, Sertung forest, 310 E, 19 Aug 1985, 1 male. A very common circumtropical species and strong flier. DISCUSSION The dragonfly fauna of the Krakataus is only a poor representation of the fauna of the ‘mainlands’ of Java and Sumatra. This can even be demonstrated with the present data, but especially when all data available about the fauna of Java and Sumatra, or even those about that of Panaitan Island, are com- pared with those from the Krakataus. The reason for the high turnover rate of the species involved must be the fact that there is hardly any water available as breeding site for freshwater invertebrates. Most specimens collected should be considered as stragglers from Java or Sumatra, and must have reached the Krakataus by active flight or transported by the wind. The fauna of Panaitan Island (c. 150 km?, c. 10 km from the mainland of Java) was found to be far more mature in 1951 and also included several stream-dwelling species. Lief- tinck (1953) concluded that the dragonfly fauna of that island was not entirely composed of species that had reached the island accidentally, but that certain species must have inhabited it since the time it was a part of Java. Not all species found on the Krakatau islands (table 1) were also collected on Sumatra and Java by the present expeditions. How- ever, they are all are eurytopic or inhabiting coastal biotopes, where they are common, and usually also abundant, in southern Sumatra and/or western Java (based on Lieftinck 1954). Nevertheless, in future one may expect an indi- genous dragonfly fauna on the Krakataus when the forests have become more mature. Several highly specialized species of Zygoptera as well as Anisop- tera, are known as breeders in phytotelmata, e.g. treeholes of decaying logs, bamboo stumps, or wa- terholes at the leaf-base of Pandanus (cf. Corbet 1983). Examples of this fauna are the members of the libelluline genus Lyriothemis Brauer (cf. Lief- tinck 1954, Kitching 1986), which breed in water- filled treeholes, and the pseudagrionine genus Am- phicnemis Selys, supposed to breed in water at the leaf-bases of Pandanus trees. This kind of Odonata may find a breeding site on the Krakataus in due time. It may even be true that representatives of 278 this kind of dragonflies already inhabit the Kraka- taus, since most of them are very hard to collect by their inconspicuous behaviour. From a zoologeographical point of view it is of interest to mention that the fauna of the Krakataus seems to be Sumatran rather than Javan, contrary to the Panaitan Island. ACKNOWLEDGEMENT I am indebted to Professor I. W. B. Thornton (Melbourne, Australia) for entrusting this collec- tion to me. REFERENCES Corbet, P. S., 1983. Odonata in Phytotelmata. In Phytotelmata: Terrestrial plants as hosts for aquatic insect communities (eds. J.H. Frank & L.P. Lounibos). pp. 29-54. Marlton, New Jersey: Plexus. Dammerman, K. W., 1948. The fauna of Krakatau 1883- 1933. Verhandelingen der Koninklijke Nederlandsche Akademie van Wetenschappen, afd. Natuurkunde (Section 2) 44: i-xii + 1-594 + plate 1-11. Davies, D. A. L. & P. Tobin, 1984. The dragonflies of the world. A systematic list of the extant species of Odo- nata. Volume 1. Zygoptera, Anisozygoptera. - Rapid Communications Societas Internationalis Odonatolog- ica, Supplement 3: 1-127. Davies, D. A. L. & P. Tobin, 1985. The dragonflies of the world. A systematic list of the extant species of Odo- nata. Volume 2. Anisoptera. - Rapid Communications Societas Internationalis Odonatologica, Supplement 5: 1-151. Fraser, F.C. 1933 Fauna of British India including Ceylon and Burma. Odonata. Volume 1. Taylor & Francis, London. Jacobson, E. R., 1909. De opneming van de Krakatau- groep in Mei 1908. 5. De nieuwe fauna van Krakatau. — Jaarverslag van den Topographischen dienst in Ne- derlandsch-Indié 1908: 192-206 (sep. 43-57), 1 table excl. Kitching, R. L., 1986. A dendrolimnetic dragonfly from Sulawesi (Anisoptera: Libellulidae). - Odonatologica 15: 203-209. Lieftinck, M. A., 1934. An annotated list of the Odonata of Java, with notes on their distribution, habits and life- history. - Treubia 14: 377-462 + 1 plate. Lieftinck, M. A., 1935. A synopsis of the Odonata (Dra- gonflies) of Sumatra. - Miscellanea Zoologica Suma- trana 92-93: 1-23. Lieftinck, M. A., 1953. Notes on some Odonata from Panaitan Island, Sunda Strait (West Java). - Idea 9: 51- 56, 2 maps. Lieftinck, M. A., 1954. Handlist of Malaysian Odonata. A catalogue of the dragonflies of the Malay Peninsula, Sumatra, Java and Borneo, including the adjacent small islands. - Treubia 22 (Supplement): i-xiii + 1-202. Ris, F, 1911. Libellulinen 5. - Collections Zoologiques du Baron Edm. de Selys Longchamps 13: 529-700 + plate V. Ris, FE, 1912. Uber Odonaten von Java und Krakatau gesammelt von Edward Jacobson. - Tijdschrift voor Entomologie 55: 157-183 + plate 6-8. Ris, F, 1916. H. Sauter’s Formosa-Ausbeute. Odonata. (Mit Notizen über andere ostasiatische Odonaten). — Supplementa Entomologica 5: 1-81, figs. 1-47, plates 1-3 excl. Schmidt, E., 1934. Odonata der Deutschen Limnologi- schen Sunda-Expedition. I. Imagines. Mit Beschrei- bungen zweier neuer Ictinus aus Celebes und Neu- Guinea. — Archiv für Hydrobiologie, Suppl. 13: 316- 397 figs. 1-93, tabs 1-3, tabs 1 + Plate 14-17 excl. Simkin, T. & R. S. Fiske, 1983. Krakatau 1883. The vol- canic eruption and its effects. - Smithsonian Institu- tion Press, Washington DC: 1-464, figs. 1-139, plate 1- 16 excl. Thornton, I. W. B. & T. R. New, 1988a. Freshwater com- munities on the Krakatau Islands. — Philosophical Transactions of the Royal Society of London (Series B) 322: 487-492, fig. 1, tab. 1. VAN ToL: Odonata Krakataus 1984-1985 Thornton, I. W. B. & T. R. New, 1988b. Krakatau inver- tebrates: the 1980s fauna in the context of a century of recolonization. — Philosophical Transactions of the Royal Society of London (Series B) 322: 493-522, figs. 1-2, tabs. 1-10. Thornton, I. W. B. & N. J. Rosengren, 1988. Zoological expeditions to the Krakatau Islands, 1984 and 1985: General introduction. — Philosophical Transactions of the Royal Society of London (Series B) 322: 273-316, figs. 1-24, tabs. 1-10, plates 1-14 excl. Yukawa, J. & Yamane, S. 1985. Odonata and Hemiptera collected from the Krakataus and surrounding islands, Indonesia. - Konty 53: 690-698. Received: 10 September 1990 Accepted: 27 September 1990 279 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 280 NEW TAXA DESCRIBED IN VOLUME 133 ACARI ASGCELOUChAMPÈLE A SANYEO EEE IT 247 Ascetolichus microthrix Pérez & Atyeo ..... 249 Ascetolichus ruidus Pérez & Atyeo ......... 249 COLEOPTERA Carphurus albipennis Wittmer ............ 109 Carphurus basilimbatus Wittmer .......... 112 Carphurus brendelli Wittmer ............. 113 Carphurus dumogaensis Wittmer .......... 110 Carphurusilyale Watters Seal: 1415 Carphurus pallidicornis Wittmer .......... 109 Carphurus partepallidus Wittmer ......... 112 Garphurus storks Wittmenes u. aan 114 Carphurus strigilatus Wittmer ............ 111 Carphurus testaceolimbatus Wittmer ....... 113 Carphurus totopallidus Wittmer .......... 111 Dromanthomorphus ranuensis Wittmer .... 118 Falsolaius semicostatus Wittmer .......... 177 Falsolaius sulawesiensis Wittmer .......... 116 Bamıssdeniicollsı Wattmer reren ee 119 EON IEN SMN IEEE GRAN AR 118 AUST ANUER ISA IEEE PETE EE: 119 Paracarphurus sulawesiensis Wittmer ...... 115 Telocarphurus nebulosus Wittmer ......... 115 COLLEMBOLA Superodontella ciconia Bedos & DETAEVEN caste. un Myc tit. rere erry tt EI 1) Superodontella flammata Bedos & lWeharyeng rn Re EER selon eee 25 Superodontella gouzei Bedos & Deharveng 21 Superodontella longispina Bedos & Deharveng Bean. dan aduk aye te 21 DIPTERA Cryptochironomus judicius Chaudhuri & Chatto- padhyayse AE ME AE AVE RO RAD De DI 154 Fletcherimyia abdita Pape ............... 48 Harnischia tenuitubercula Chaudhuri & Chattopad- AMENER 20 Manag LOA AEN Ien o 158 Microcerella adelphe Pape ............... 52 Microcerella aulacophyto Pape ............ 49 (nom. nov. for Aulacophyto auromaculata Townsend) Microcerella austrohartigia Pape .......... 51 (nom. nov. for Austrohartigia bicoloricauda Lopes) Microcerella bermuda Pape .............. DIL WMicnocenellalboeta Papel. 22: 222.2... 50 (nom. nov. for Boettia ecwatoriana Lopes) Microcerella boettcherimima Pape ......... 49 (nom. nov. for Boettcherimima hypopygialis Lopes) Microcerellaicanchiabape eae ee. 49 (nom. nov. for Carchia andina Lopes) Macnocercllagujuytapapem ren. EEE 49 (nom. nov. for Jujuyia alvarengai Lopes) Paracladopelma aratra Chaudhuri & Chattopadhyay 4/404 ees ore 160 Paracladopelma sacculifera Chaudhuri & Chattopadhyay sunt ne Cee ee eee 161 Polypedilum circulum Chaudhuri & Chattopadhyay ines er PR eee ee 162 Sarcophaga (Discachaeta) amputata Pape .... 39 Sarcophaga (Pierretia) inlicida pape ........ 41 Spirobolomyia latissima Pape ............. 54 Tripanurga guatemala Pape .............. 57 LEPIDOPTERA Biselachista spinigera Sruoga ............. 79 Cosmiotes kopetdagica Sruoga ............ 83 Cosmsotenpallenworuoga Garena TIE 83 Deltas bobaga van Mastrigt .............. 200 Deltas catocausta eeft van Mastrigt ........ 201 Delias luctuosa kuning van Mastrigt ....... 202 Deltas nieuwenhuisi van Mastrigt ......... 202 (nom. nov. for Deltas leucias roepkei Nieuwen- huis & Howart) Deltas nieuwenhuisi poponga van Mastrigt 204 Deltas oktanglap van Mastrigt ............ 201 Delrastsiert var Mas trict =. Sun u GSC 200 DelasisinakvantMastrigth seater e 200 Delias wollastoni abmisibilensis van Mastrigt 202 Elachista fuscofrontella Sruoga ............ 76 Elachista megagnathos Sruoga ............ 79 Elachista multipunctata Sruoga ........... 75) Euchromius cornus Schouten ............. 265 RenttiabilobaSruogan OICR TI ANO 76 Stigmella rolandi van Nieukerken ......... 239 Trifurcula austriaca van Nieukerken ....... 245 Trifurcula coronillae van Nieukerken ....... 21197, Trifurcula iberica van Nieukerken ......... 228 Trifurcula josefklimeschi van Nieukerken 225 Trifurcula luteola van Nieukerken ......... 215 Trifurcula puplesisi van Nieukerken ....... 215 Trifurcula silviae van Nieukerken ......... 229 Trifurcula victoris van Nieukerken ......... 220 ODONATA Leptogomphus pasta van Tol ............. 97 BhylioeyelalhamaraaBelle Era 29 Progomphus gel NBE 27 PYCNOGONIDA Austmnodecus staples Stock 2... 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Belle, A revision of the New World genus Newraeschna Hagen, 1867 (Odonata: Aeshnidae) published in volume 132 (2), page 283, unfortunately are depicted too large. The reproductions below show these figures in the correct size. Figs. 72-73. — Neuraeschna titania spec. nov., & holotype; 73, Neuraeschna titania spec. nov., 9 paratype from same locality as holotype. vili Tijdschrift voor Entomologie Volume 133, no. 2 Articles 121 B. Aukema Taxonomy, life history and distribution of three closely related species of the genus Calathus (Coleoptera: Carabidae). 143 J. Belle & J. van Tol Anomalagrion hastatum (Say), an American damselfly indigenous to the Azores (Odonata, Coenagrionidae). 149 P. K. Chaudhuri & S. Chattopadhyay Chironomids of the rice paddy areas of West Bengal, India (Diptera: Chironomidae). 197 H. J. G. van Mastrigt New (sub)species of Delias from the central mountain range of Irian Jaya (Lepidoptera: Pieridae). 205 E. J. van Nieukerken The Trifurcula subnitidella group (Lepidoptera: Nepticulidae): taxonomy, distribution and biology. 239 E. J. van Nieukerken Stigmella rolandi sp. n.: a widespread southern European species on Rosa (Lepidoptera: Nepticulidae) 245 T.M. Pérez & W. T. Atyeo New taxa of feather mites (Acarina, Pterolichidae) from Megapodes (Aves, Megapodiidae). 251 D.L. J. Quicke & C. van Achterberg The type specimens of Enderlein's Braconinae (Hymenoptera: Braconidae) housed in Warsaw. 265 R. T. A. Schouten A new species of Euchromius Guenée, 1845 (Lepidoptera: Pyralidae: Crambinae) from Australia. 269 J. H. Stock A new species of Austrodecus (Pycnogonida) from New South Wales, Australia. 273 J. van Tol Zoological expeditions to the Krakatau Islands, 1984 and 1985: Odonata. Descriptions and records of Malesian Odonata, 2. Book announcements and reviews 268 T.B. Larsen, The butterflies of Egypt; O. |. Merzheevskaya, Larvae of owlet moths (Noctuidae); P. W. Oman, W. J. Knight & M. W. Nielson, Leafhoppers (Cicadellidae): a bibliography, generic check-list and index to the world literature 1956-1985; D. |. Williams & G. W. Watson, The scale insects of the Tropical South Pacific region, 3. © Nederlandse Entomologische Vereniging, Amsterdam Published 14 December 1990 ISSN 0040-7496 té i 1 RS nr 3 2044 093 383 164 wanen ap eijsden AA rarr eneen IA VA sar pres ear nenn vernemen ver mun (mene rn neer er Oo Woets venen Rr ee ee ¢ severe > 5 Rigo DR PAC NEA TIR ATI MT 3 N 2 ~ “ gg ne wie i ea eres PORCO SE IPP Ve vr KIN PRET AAP ennn armere eet ee fume oe mar man mien reen eeN are Ne rar eere en TERRA ever repens seer aen TA it por: rate cero verde VS N een ine pa etek Ar Run sage hee ere nme n Rapina spingono et tn eig my nz N WI EN OA ETRE ITAL RIA ATE TA diend 7 n e ovvie en NEEN AIAR E L m pau ee pr de ERROR DI LU Ue Ee AE vene Gemen MEE ENI Ann cera ieri ae cagna MR pre ven men PT rar ED ORPI Ft ene eert ee REI eng ag perk kerr en arden E endorse ve AN nende enaar =~ D SRN Bang og coe rt Weer ten eRe UNE peace nn tn Te NT ee Zn ne ee a omroeren SL RAR ET o San ei peren YG N neun er waters Fe. 5 DEP D TTI SE ; ; re > IAA RES pros MONA dO ti vr re ne EP RON OE Sn è i eenn WPS avena pase fect fay a SEAN EN enen megen Can te PA FAUNE PS en oe AT ART + LE ES me Ur eut a RN rn er EA - : werker waver reren È EEG ee : to iaia Gy eee eg ar“ one anni vr en senna ay Ar D RANA ee PS EN US = EE de NET EA GD fp PRT APS > : peers tan SAINT Spe ar Te Apster 7 ea Ay EUA A END EDEN ee NE ASD Kanar see PUM Er A A RA ek eue EN : an era EEND OKEE Dr Erve er RAA PET PNM Cl E mp CN MER ICS irda etl Rent Res Ania ar pra tr ta ve + SN PRET VERLATE AAA Ne Ad D D a va AD ASE A RÉ LR MS ER Ta UNE jn BV . sr LE PO NP RE RAP D GR SI i mn April dits ENG evn enn ae su 2. 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