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CES EE rs Minty ee dn, Pa aa ete ve D DEZE? meri te heinen re Pori re Lars ÉRIC LIN ‘4 = SE ei HARVARD UNIVERSITY ta Library of the Museum of Comparative Zoology i & 7 N°) re A 4 4 2 \ ' + ; | i i | ty volume 156, no. 1, 1775 ISSN 0040-7496 Tijdschrift voor > Entomologie A journal of systematic and evolutionary entomology since 1858 so So SE È. Ry < u = ti ale = ASI 3 = > 7 5 LF N EIS Published by the Nederlandse Entomologische Vereniging: _ ze Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address 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 Published with index of volume 135 (1992). Graphic design Ontwerpers B.V., Aad Derwort, ’s-Gravenhage DISTRIBUTION AND ECOLOGY OF HYDRAENA ]. G. M. CUPPEN ee RARY Department of Nature Conservation, Wageningen Agricultural University AUG 0 2 1993 KUGELANN IN THE NETHERLANDS (COLEOPTERWIVERS/ Ty HYDRAENIDAE) Cuppen, J. G. M., 1993. Distribution and ecology of Hydraena in The Netherlands. — Tijdschrift voor Entomologie 136: 1-10, figs. 1-16, tables 1-5. [Issn 0040-7496]. Published 1 July 1993. The distribution patterns of the species of the genus Hydraena in The Netherlands are figured. Their habitats are briefly described and compared with references. Physical/chemical data, phe- nologies and co-existence of species, based on observations by the author, are presented. Hydraena species mainly living in stagnant waters have relatively wide distributional areas in The Netherlands. Hydraena species from running waters are confined to the most eastern and southern part of the country and some of them may have disappeared at the beginning of this century. Chlorinity explains best the different distribution patterns of the stagnant water Hydraena. All stagnant water species can co-exist and their phenology is more or less the same with a high (early) spring maximum and most often a (lower) autumn maximum, indicating an univoltine life cycle. J. G. M. Cuppen, Department of Nature Conservation, Wageningen Agricultural University, Ritzema Bosweg 32a, 6703 AZ Wageningen, The Netherlands. Key words. — Hydraena; distribution maps; ecology; co-existence; The Netherlands. Beetles of the genus Hydraena Kugelann are small water beetles (1.5-2.5 mm) characterised by parallel sides, club-shaped antennae and very long maxillary palps. Sexes can be distinguished easily as males have six visible abdominal sternites, while females have seven. Many species show secondary sexual differen- ces, which sometimes offer useful characters for the identification of the species. Many species, however, can only be reliably identified in the male sex by their genitalia. In the past the study of genitalia has been neglected in The Netherlands owing to the lack of il- lustrated (Dutch) keys and by the former practice of relying entirely on external characters. An illustrated key to the Dutch species is now available (Cuppen 1992) to which one is referred for identification. The present survey is the first covering all species of Hydraena in The Netherlands and gives distribution maps based on old records, which have been checked in relation with taxonomic uncertainties, and recent- ly collected material. A survey of phenology and ecol- ogy is presented on the basis of literature and own field-work on mainly the stagnant water Hydraena. MATERIAL AND METHODS The material of this study includes the collections ‘Mededeling E. I. S. Nederland, no. 67 of the National Museum of Natural History, Leiden (RMNH), the Zoological Museum, Amsterdam (ZMA), the Departments of Entomology and Nature Conservation of the Wageningen Agricultural University, and the private collections of H. Cuppen (Apeldoorn), B. Drost (Wadenoyen), G. van Ee (Haarlem), J. Huijbregts (Leidschendam), B. van Maanen (Wageningen), H. Vallenduuk (Boxtel), C. Visser (Wageningen), B. van Vondel (Hendrik-Ido- Ambacht), O. Vorst (Utrecht) and the author. Some data from the Provincial Water Authorities of Noord- Holland and Utrecht, and from the Research Institute for Nature Management are also included. The maps are based on specimens, identified by the author. For Hydraena riparia Kugelann, Hydraena as- similis Rey and H. melas Dalla Torre in the H. ripa- ria-complex (see Jäch 1988), the maps are based on records of males only. The distribution of the Hydraena species in The Netherlands is plotted in the 10 km-squares of the UTM-grid (figs. 1-12) in which small dots refer to rec- ords before 1950 and large dots to records since 1950. On most localities visited by the author a water sample has been taken, which has been analysed the same or the next day on pH, electrical conductivity, chlorinity and total hardness. The results, viz. the ranges, 90% ranges and median values are presented in tables 1-4. TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-4. Distribution of Hydraena species in the Netherlands. — 1, Hydraena testacea Curtis; 2, Hydraena palustris Erichson; 3, Hydraena britteni Joy; 4, Hydraena riparia Kugelann (males only). Data on phenology and ecology are mainly based on the authors observations from The Netherlands and are compared with data from literature. Owing to the paucity of records for most species, statistical treatment of ecological data was not possible. For some species only old records are available and there is little information concerning ecology as labelling was insufficient in the past. HABITAT AND DISTRIBUTION All Dutch Hydraena species are widely distributed in Europe (lenistea 1978, Hebauer 1980, Horion 1949, Berthélemy 1986, d’Orchymont 1925, Jäch 1988). Detailed distribution maps for Hydraena, using any grid system, are given by Nilsson (1984) for the northern part of Sweden (50 km-squares) and Foster (1990) for the British Isles (10 km-squares). Relatively small on an European scale are the distri- bution areas of H. assimilis, H. melas, H. belgica d’Orchymont and A. excisa Kiesenwetter. The Dutch localities of H. excisa form an extreme western out- post of this eastern European species. Running waters including spring-fed streams are confined to the southern, central and eastern part of The Netherlands and therefore many rheophilic spe- cies of Hydraena are restricted to these parts of the country. Stagnant waters, however, occur all over the country. Brackish waters are mainly found in the south-western and northern part of the country (and before 1950 also around the Zuiderzee). The sandy soils in the southern, central and eastern part of the country, and the coastal dune area have usually a chlorinity of less than 100 mg/l, while the clay soils in the western and northern part have a chlorinity of more than 50 mg/l and gradually pass into the brack- ish areas. The main soil types per 10 km UTM grid- square are presented by Van Tol (1981). NOTES PER SPECIES H. testacea Curtis (fig. 1). This species is recorded from 89 squares since 1950 and additionally from 10 squares before 1950. It is the commonest species of the genus Hydraena in The Netherlands, collected mainly in the northern, east- ern and southern part of the country. From the west- ern part of the country there is only one recent record, apart from some pre 1950 records, despite large sampling programmes in recent years. The lack of records from Flevoland is probably due to under- sampling. South-Limburg apparently has no proper habitats for this species. Most records of H. testacea in the present investiga- tion are from temporary, semi-permanent and per- manent stagnant waters with a well developed margi- nal vegetation dominated by Carex, Glyceria maxima (Hartman) Holmberg, Phragmites australis (Cav.) Steudel, Calamagrostis canescens (Weber) Roth or Juncus effusus L. Certainly the marginal vegetation is not a necessity as the species was frequently found in waters without any marginal vegetation or with only a submerged vegetation. Also, in slowly running wa- Cuppen: Hydraena in The Netherlands ters such as regulated streams, H. testacea occurs fre- quently. In fast running waters the species is-extreme- ly rare in The Netherlands. This species is generally mentioned from stagnant waters, often with a well developed marginal vegetation as well as slowly run- ning wa.crs (d’Orch,.nont 1925; Horion 1949; Hrbacek 1951; Balfour-Browne 1958). Derenne (1952) mentions its occurrence in slowly running wa- ters dominated by algae. The occurrence in fast run- ning waters is only mentioned by Cuppen (1985) and Horion (1949) in Fontinalis in clear mountain- brooks. The habitat investigations by the author fair- ly well agree with literature for this species. H. palustris Erichson (fig. 2). This is an uncommon species recorded from 35 squares since 1950 and additionally from two squares before 1950. The general distribution pattern of H. palustris resembles that of H. testacea, but with much larger gaps and some isolated populations in the coastal dune area. The only area with a more or less continuous cover of squares is formed by the fresh water part of the river district. With one exception (in a regulated stream), all Dutch records of H. palustris are from temporary, semi-permanent and permanent stagnant waters. The species prefers temporary and semi-permanent, over- grown ditches and pools with Carex, Phragmites aus- tralis, Phalaris arundinacea L., Calamagrostis canescens and Glyceria maxima, in which the soil is covered by organic debris of the dominant plants. Most localities are unshaded or only partly shaded by Salix. In per- manent ponds, ditches and canals H. palustris has been found only where a well developed marginal vegetation of helophytes is present. Generally H. pa- lustris is considered to be a species of stagnant waters (d’Orchymont 1925; Derenne 1952), which are cov- ered by aquatic macrophytes (Horion 1949; Hebauer 1980) or mosses (Balfour-Browne 1958) as forest- pools, fens, ponds and ditches. The rare occurrence in running water is only mentioned by Horion (1949) and Hebauer (1980). Hebauer (1980) mentions a preference for acid waters, which could not be con- firmed in the present investigations. H. britteni Joy (fig. 3). This species was recorded from The Netherlands from 18 squares in 1982 by Cuppen & Cuppen. At present this rather common species is known from 73 squares since 1950 and additionally. from three squares before 1950. The low number of specimens in museum collections in comparison with other Hydraena species is very remarkable. The large in- crease of the number of records is caused by specific TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 sampling in autumn, winter and early spring. H. brit teni is confined to the southern and eastern part of the country with one old record from the province of Groningen (Haren; coll. ZMA). H. britteni fails in South-Limburg and occurs locally in the river area be- hind dikes of the rivers Rhine and Waal (Drost 1989). Records of H. nigrita by Everts (1898; 1922) belong to this species. H. britteni is considered to inhabit smaller stagnant waters, such as vegetation-rich pools and ditches, of- ten rich in mosses or with decaying leaves (Hrbacek 1951; Derenne 1952; Hebauer 1980; Nilsson 1984). Dutch habitats of H. britteni are characterized by Cuppen & Cuppen (1982) as temporary, weakly acid, stagnant waters with peaty soils covered by coarse organic debris. Such habitats are alder-brooks, forest-drains and -pools, marshes and swampy ditch- es dominated by Phragmites australis, Phalaris arundi- nacea, Calamagrostis canescens, Glyceria maxima or Carex. The acidophilic character of the habitat of H. britteni mentioned by Hebauer (1980) is confirmed above. The occurrence in a variety of running waters (Hebauer 1980; Balfour-Browne 1958), especially those with grassy edges in partial shade (Foster 1990) and cold springs (Nilsson 1984) is only confirmed for small sluggish streams receiving seepage-water. The incidental records in The Netherlands in larger streams most probably concern specimens that are washed out from their normal habitats during high water or flyers from habitats which have been dried out during the summer. There are some records of H. britteni, sifted from litter, in the summer period in dried out habitats (Vorst, pers. comm.). Landin (1976) mentions the presence of A. britteni and lar- vae of Hydraena above the water line. H. riparia Kugelann (fig. 4). After the splitting up of H. riparia and H. assimilis by Jäch (1988), a comparison between literature and the present data is difficult. H. riparia is known from 38 squares since 1950 and additionally from 13 be- fore 1950. The map gives only records of males and the species can be considered as rather common. Most records are confined to the river area with some recent records from the northern and western part of the country, and some old records from the south. Jäch (1988) mentions A. riparia from very fast flowing streams and not in stagnant waters, while Nilsson (1984) mentions exposed lakeshores among | | I > N ® 5 6 # 8 © (EI 9 10 11 el 12 Figs. 5-12. Distribution of Hydraena species in The Netherlands. — 5, Hydraena assimilis Rey (males only); 6, Hydraena melas Dalla Torre (males only); 7, Hydraena pygmaea Waterhouse; 8, Hydraena pulchella Germar; 9, Hydraena minutissima Stephens; 10, Hydraena gracilis Germar; 11, Hydraena belgica d’Orchymont; 12, Hydraena excisa Kiesenwetter. sand and gravel in Sweden. In England Balfour- Browne (1958) records the species from both stag- nant and running waters. The Dutch data contradict the findings of Jach (1988) and Nilsson (1984): there are no records of H. riparia in fast flowing waters nor on exposed lake shores. The species is mainly con- fined to stagnant waters and is occasionally found in slowly running, small streams. In fact most records are from temporary and semi-permanent habitats with a well developed vegetation of Phragmites australis, Glyceria maxima or Carex and soils covered by coarse organic debris or, in the case of running waters, small shady brooklets less than one meter wide. The occur- rence in larger ditches and ponds has been found only when there exists a well developed marginal vegetation or, single specimens have been found, when the ap- propriate habitats are within close vicinity. H. assimilis Rey (fig. 5). This very rare species is recorded from 9 squares since 1950 in the south-eastern part of the country (provinces of Limburg and Gelderland), but very probably some pre-1950 females from Limburg be- long to this species. The map gives only records of males. The few Dutch records of this species originate from small ditches and pools, receiving seepage-wa- ter, with or without vegetation, and from some small, relatively fast flowing streams. The only reference (Jäch 1988) mentions the same habitats as for A. ri- paria as the most probable. H. melas Dalla Torre (fig. 6). This very rare species was recorded for the first time from The Netherlands by Cuppen (1981) from Beertsenhoven. There are additional records of single males from Epen (coll. RMNH and coll. O. Vorst) and the Middelsgraaf near Echt (coll. J. Cuppen). The few Dutch records of H. melas are from spring-fed, small streams and, in one case, a larger, sun-exposed, slowly running stream. The references for H. meias are contradictory: Hrbätek (1951) and Hebauer (1980) mention preference for cold streams in mountainous areas, as well as acid ditches with much detritus in river valleys, while Jach (1988) men- tions mainly stagnant waters and small springs, but occasionally in larger streams and rivers. [H. nigrita Germar] The specimens from Warnsveld and Breda, men- tioned by Everts (1898, 1922), are deposited in the RMNH-collection and appear to belong to H. britteni. For this reason H. nigrita has to be removed from the Dutch list. CuppEn: Hydraena in The Netherlands H. pygmaea Waterhouse (fig. 7). This very rare species is recorded from three squares since 1950 in South-Limburg (Bunde, Epen, Houthem, Noorbeek en Cottessen) with the last record from 1991. The few recent Dutch records of H. pygmaea are from small (width less than 1 m), relatively fast flow- ing streams, which are heavily shaded in one case and unshaded in two. Generally this species is confined to running waters in hills and mountains (Horion 1949; Hrbätek 1951; Derenne 1952; Balfour-Browne 1958). Apparently H. pygmaea prefers sites with aquatic mosses and periphyton on wires and boulders (Hebauer 1980, Foster 1990). Hebauer (1980) refers to the species as cold-stenotherm and crenophilic, confined to the upper courses of streams and rarely found in the middle course. The Dutch data confirm the restriction of H. pygmaea to the smallest streams. H. pulchella Germar (fig. 8). Forty specimens of this very rare species were taken in four squares in South-Limburg and the vicinity of Winterswijk, all at the end of the 19th century or the beginning of the 20th century. A large series from 1921 (Wylre) represents the last record. H. pulchella is regarded to be extinct in The Netherlands. The Dutch records of H. pulchella suggest that, in the past, this species mainly occurred in the larger, fast-flowing streams (rivers Geul, Gulp and Ratumse Beek). Generally H. pulchella is confined to running waters, especially mossy stones and grassy edges (Hrbatek 1951; Derenne 1952; Balfour-Browne 1958; Hebauer 1980). Foster (1990) mentions ex- posed muddy pools in a small stream with riffles and pools, and amongst fine tree roots in a clayey gorge in a river. In comparison with H. pygmaea, this species is more restricted to middle courses and open, exposed parts of a stream. Hebauer (1980) refers to it as a rhe- obiontic, eurythermous species. H. minutissima Stephens (fig. 9). This very rare species is only known from two squares with 15 specimens altogether. The last record is from 1920 (Gulpen) and the species is regarded to be extinct in The Netherlands. The few (old) records from The Netherlands prob- ably all originate from the middle courses of the Geul and the Gulp, which are relatively fast flowing streams. Abroad this running water species has been found in small and large streams between gravel and aquatic mosses (Hrbacek 1951; Derenne 1952; Balfour-Browne 1958; Hebauer 1980; Valladares TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 1989). The distribution of H. minutissima seems to depend on strong current, rather than on tempera- ture-regime (Hebauer 1980). H. gracilis Germar (fig. 10) There are only three specimens from two squares known from The Netherlands. A record from 1923 (coll. ZMA) is the last and the species is regarded to be extinct in The Netherlands. Most records of this spe- cies by Everts (1898, 1922) appear to belong to either H. belgica or H. excisa. Generally A. gracilis is considered to be the most common running water Hydraena in western and central Europe, which occurs from the upper course till the lower course, as well in cold as in exposed streams and as well in mountains as on plains (Pretner 1930, Hebauer 1980, Valladares 1989). Also, with reference to the current velocity, the species shows a wide amplitude as Derenne (1952) even mentions stagnant waters. In these habitats the species can be found in mosses, under stones and in gravel (Horion 1949, Balfour-Browne 1958, Nilsson 1984). Despite this wide ecological amplitude H. gracilis has only been found twice in probably the middle and lower course of the Geul. H. belgica d’Orchymont (fig. 11) The only Dutch record is from the middle course of the Voer river (width of 3-5 m), probably from the beginning of the 20th century (coll. RMNH). No fur- ther details are known. H. belgica most probably no longer occurs in The Netherlands. Everts (1922) re- fers to these specimens as H. gracilis. The habitat of H. belgica in The Netherlands cor- responds with Derenne (1952), who simply refers to streams as its habitat. Hrbäcek (1951) mentions small (shady) streams with a moderate slope. H. excisa Kiesenwetter (fig. 12). This very rare species is known from only two Table 1. Range, 90% range and median of observed pH- values on localities of Hydraena species in The Netherlands. N is the number of observations. range 90% range median N Hydraena assimilis 6.3 - 9.0 6.8 6 Hydraena britteni 47-78 5.5-7.3 6.5 88 Hydraena melas 7.1 - 9.0 2 Hydraena palustris 3.5-9.2 5.3-7.7 6.8 45 Hydraena pygmaea 7.9 1 Hydraenariparia 3.3-83 6.2-7.8 7.0 174 Hydraena testacea 3.5-9.9 5.5 - 8.0 6.8 151 squares since 1950 in the vicinity of Winterswijk. Tolkamp (1980) lists H. excisa for the first time for The Netherlands, without mentioning it as an addi- tion to the Dutch fauna. However, Everts (1898) al- ready recorded H. gracilis var. erosa and var. excisa from Winterswijk, but the material could not be found in the museum collections. The first specimens from Winterswijk, labelled as H. gracilis, in these col- lections date from 1918 and all later references (Everts 1922, Laijendecker & Nieser 1971) belong to H. excisa. The recent Dutch records are from the Ratumse Beek (width about 3-5 m with shaded and unshaded sections) and some smaller tributaries. The species has been found here amongst gravel in relatively fast flowing parts of the stream (Laijendecker & Nieser 1971, Tolkamp 1980). Elsewhere in Europe H. excisa occurs in small, shady streams with a moderate slope (Hrbacek 1951) or sun-exposed streams (Hebauer 1980). According to the latter author the species is not cold-stenotherm and has no preference with re- spect to calcium. Within the streams it can be found in mosses or between and under stones (Knie 1974; Hebauer 1980). PHYSICAL AND CHEMICAL DATA The ranges of the observed pH-values (table 1) are large for most Hydraena species. However, the me- dian values of pH for all frequently encountered Hydraena species are between 6.5 and 7.0, with 90% of the observed values within 1.3 pH-unit above or below the median pH of that species. A. britteni has the lowest median pH-value and H. riparia the hig- hest. As the 90% ranges of pH for the different spe- cies considerably overlap and the median values are very close to each other, the pH can not be conside- red a very important factor for the explanation of the different geographic distribution patterns of these Hydraena species in The Netherlands. Ranges for the values of the electrical conductivity (table 2) are very large for A. riparia and H. testacea and large for A. britteni and H. palustris. The 90% ranges for the most common species reduce the upper limit for electrical conductivity considerably with the smallest range for H. britteni and a larger, more or less the same, range for the other species. The median val- ue of electrical conductivity is distinctly higher for H. riparia in comparison with the other species, which partly can be explained by the different distribution patterns of these species. However, also in this respect there is a considerable overlap in ranges. Ranges, 90% ranges and median values of total hardness (table 3) are more or less the same for the common Hydraena species and certainly cannot ex- plain their different geographic distribution patterns. Table 2. Range, 90% range and median of observed values of electrical conductivity (US cm') on localities of Hydraena species in The Netherlands. N is the number of observa- tions. range 90% range median N Hydraena assimilis 330- 710 490 6 Hydraena britteni 107-1260 192- 730 450 88 Hydraena melas 420- 620 2 Hydraena palustris 124-1590 150 - 1030 440 45 Hydraena pygmaea 580 I Hydraena riparia 55 - 2030 228-1140 560 174 Hydraena testacea 87-2030 174-1030 460 151 Table 3. Range, 90% range and median of observed values for total hardness ("D) on localities of Hydraena species in The Netherlands. N is the number of observations. range 90% range median N Hydraena assimilis 6-14 10 6 Hydraena britteni 1-23 3-16 8 46 Hydraena melas 9 1 Hydraena palustris 2-25 3-21 6 33 Hydraena riparia 1226. AD] 9 Vo, Hydraena testacea We) Mall 6 88 Based on only 6 observations, H. assimilis has a rela- tively small range with a high median value. Of the four investigated parameters, chlorinity is the most informative with respect to the distribution patterns of Hydraena (table 4). All species are con- fined to fresh water with only one sample with a chlo- rinity of more than 200 mg/l for H. britteni, H. palus- trisand H. testacea, while for H. riparia there are five. Chlorinity is certainly highly responsible for the ab- sence of Hydraena species in the western and north- ern part of the country of which the coastal dune area is excluded due to the presence of fresh waters. Median and 90% ranges of the most common Hydraena species are also in agreement with their general distribution patterns: H. riparia occurs main- ly between the large river systems, H. palustris is scat- tered all over the country with the exception of brack- ish parts, H. testacea occurs on sandy soils in the Cupren: Hydraena in The Netherlands Table 4. Range, 90% range and median of Cl--values (mg | ') on localities of Hydraena species in The Netherlands. N is the number of observations. range 90% range median N Hydraena assimilis 36.5 - 102.8 41.9 6 Hydraena britteni 6.8 - 270.0 14.5 - 92.7 36.5 88 Hydraena melas 19.8 - 41.9 2 Hydraena palustris 6.8 - 355.0 17.1 - 130.0 60.7 45 Hydraena pygmaea 19.6 1 Hydraena riparia 7.1 - 534.0 18.8 - 172.2 64.4 174 6.9 - 492.4 16.6 - 126.3 49.2 151 Hydraena testacea eastern, southern and northern parts of the country and between the large rivers and A. britteni is mainly found on the sandy soils in the southern, central and eastern part of the country. Co-EXISTENCE In table 5 the co-existence of five Hydraena species in The Netherlands is shown. The number of co-exis- tences for each species exceeds the total number of oc- currences because more than two species often co- exist. H. assimilis is found alone or in company with H. testacea. H. britteni lives most often alone or in co- existence with H. testacea. H. palustris lives most often in company with H. riparia and/or H. testacea, and only occasionally alone. H. riparia occurs most often alone, very frequently with H. testacea and frequently with H. palustris. H. testacea most often lives alone, very frequently with H. riparia and frequently with H. britteni and/or H. palustris. Proportionally H. britteni is most often the only Hydraena species at a locality, indicating that its hab- itat requirements may be more exclusive than those of the other Hydraena species. H. palustris rarely occurs alone, and it shares its habitat quite often with A. ri- paria and/or H. testacea. H. testacea, though frequent- ly occurring alone, co-exists quite often with all other Hydraena species, indicating that it can live in quite different habitats. H. testacea can be considered as the most euryoecious Hydraena species in The Netherlands. Table 5. Association of Hydraena species in The Netherlands. N is the number of observations. N Alone with H. with H. with H. with A. with A. assimilis brittent palustris riparia testacea Hydraena assimilis 10 4 = 2 = 1 6 Hydraena britteni 104 78 2 = 3 5 21 Hydraena palustris 52 11 - 3 = 28 19 Hydraena riparia 196 126 ] 5 28 — 51 Hydraena testacea 164 82 6 21 19 51 = TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 PHENOLOGY Phenological data at monthly intervals of adult Hydraena species are given for H. palustris, H. britte- ni, H. riparia and H. testacea (figs. 13-16). For the re- maining species the number of records of adults is too low to present meaningful diagrams. Keys for the identification of the larvae of Hydraena are not availa- ble and the larvae of only very few Hydraena species have been described so far (Hrbácek, 1943-44). As the number of collected larvae of this genus is very low and the above mentioned species quite often co- exist, it is impossible to include these data in the figu- res. H. palustris has been collected throughout the year with a distinct maximum in April, which probably coincides with the breeding period. Cuppen (1983) reports a different phenology for H. palustris with a spring maximum in May and a (lower) autumn max- imum in October. The same phenology was found by Landin (1976) in Lake Sagsjon near Stockholm, though the autumn peak occurred in September. Adults in copula, however, have never been observed. Teneral adults have been observed between 7.viii and 18.xi, indicating a rather long emergence period, and without a clear autumn maximum. This type of life cycle agrees with life cycle type I of Nilsson (1986). H. britteni can be collected throughout the year with clear maxima in March/April and November (the absence of records in January is certainly artifi- cial). A similar phenology has been reported by Cuppen (1983) in a study of a population of A. brit- teni in a seepage-marsh near Apeldoorn (The Netherlands) during one year. Landin (1976) found a spring maximum in the beginning of May and a (low- er) autumn maximum in September. The spring maximum coincides with the breeding period as many adults in copula have been observed between 340 | 120 50 100 + 40 u ao 30 so} 20 40 | 10 20 IL 0 0 J F M À M J IAS ON DO J 13 F M À MJ JF M À M J AES ORNE 14 Figs. 13-16. Number of specimens Aydraena spe- cies at monthly intervals. — 13, Hydraena palustris Erichson; 14, Hydraena britteni Joy; 15, Hydraena riparia Kugelann; 16, 16 Hydraena testacea Curtis. JAS OWN D 28.xii and 10.iv and the species can be considered as a winter breeder. Some pairs in copula even have been found in ice-covered pools. Teneral adults have been found in a very few occasions between 30.vii and 13.x before the November maximum. Cuppen (1983) mentions teneral adults in October and November. This low number of observations of teneral adults can be explained by the drying out of most habitats of A. britteni in the summer period. Adults probably leave their pupal chambers after the rising of the water lev- el due to autumn rains, when they are already fully developed. The November maximum coincides with the filling up with water of the habitats of H. britteni during the period in which their gonads develop. This type of life cycle with winterbreeding is not mentioned by Nilsson (1986). H. riparia has been collected throughout the year with two peaks in abundance in April and August/October. The spring maximum probably co- incides with the breeding period but pairs in copula have been found only once on 4.iii. Teneral adults have been observed in great numbers between 23.vii and 13.xi (mainly August and the beginning of September), and therefore coinciding with the late summer maximum. The life cycle agrees with the life cycle type I of Nilsson (1986). H. testacea has been collected quite evenly through- out the year with a maximum in April. This April maximum probably coincides with the breeding peri- od though pairs in copula never have been found. Teneral adults have been found in abundance between 24.vii and 14.xi with a distinct maximum in September. Also in this species its life cycle is in agree- ment with type I of Nilsson (1986). Based on the museum material, all other species have been collected mainly between May and September indicating more the activity of the collec- tors than the activity of the beetles. This also applies to the data for the above-mentioned species in the museum collections. The low number of records of Hydraena in museum collections is certainly partly based on the low activity of collectors in autumn, winter and spring. DISCUSSION The various distribution patterns of Hydraena in The Netherlands are in the first place determined by stream-velocity or a parameter associated with it. Most species are rheophilic or rheobiontic and there- fore confined to the southern and eastern part of The Netherlands. The maps suggest that nearly all species of this group are restricted to areas with fast running waters and do not occur in areas with more slowly running lowland streams. Paucity of recent records of these species permits no further conclusions about Cuppen: Hydraena in The Netherlands their preferred habitats. As running water species of Hydraena have no obvious morphological adaptations to that type of habitat it is difficult to judge which pa- rameters are responsible for their restricted occurren- ce. A permanent high oxygen content of the water could be a main factor, as most species disappeared after pollution and subsequent decrease of oxygen. For the stagnant water species chlorinity seems to be an important parameter for the explanation of their distribution patterns. All species are mainly re- stricted to waters with a chlorinity less than 200 mg/l. Other, less well known parameters, as soil type or the occurrence of seepage, probably also play an impor- tant role in their distributions. Quite often, however, two or three species can co-exist in the same locality, indicating that the different distribution patterns are not gouverned by the measured parameters. Until 1991 twelve species of the genus Hydraena have been found in The Netherlands. Most of them, if not all, have been collected before 1950, but many of them were not recognized until recently after dis- section of the male genitalia. From these twelve spe- cies only eight can be considered to occur at present in The Netherlands. H. pulchella, H. minutissima, H. gracilis and H. belgica have been collected for the last time at the beginning of this century and probably these species were confined to large streams as the Geul, Gulp, Voer (and Ratumse Beek) in the south- ern (and eastern) part of the country. Their disap- pearance is probably due to water pollution and, to a minor extent, to regulation of streams. Some of these species can, after improvement of water quality, re- turn to The Netherlands from neighbouring streams in Belgium and Germany. However, colonisation power of Hydraena species seems to be limited. The remaining running water species (H. melas, H. excisa, H. assimilis and H. pygmaea) are all very rare and under serious threat. Especially H. excisa, which occu- pies an extreme western outpost in The Netherlands, will not easily return after extinction. The chances for survival of the other three species, which occur main- ly in smaller streams or ditches receiving seepage-wa- ter, seem to be better, as most localities are situated in nature reserves. Hydraena species of stagnant waters are widely distributed and are not under threat. However, a decline of H. testacea and H. riparia in the western parts of The Netherlands is suggested by the available data. ACKNOWLEDGEMENTS This study would not have been possible without the ma- ny specimens, set at my disposal, from private and museum collections mentioned under material. Dr. G. N. Foster and anonymous referees are thanked for commenting on a first draft of this paper. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 REFERENCES Balfour-Browne, F., 1958. British water beetles III. — Ray Society, London, 210 pp. Berthélemy, C., 1986. Remarks on the genus Hydraena and revision of the subgenus Phothydraena (Coleoptera: Hydraenidae). — Annales de Limnologie 22: 181-193. Cuppen, H. P. J. J., 1983. Een oecologisch onderzoek naar de macrofauna van een temporair kwelmoeras op de Oost-Veluwe. I. De waterkevers. — Regionale Milieuraad Oost-Veluwe, Apeldoorn, 17 pp. Cuppen, J. G. M., 1981. Hydraena bohemica Hrbacek, nieuw voor Belgié en Nederland (Coleoptera, Hydrae- nidae). — Phegea 9: 61-64. Cuppen, J. G. M., 1985. Rediscovery of Hydraena testacea Curtis (Coleoptera: Hydraenidae) in Scotland. — Entomologist’s Gazette 36: 181-182. Cuppen, J. G. M., 1991. Genus Hydraena Kugelann. In: De waterkevers van Nederland (Coleoptera) (eds B. Drost, H. P. J. J. Cuppen, E. J. van Nieukerken & M. Schreijer). — Natuurhistorische Bibliotheek Koninklijke Neder- landse Natuurhistorische Vereniging 55: 280 pp. Cuppen, J. G. M. & H. P. J. J. Cuppen, 1982. Hydraena britteni Joy new for The Netherlands (Coleoptera: Hy- draenidae). — Entomologische Berichten, Amsterdam 42: 45-48. Cuppen, J. G. M. & K. Dettner, 1987. The larvae of the predaceous water beetle Laccornis oblongus (Stephens) (Coleoptera: Dytiscidae), with notes on ecology and dis- tribution. — Aquatic Insects 9: 211-220. Derenne, E., 1952. Les Hydraena de Belgique. — Bulletin et Annales de la Société Entomologique de Belgique 88: 195-218. Drost, M. B. P., 1989. Helophorus croaticus and H. pumilio in The Netherlands, with description of their larvae (Coleoptera: Hydrophilidae). — Entomologische Berich- ten, Amsterdam 49: 1-7. Everts, E., 1898. Coleoptera Neerlandica. I.: i-viii, 1-676. — Nijhoff, ’s Gravenhage. Everts, E., 1922. Coleoptera Neerlandica. III: i-xviii, 1- 667. — Nijhoff, ’s Gravenhage. Foster, G. N., 1990. Atlas of British water beetles. Preliminary edition. Part 6. — Balfour-Browne Club Newsletter 48: 1-18. Hebauer, F., 1980. Beitrag zur Faunistik und Okologie der Elminthidae und Hydraenidae in Ostbayern (Coleo- ptera). Mitteilungen der Miinchener Entomologischen Gesellschaft 69: 29-80. Horion, A., 1949. Faunistik der Mitteleuropäischen Kafer 10 II. — Vittorio Klostermann, Frankfurt am Main, 388 pp. Hrbätek, J., 1943-44. O larvách rodu Hydraena (Coleoptera, Hydrophilidae). — Sbornik Entomologic- kého Oddeleni Narodniho Musea v Praze 21-22: 84-89. Hrbätek, J., 1951. Revue des espèces du genre Hydraena Kug. sur le territoire de la république Tchecoslovaque (Col. Hydroph.). — Casopis Ceskoslovenske Spolecnosti Entomologickä 48: 201-226. Ienistea, M. A., 1978. Hydradephaga und Palpicornia. — In: Illies, J., Limnofauna Europaea. Gustav Fischer Verlag, Stuttgart, 532 pp. Jach, M. A., 1988. Revisional notes on the Hydraena ripar- ia species complex (Coleoptera: Hydraenidae). — Aquatic Insects 10: 125-139. Knie, J., 1974. Ein Beitrag zur Verbreitung und Okologie von Hydraena excisa Kiesw. — Decheniana 127: 263-264. Landin, J., 1976. Seasonal patterns in abundance of water beetles belonging to the Hydrophiloidea (Coleoptera). — Freshwater Biology 6: 89-108. Laijendecker, G. & N. Nieser, 1971. Waterkevers en water- wantsen uit de omgeving van Winterswijk (Coleoptera et Heteroptera aquatica). — Entomologische Berichten, Amsterdam 31: 3-12. Nilsson, A. N., 1984. The distribution of the aquatic beetle family Hydraenidae (Coleoptera) in northern Sweden, with an addenda to the Elmidae. — Fauna Norrlandica 4: 1-12. Nilsson, A. N., 1986. Life cycles and habitats of the north- ern European Agabini (Coleoptera, Dytiscidae). — Entomologica Basiliensia 11: 391-417. d’Orchymont, A., 1925. Les Hydraena de Belgique. — Bulletin et Annales de la Société Entomologique de Belgique 65: 45-53. Pretner, E., 1930. Ueber die angebliche Variabilitat der Hydraena gracilis Germar. — Coleopterologisches Zen- tralblatt 5: 81-105. Tol, J. van, 1981. An introduction to The Netherlands. — Nieuwsbrief European Invertebrate Survey - Nederland 10: 5-12. Tolkamp, H. H., 1980. Organism-substrate relationships in lowland streams: 1-211. Centre for Agricultural Publishing and Documentation, Wageningen. Valladares, L. F., 1989. Los Palpicornia acuaticos de la pro- vincia de Léon. II. Hydraena Kugelann, 1794 y Limnebius Leach, 1815 (Coleoptera: Hydraenidae). — Boletin de la Asociacion Espafiola de Entomologia 13: 313-330. Received: 12 November 1991 Revised version accepted: 10 September 1992 RYSZARD HAITLINGER Department of Zoology, Wroclaw Academy of Agriculture, Poland A NEW SPECIES OF THE GENUS DIPLOTHROMBIUM BERLESE (ACARI, PROSTIGMATA, JOHNSTONIANIDAE) FROM POLAND, BASED ON THE LARVA Haitlinger, R., 1993. A new species of the genus Diplothrombium Berlese (Acari, Prostigmata, Johnstonianidae) from Poland, based on the larva. — Tijdschrift voor Entomologie 136: 11-13, figs. 1-8. [issN 0040-7496]. Published 1 July 1993. Diplothrombium ludwinae sp. n. (larva) is described from Poland and a key is provided to the larval stages of the genus. Dr. R. Haitlinger, Department of Zoology, Wroclaw Academy of Agriculture, ul. Cybulskiego 20, 50-205 Wroclaw, Poland. Key words. — Acari, Johnstonianidae, new species, Poland. The genus Diplothrombium Berlese was established from the U.S.A.(Newell 1957, Feider 1959, Robaux in 1910. Since then, only four species based on larvae 1977, Southcott 1987). Larvae of Diplothrombium have been described, viz. D. monoense Newell, D. cas- were found attached to aquatic beetles (Coleoptera) cadense Newell, both from the U.S.A., D. moldavi- (Newell 1957) and Tipulidae (Diptera) (Feider cum Feider from Romania and D. newelli Robaux 1959). Up to now, only one species was known from all visible Figs. 1-8. Diplothrombium ludwinae sp. n. -- 1, idiosoma, dorsal view; 2, scutum; 3, idiosoma, ventral view; 4, palp; 5, palp- tarsus; 6, leg I, tarsus-trochanter; 7, leg II, tarsus-trochanter; 8, leg III, tarsus-trochanter. TIJDSCHRIFT voor ENTOMOLOGIE, VOLUME 136, 1993 Table 1. Measurements in mp of Diplothrombium ludwinae sp. n., larva, holotype. Length of idiosoma 904 Width of idiosoma 752 Length of scutum 108 Width of scutum 76 Distance between bases of AL (AW) 46 Distance between bases of PL (PW). 56 Length of anterior scutala (AL) 42 Length of posterior scutala (PL) 60 Distance between ASE (SBa) 10 Distance between PSE (SBp) 40 Length of anterior sensilla (ASE) 16 Length of posterior sensilla (PSE) 134 Distance between ASE ans PSE (ISD) 38 Distance between AL and PL (AP) 48 Distance between AL and ASE (AAS) 20 Ocular sclerite 34 Length of dorsal setae (DS) 78-90 Coxala I 56 Coxala II 58 Coxala III 40 fd 44 gd 36 Tal 104 Til 46 Ge I 44 TfI 52 BfI 46 Tre ll 40 Cx I 88 Ta II 84 Ti II 44 Ge II 40 Tf II 40 Bf II 40 Tr I 34 Cx II 76 Ta III 92 Ti Ill 62 Ge III 46 Tf Il 32 Bf III 62 Tr III 46 Cx III 86 Tale) > dal (A) 3.25 Til: Gel 1.04 Ti ll: Ge II 1.10 Ta III : Ge III 2.00 AW : ISD 1,211 ISD : AP 0.79 Til: AW 1.00 Ti II: AW 1.35 AW : AL 1.09 Europe. In the present paper a new species is de- scribed from Poland, and a key to the larvae of Diplothrombium is provided. Terminology for setae and structures follows Southcott (1988). All measure- ments are in micrometers (Um). Diplothrombium ludwinae sp. n. (figs. 1-8) Type material. — Holotype larva: Poland, Zawoja- Markowe, 800m (voi. Bielsko-Biala), 9.VI.1983, beech-co- niferous forest from plants, leg. R. Haitlinger (in Museum of Natural History, Wroclaw University). Description Idiosoma longer than wide, oval, anterior part of idiosoma somewhat deformed. Dorsum with 27 bar- bed setae arranged in five rows: 6, 7, 6, 5 and 3; each setae placed on oval and small platelet (fig. 1). Eyes small. Scutum longer than wide; behind bases of PSE with transverse crista which divides scutum into une- qual parts. Scutum with two pairs of scutalae (AL, PL); posterior setae PL longer than AL; further with two pairs of sensilla (ASE, PSE) from which posterior sensilla PSE many times longer than ASE. Between basis of ASE and posterior margin of scutum a longi- tudinal crista. All scutalae and sensilla smooth. Anterior margin with nasus (fig. 2). Ventral side of idiosoma with 74 setae, each placed on small platelet; posterior setae somewhat longer and placed on larger platelets. Anus located in middle part of opisthosoma. Two setae between coxae III. Gnathosoma short, its base with two bifurcate se- tae. Palpfemur and palpgenu each with one barbed seta; palptibia with three setae, two of these are barbed. Palptarsus long and with eight setae, five of which are barbed (figs. 4, 5). Legs short, length (including coxae, excluding claws): I 420, II 358, III 426; coxa I with one slightly barbed setae; medial coxala I bifurcate and separate from coxa; coxa II with one barbed seta; coxa III with one smooth seta. Tarsus II with enlarged solenidion and small famala. Number of solenidions and other setae on tibia, genu, telofemur, basifemur and tro- chanter | e 111 2250246, @ey72So NI 2 Sor aan lS or ONCS Or SOR ZN 2, Ne 1500 = IM Il Sor ©, Ge 2 So + 4, MI So + 3, Bir QI Measurements of holotype, sce table 1. Remarks. — D. /udwinae sp. n. is similar to D. mol- davicum Feider and D. cascadense Newell. It can be distinguished from the first species by the absence of a transverse bar on scutum above the posterior sensil- la, barbed coxalae II, shorter legs I-III and tarsi I-III, especially I and III; from the second species by the shape of the posterior part of the scutum and the smooth scutalae. Etymology. — Name derived from Ludwina. Key to species of Diplothrombium larvae il. Scutum with three transverse bars, two of them placed before the bases of posterior scutal sensilla teek cl BL D. moldavicum Feider, 1959 — Scutum without transverse bars before the bases o@posterorseutalisensi]t tn 2 2. Scutum with longitudinal and transverse bars be- low the bases of posterior sensilla ..................... 3 — Scutum without transverse and longitudinal bars below the bases of posterior sensilla .................. 4 3. Scutalae barbed, distance between bases of poste- rior scutalae PW distinctly larger than the dis- tance between bases of anterior scutalae; AW = 1.85 RENNES D. cascadense Newell, 1957 — Scutalae smooth, distance between bases of poste- rior scutalae somewhat larger than the distance between bases of anterior scutalae; AW = 1.22 … PE NSA RAI a RANA REACH AVIO TARE D. ludwinae sp. n. 4. Medial coxala I separated from coxa, genua I with 8 solenidions ............... D. newelli Robaux, 1977 — Medial coxala I on coxa I, genua I with 15 sole- DION NO D. monoense Newell, 1957 REFERENCES Feider, Z., 1959. Prima specie a genului Diplothrombium HAITLINGER: New species of Diplothrombium (Acari) din R. P. R. si Europa sub forma de larva (Diplothrombium moldavicum n. sp.). — Academia R. P. R. Filiala Iasi Studii si Cercetari Stiintifice si Stiinte Agricole 10: 261-268. Newell, I. M., 1957. Studies on the Johnstonianidae (Acari, Parasitengona). — Pacific Science 11: 396-466. Robaux, P., 1977. Observations sur quelques Actinedida (=Prostigmates) du sol d’Amérique du Nord. IX. Nouvelles formes larvaires de trombidions (Acari). — Acarologia 19: 258-271. Southcott, R. V., 1987. The classification of the mite fami- lies Trombellidae and Johnstonianidae and related groups, with the description of a new larva (Acarina: Trombellidae: Nothrotrombidium) from North America. — Transactions of the Royal Society of South Australia 111: 25-42. Southcott, R. V., 1988. Two new larval mites (Acarina: Erythraeidae) ectoparasitic on north Queensland cicadas. — Records of the South Australian Museum 22: 103-116. Received: 20 December 1992 Accepted: 12 April 1993 13 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 BOOK REVIEW Ossiannilsson, F., 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. — Fauna Entomologica Scandinavica 26: 1-346, figs. 1-1415 + 1-9. [ISBN 90 04 09610 8]. Published by E. J. Brill, Leiden. Price NLG 180.00 (appr. USD 105.00). The important series ‘Fauna Entomologica Scandinavica’, with much more relevance to other NW European countries than can be expected from the title, continues with another volume on Homo- ptera, viz. the Psylloidea. The author, F. Ossiannils- son, is a well-known specialist of this group and an author of Homoptera studies for more than fifty years. He also contributed earlier to this series with three volumes on the Auchenorrhyncha (1978- 1983). The present volume includes for each species syn- onymy, descriptions, including of larval stages, dis- tributional data and biology. Especially from the biol- ogy paragraphs, it is clear that the author is able to summarize his knowledge, based on the literature and on his own experience as well, in a concise and yet easily readible form. Moreover, a wealth of original data are provided for many of the species. Terminology is clearly explained in the introducto- ry chapters, while various keys guide the reader to the names of families, subfamilies, genera and species. Not less than 1424 figures illustrate all kinds of rele- vant characters of the 98 species included. Psyllids can cause, directly or indirectly, consider- able damage to their host-plants, and some species are specific to host-plants of economic importance (car- rot, pear). The present work is, therefore, not only strongly recommended to museums or amateur ento- mologists, but also to „vraries of applied entomolo- gists. Unfortunately, the publisher seems to discon- tinue his policy of producing this series for a reasonable price, although I certainly understand the connection with the limited number of copies that can be sold. Nonetheless, the price will inevitably af- fect the possibilities for the individual entomologist to purchase a personal copy. [J. van Tol] I. LANSBURY Oxford University Museum, United Kingdom STRONGYLOVELIA (VELIIDAE) AND METROBATOPSIS (GERRIDAE) AND ASSOCIATED PLEUSTON HEMIPTERA OF WEST NEW BRITAIN I. Lansbury, 1993. Strongylovelia (Veliidae) and Metrobatopsis (Gerridae) and associated pleuston Hemiptera of West New Britain. — Tijdschrift voor Entomologie 136: 15-22, figs. 1- 26 [issn 0040-7496]. Published 1 July 1993. A brief survey of the pleuston bugs of the Von River, West New Britain are given. Strongylovelia priori sp. n. is described (Veliidae: Haloveliinae) with distributional data from other localities. Two species of Rhagovelia (Veliidae: Rhagoveliinae) are listed. Metrobatopsis flavonotatus Esaki (Gerridae) is described and figured as its specific identity is uncertain. Other gerrid genera and species are listed, viz. Ptilomera, Limnometra and Tenagogonus. Dr. I. Lansbury, The University Museum, Parks Road, Oxford ox1 3pw, United Kingdom. Key words. — Hemiptera; Strongylovelia, Metrobatopsis, New Britain; new species. During a single visit to the Von River near Dami, West New Britain, the pleuston species diversity was found to be rather unusual as it included species char- acteristic of both lentic and lotic habitats. The river runs through a deeptly cut channel of what appears to be volcanic debris and silt. Substrate varying between coarse sand and gravel in slower shallow stretches and much deeper extremely fast stretches with large stone and boulders. The banks are steep, banded alternate- ly with black layers and broader intervening pale brown bands of silt. The habitat is deeply shaded by overhanging secondary forest and in places a dense sub-shrub layer. Supplementary data from other regions of Papua New Guinea are given for Strongylovelia priori sp. n., Metrobatopsis flavonotatus Esaki and other species of Veliidae and Gerridae. Data for Rhagovelia (Veliidae), Ptilomera, Metrobatopsis and possibly some species of Tenagogonus are found on slow-fast lotic habitats, whereas Limnometra species (all Gerridae) most com- monly found on lentic habitats. Published data on Strongylovelia give no indication of habitat preferences. SYSTEMATICS Veliidae, Haloveliinae Strongylovelia priori sp. n. (figs. 1-19, table 1) Type material. — Holotype d : Papua New Guinea, West New Britain, Balima River near Ulamo, 19.viii.1989, R. N. B. Prior. — Paratypes (all Papua New Guinea, West New Britain): Von River, Banaule near Dami, 21.11.1990, 129; Tamari Creek, freshwater/saline habitat, 11.xii.1988, R. N. B. Prior, 19. The holotype and a series of paratypes in OXUM, other paratypes in RMNH. Description Adult apterous. Male 1.6 mm long, maximum width 0.77 mm, female 1.58-1.76 mm long, maxi- mum width 0.86-0.99 mm. Coloration. — Female: Head black, latero-posterior margins and cephalic trichobothria dark yellow tinged pale red. Eyes silvery grey. Antennae black, first segment ventrally pale yellow. Rostrum shining dark brown to black. Pronotum black, anterior mar- gin narrowly pale yellow. Mesonotum broadly pale yellow, lateral margins black. Metanotum with two (1+1) yellow-orange blotches narrowly separated from yellow mesonotum. Anterior tergites uniformly black, distal tergites faintly iridescent. Connexivum posteriorly broadly pale yellow. Propleura narrowly black posterior of eyes, mesopleura broadly pale yel- low with upper margin and most of metapleura black. Metasternum pale yellow. Sternites laterally yellow, posteriorly uniformly blackish-brown, ventrally black (fig. 3). Legs: Front leg, trochanter and femur pale yellow, distally femur narrowly annulated dark brown. Inner margin of tibia anteriorly pale yellow, remainder and tarsi dark brown. Middle and hind legs black, tro- chanter and proximal part of hind femur pale yellow. Male similar to female, but pale yellow pronotal band narrower. Meso and metanotum and meso and metapleura as in female. Tergites black, not iridescent posteriorly. Connexivum and sternites black. Structure. — Female subovate, length 1.82x greatest 15 Figs. 1-5. Strongylovelia priori sp. n., 1-2, 5 holotype d , 3-4 paratype ?.—1, dorsal habitus; 2, ventral aspect; 3, dorsal hab- itus; 4, side view; 5, side view Scale line 0.5 mm. width across thorax. Head length about 1.6x greatest width. Eye width about 0.4x width of head between eyes. Thoracic dorsum clearly raised above tergites (fig. 4). Mesonotum and metanotum with sparse ad- pressed pubescence. Pro and mesopleura with much longer prominent curled black pubescence. Apical tergites broad, covered with adpressed greyish pubes- cence, distal tergites less pilose, shining greenish iri- 16 descent. Connexivum clearly raised above tergites, apically sinuate, curving and more erect posteriorly. Connexivum with long black curled hairs. Viewed laterally, connexivum posteriorly with a cluster of erect black hairs (figs. 3-4). Female genitalia (figs. 18-19). Tergum 8 large, first gonocoxae elongate. First gonapophyses distally with scattered prominent spines. Second gonapophyses ES \ I it N Ni) N MN AIN \ | I | | 3 Figs. 6-9. Strongylovelia priori sp. n. — 6, holotype d, anten- na; 7, same, hind leg; 8, paratype female, antenna; 9, same, hind leg. Scale line 0.5 mm. connected by a lightly sclerotised bridge, distally with fringes of fine hairs. Proctiger viewed laterally elon- gate, subquadrate from dorsal aspect. Male elongate, about 2x greatest width across tho- rax. Head length slightly less than greatest head width between eyes. Greatest eye width about 0.4x head width. Tergites slightly raised, connexivum continu- ous with tergites. Lateral margins of prothorax, ter- gites and connexivum covered with long hairs. Sternites not pilose with scattered minute spicules, more prominent and in organised row on distal seg- ment (figs. 1, 2, 5). Male genitalia (figs. 14-17): Genital segment hid- den within abdomen and moderately sclerotised, proctiger prominent. Parameres symmetrical, viewed laterally long and slender, from dorsal aspect sinuate. Legs: Male front femur moderately incrassate, tibia distally with a row of fine spinose hairs (fig. 10). LL = ZE LANSBURY: Strongylovelia of West New Britain Table 1. Proportions of leg segments of Strongylovelia prio- ri sp. n. Femur Tibia Tarsil Tarsi II Male front leg 60 60 8 20 Female front leg 63 63 8 22 Male middle leg 130 108 44 25 Female middle leg 141 105 40 25 Male hind leg 87 ij 13 20 Female hind leg 80 72 13 15 Middle femur about half length of male (21:41) (fig. 12). Hind femur slightly more robust than middle fe- mur, ventrally with numerous long spine-like hairs (fig. 7). Female front femur slightly curved, tibia dis- tally with a row of fine spinose hairs (fig. 11). Middle femur about half length of female (fig. 13). Hind fe- mur not noticeably thicker than middle femur, ven- trally fringed with fine hairs (fig. 13). Antennal segments 1-4 male (fig. 6) 26 : 37 : 63 : 52. First segment about 0.66x head length, second segment very robust. Female segments 1-4 (fig. 8) 27 : 33 : 50 : 42. First segment just over half median — head length. Remarks Strongylovelia priori sp. n. is similar in general ap- pearance to S. formosa Esaki (1924) described from two females from Taihoku, Northern Formosa. The hind femora of formosa are almost entirely pale and the pronotum and connexivum of the female are uni- formly dark brown to black. Lundblad (1933) figured the male of formosa from Central Sumatra, Singkarak. The parameres of formosa sensu Lundblad are proxi- mally very broad compared with priori. Lundblad gives the relative lengths of male antennal segments 1- 4 as 50 : 53 : 92 : 67. He states ‘In his Figure 1, Esaki, has shown the Ist segment too short, because it has apparently been drawn from an antenna that has not been detached. The condyle is therefore not visible. In fact the Ist and 2nd segments agree so closely in their length that no differences can be detected except by measuring them’. Figures 6 and 8 of priori are drawn from slide mounted partially cleared prepara- tions. Esaki described a second species, S. albicollis from New Guinea, Erima, Astrolabe Bay, based on a apter- ous female 1.5 mm long and a macropterous female 2 mm long. The apterous albicollis is distinguished from priori by the broadly yellowish white pronotum and the two yellowish white blotches on the black mesonotum. The yellowish white coloration of part of the connexivum is similar to priori. The variation of coloration between male and females of 107 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 10-13. Strongylovelia priori sp. n. — 10, holotype à, front leg; 11, paratype female, front leg; 12, holotype male, middle leg; 13, paratype female, middle leg. Scale line 0.5 mm. Strongylovelia and iridescence of the distal tergites have not previosuly been commented upon. Andersen (1982) figures the apterous female of S. for- mosa and forewing of macropterous form showing the extreme reduction of the venation. The longest series of Strongylovelia were collected from shaded moderately slow lotic stretches of the Von River, occurring with Metrobatopsis (Gerridae). Because of their small size, they were impossible to see on the surface of the river. Etymology. — A noun in the genitive case. This elu- sive haloveline is dedicated to Dr. R. N. B. Prior for his invaluable assistance whilst in West New Britain and for the collections he made for me during his ten- ure at Kimbe, West New Britain. 18 Rhagoveliinae Rhagovelia biroi Lundblad Rhagovelia biroi Lundblad, 1936: 13-14: type series New Guinea, Erima, Astrolabe Bay and Stephansort and East New Britain, Herbertshöhe [Kokopo near Rabaul] in HNHM [not examined]; Polhemus & Polhemus 1988: 165. Material. — Papua New Guinea, West New Britain, Von River, Banaule near Dami, 21.iii.1990, I. Lansbury, 17d 109, all apterous. This series were found principally along the margins amongst fallen branches. It is moderately com- mon at Tamari, Bialla and Bilomi River, Kimbe Bay, West New Britain. Polhemus & Polhemus (1988) place biroi in the ‘novacaledonica group’. LANSBURY: Strongylovelia of West New Britain Figs. 14-19. Strongylovelia priori sp. n., 14-17, holotype d genitalia; 18-19, paratype female — 14, segment 8 and ancilliary structures, scale line 0.25 mm; 15, genital capsule dorsal aspect; 16, same, side view; 17, part of segment 8. (Abbreviations S8, 8th segment; Pr, proctiger; pg, pygophore; c, clasper (paramere)). — 18, side view of ovipositor; 19, dorsal aspect of oviposi- tor. (Abbreviations t8, tergite 8; gx1, first gonocoxa; gol-go2, first and second gonapophysis; pr, proctiger). Scale lines 0.5 mm. Rhagovelia papuensis Lundblad Rhagovelia papuensis Lundblad, 1936: 28-30: type series New Guinea, Erima, Astrolabe Bay and Stephansort and East New Britain, Herbertshéhe [Kokopo near Rabaul] in HNHM [not examined]; Andersen 1982: 152; Polhemus & Polhemus 1988: 165. Material. — Papua New Guinea, West New Britain, Von River, Banaule near Dami, 21.iii.1990, I. Lansbury, 19 ap- terous. Found in small numbers at Tamari Creek and Balima River, Bialla where it was found in dense shade. Andersen (1982) figures the apterous male of papuensis. Polhemus & Polhemus (1988) include about twenty species in the ‘papuensis group’, which includes also À. australicus Kirkaldy, an australian endemic found in the N. Queensland rain forest habitats. Gerridae, Trepobatinae Metrobatopsis flavonotatus Esaki (figs. 20-26) M. flavonotatus Esaki, 1926: 144-146: type series New Guinea, Erima, Astrolabe Bay, in HNHM [not examined]; Hungerford & Matsuda 1959: 31-35 [lectotype d desig- nated]; Matsuda 1960: 367-371, 628-629. Material. — Papua New Guinea, West New Britain, Von River, Banaule near Dami, 21.11.1990, I. Lansbury, 13d 19 apterous, 19 macropterous and immatures; New Hanover, Lavonga Mission, small freshwater stream, 31.x.1989, R. N. B. Prior, 45 29 apterous and numerous immatures. TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 20-22. Metrobatopsis flavonotatus Esaki. — 20, macropterous female; 21, apterous male, dorsal habitus; 22, apterous male, side view. Scale line 1 mm. 20 Figs. 23-26. Metrobatopsis flavonotatus Esaki, male. — 23, front leg; 24, antennae; 25, genital capsule side view; 26, genital segment, ventral aspect. LANSBURY: Strongylovelia of West New Britain te SS TAS 26 Esaki (1926) and Hungerford & Matsuda (1959) de- scribe the variation in colour pattern and general structural features, the latter describing the male genitalia. The series from West New Britain and New Hanover differ slightly in the structure of the male genitalia. Description. — Adult apterous. Male 2.34-2.64 mm long, maximum width 1.0-1.1 mm; females in- cluding macropterous form with damaged wings (fig. 20) 2.6 mm long, 1.32 mm wide. Coloration: Male. Head, pronotum and mesonotum velvety black. Basal inner margin of head adjacent to eyes reddish brown. Pronotum and mesonotum with a prominent pale yellow longitudinal stripe, distal margin of mesonotal stripe evanescent blue. Metanotum, ter- gites, connexivum and metacetabulae velvety black with bluish evanescent patches, very noticeable on metano- tum and tergites. Genital segments black, proximally ir- idescent blue, lateral margins pale yellow (figs. 21-22). Pro and mesopleura pale yellow, distally the latter black overlaid with iridescent blue. Lateral margins of ster- nites blackish brown becoming paler ventrally. Genital segment ventrally pale yellow, distally black. Process of pygophore, suranal plate and paired processes shining black. Antennae black, proximal third of first segment pale yellow. The pale band is much more conspicuous when the gerrid is alive, likewise the proximal pale yel- low band on front femur. Trochanter of middle and hind legs pale yellow, remainder of legs black. Macropterous female (fig. 20) similar to male. Pronotum with a yellowish patch antero-mesially. Remnants of wings dark brown, venation black. 21 TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Male structure. Antennal segments 1-4 (fig. 24) 39.5: 25: 11.5 : 14. The ratios agree with Hungerford & Matsuda (1959); they are however, rather more ro- bust than those of the the female, contrary to Hungerford & Matsuda (1959). The inner median and distal surface of first segment with an area of pit- like structures. Front leg (fig. 23) femur proximally ro- bust, narrowing distally, tibia with a very prominent blunt pilose projection distally. Eighth segment long, laterally slightly sinuate, but not constricted as figured by Hungerford & Matsuda (1959), but does resemble figure by Matsuda (1960). Segment dorso-anteriorly with transverse striations (fig. 21). Ninth segment (figs. 25-26) with a prominent curved process usually ventrally but in some males the process is in a latero- ventral position. Suranal plate with a pair of sharply acuminate processes usually not visible (fig. 25). Gerrinae Tenagogonus (T.) kampaspe (Kirkaldy) Gerris kampaspe Kirkaldy, 1900: 804: type series New Guinea, Rigo and Kellesi in SNOW [not examined]. Limnometra kampaspe, Lundblad 1933: 371 [listed]. Tenagogonus kampaspe, Hungerford & Matsuda 1958: 386- 388 [redescription, locality Dilo Loria, vi, vii. 90]; Matsuda 1960: 212, 501 [species groups]. Material. — Papua New Guinea, Milne Bay Province, Naura, lotic stony river, 29.ix.1989, R. N. B. Prior, 14 macropterous; West New Britain, Von River, Banaule near Dami, 21.iii.1990, I. Lansbury, 32 apterous. Overall length of male including wings 5.39 mm, apterous female between 5.8-6.0 mm long. Limnometra monochroma Nieser & Chen Limnometra monochroma Nieser & Chen, 1992: 21-23. Material (type series). — Papua New Guinea, West New Britain, Buluma nr Dami, rain water pit, 17.1.1989, R. N. B. Prior, 26 19 (incl. 1d holotype) (OxUM); West New Britain, Von River, Banaule near Dami, torrential habitat, 21.11.1990, I. Lansbury, 22 (OXUM). Ptilomerinae Ptilomera breddeni Hungerford & Matsuda Ptilomera breddeni Hungerford & Matsuda, 1965: 451-454: 22 types New Guinea, SE Haveri; SE Paumenu River; Milne Bay (SNOW); Andersen 1982: 183, 191, 211. Material. — Papua New Guinea, West New Britain, Von River, Banaule near Dami, 21.iii.1990, I. Lansbury, 16 macropterous, 105 109 apterous. Collected in small num- bers on faster stretches and easily able to evade capture. REFERENCES Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera Gerromorpha). — Entomonograph 3: 1-455, figs. Esaki, T., 1924. On a new genus and species of the Gerridae from Formosa. — Annals Entomological Society of America 17: 228-230. Esaki, T., 1926. The water-striders of the subfamily Halobatinae in the Hungarian National Museum. — Annales Musei Nationalis Hungarici 23: 117-164. Hungerford, H. B. & R. Matsuda, 1958. The Tenagogonus - Limnometra complex of the Gerridae. — University of Kansas Science Bulletin 39: 371-457. Hungerford, H. B. & R. Matsuda, 1959. Concerning the genus Metrobatopsis Esaki with descriptions of a new spe- cies (Heteroptera: Geriidae). — Bulletin Brooklyn Entomological Society 44: 29-36. Hungerford, H. B. & R. Matsuda, 1965. The genus Ptilomera Amyot and Serville (Gerridae: Hemiptera). — University of Kansas Science Bulletin 45: 397-515. Kirkaldy, G. W., 1900. On some Rhynchota, principally from New Guinea (Amphibicorisae and Notonectidae). — Annali del Museo Civico di Storia Naturale G. Doria 20: 804-810. Lundblad, O., 1933. Zur Kenntnis der aquatilen und semi- aquatilen Hemipteren von Sumatra, Java and Bali. — Archiv fiir Hydrobiologie, Supplement 12: 1-195, 263- 489, pls. 1-21, figs. 1-142. Lundblad, ©., 1936. Die altweltlichen Arten der Velidengattungen Rhagovelia and Tetraripis. — Arkiv for Zoologi 28A: 1-63. Matsuda, R., 1960. Morphology, evolution and a classifica- tion of the Gerridae (Hemiptera-Heteroptera). — University of Kansas Science Bulletin 41: 25-632. Nieser, N. & P. Chen, 1992. Revision of Limnometra Mayr (Gerridae) in the Malay Archipelago. Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), II. — Tijdschrift voor Entomologie 135: 11-26, figs. 1-60, tab. 1 Polhemus, J. T. & D. A. Polhemus, 1988. Zoogeography, ecology and systematics of the genus Rhagovelia Mayr (Heteroptera: Veliidae) in Borneo, Celebes and the Moluccas. — Insecta Mundi 2: 161-230. Received: 17 March 1992 Accepted: 20 March 1992 I. LANSBURY Oxford University Museum, United Kingdom RHAGOVELIA OF PAPUA NEW GUINEA, SOLOMON ISLANDS AND AUSTRALIA (HEMIPTERA-VELIIDAE) Lansbury, 1., 1993. Rhagovelia of Papua New Guinea, Solomon Islands and Australia (Hemiptera-Veliidae). — Tijdschrift voor Entomologie 136: 23-54, figs. 1-150, tables 1-26. [ıssn 0040-7496]. Published 1 July 1993. Three species of Rhagovelia are known from ‘New Guinea’. R. papuensis Lundblad and R. biroi Lundblad are redescribed and figured. R. peggiae Kirkaldy is only known by the apterous female from Moroka and remains unrecognised at present. New species described from Papua New Guinea, R. priori sp. n.; R. hirsuta sp. n.; R. aureo- spicata sp. n.; À. herzogensis sp. n.; R. crinita sp. n.; R. thysanotos sp. n. and R. caesius sp. n. No records have so far been published of Rhagovelia from the Solomon Islands, R. browni sp. n. described from Guadacanal; R. amnicus sp. n. and R. fulvus sp. n. described from Malaita. I. Lansbury, Hope Entomological Collections, University Museum, Oxford, OX1 3PW, United Kingdom. Key words. — Papua New Guinea; Solomon Islands; Veliidae; Rhagovelia; key; new species. The Rhagovelia (Veliidae) are highly specialised pleuston bugs of lotic waters throughout most of the tropics. Andersen (1982) describes the general mor- phology and biology of Rhagovelia. Polhemus & Polhemus (1988) discuss the relative merits of various interpretations of the thoracic and abdominal struc- ture and their interpretation is followed in this re- view. The Rhagovelia of the ‘northern’ region of Papua New Guinea described in this review are from West Sepik (Torricelli Mountains), Madang, Eastern Highlands, Morobe, Oro or Northern, Milne Bay and West Britain Provinces. Additionally series from Malaita and Guadalcanal (Solomon Islands) are in- cluded. Lundblad (1936) described R. papuensis and R. bi- roi from Madang and East New Britain Provinces. Kirkaldy (1901, 1908) described R. peggiae and R. australica from Papua New Guinea and Australia, north Queensland, respectively. Polhemus & Polhemus (1988) revised the genus from Borneo, Celebes and the Moluccas adding 26 new species. They divided the Rhagovelia of the region into eight monophyletic infrageneric species groups using wing venation, thoracic morphology of macropterous forms and genitalia, and included some extralimital species in their groups. Of the taxa included in this review, R. papuensis, R. biroi Lundblad, R. amnicus sp. n., À. browni sp. n., R. herzogensis sp. n. and R. caesius sp. n. are known by both macropterous and the more common apterous forms. The remaining species de- scribed are only known by the apterous form. The Papua New Guinea species are tentatively as- signed to the ‘Polhemus’-groups (1988): Papuensis-group: broadly characterised by the greatly enlarged pronotum covering all but the poste- rior margin of the mesonotum. Included species R. papuensis Lundblad and R. priori sp. n. Additional species from the Solomon Islands R. fulvus sp. n., R. browni sp. n. and À. amnicus sp. n., and the australian endemic R. australica Kirkaldy. Bacanensis-group: Polhemus & Polhemus (1988) suggest that this group is closely allied to the papuen- sis-group. The forewing venation is similar to R. pa- puensis. Included species R. herzogensis sp. n., and also R. aureospicata sp. n., as the apterous males of À. her- zogensis and R. aureospicata are similar. The females are much easier to distinguish. The dark areas of the dorsum of both sexes having prominent fields of golden or silvery spicules. Novacaledonica-group. Included species R. nova- caledonica Lundblad, À. biroi Lundblad and possibly R. hirsuta sp. n. may belong to this group. As defined by Polhemus and Polhemus (1988) macropterous forms have four closed cells and an adventitious fifth cell on the forewing. The R biroi from Papua New Guinea examined in this study agree in all respects with Lundblad’s original description of the apterous form, but the forewing venation differs considerably from that figured by Polhemus & Polhemus (1988). Rhagovelia crinita sp. n. and À. thysanotos sp. n. dif- fer from other known species from Papua New Guinea. The male front tibiae are explanate with a cluster of long hairs / spines along the inner margin. Neither species is known by the macropterous form. Both have short pronotums compared with the me- 25 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 sonotum, the males are dorsally hairy and the genital capsules are very large. The female distal tergites are laterally variably pilose and the 8th and 9th abdomi- nal segments are sharply depressed. Rhagovelia caesius sp. n. Polhemus & Polhemus (1988: 167) comment on this group in northern New Guinea and mention another species from Negros in the central Philippines. Data relating to material col- lected by R. N. B. Prior suggests that R. caesius is found in slow flowing habitats in shady areas. It is a remarkable species which differs from other Papua New Guinea Rhagovelia by the absence of prominent projections on the hind femora of both sexes. Unlike other species, caesius is sexually dimorphic. The males dorsally are covered with long hairs, the females fine- ly tumescent, lacking long hairs. Females are abnor- mally flattened compared with males. The males rid- ing on the females, such phoretic behaviour has not been observed in other Rhagovelia species. There is some doubt that caeszus can be included in the same section of Rhagovelia s.s. as other species from this re- gion of Papua New Guinea. Kirkaldy (1901) described Rhagovelia peggiae, ‘Type’ apterous female from Central Province, Moroko [sometimes Meroka] 1300 m, 7.xi.1893, Loria. The type is thought to be in Genoa (MCSN). It is described as being very hairy and pubescent. Pronotum distinctly sutured from mesonotum, this suggests that the mesonotum may be longer than pro- notum. Anterior margin of prontum yellow other than lateral margins. The hind femora are ‘blue- greenish lustre’. The connexivum is much reflexed in- wards, meeting about the apex of the 7th connexival segment and continuing in contact up to the apex [somewhat as in R. distincta Champion, except that they do not diverge after meeting]. The description suggests that peggiae shares some features with hirsuta sp. n. A key to species excluding R. peggiae is given which will only work for apterous forms of both sexes. The australian endemic À. australica Kirkaldy is included in the key. Key to the species of Rhagovelia of Papua New Guinea, Solomon Islands and Australia 1. Pronotum covering all but a transverse narrow band of mesonotum (figs. 39, 63, 69) .............. À — Median pronotal length shorter than median length of mesonotum (figs. 73, 84) 2. Pro-mesopleura and proepisternum bare lacking fields of minute black spicules australica Kirkaldy Areas of propleura, pro-meso and metasternum and jugum or lateral margins of pro-meso and metanotum with fields of black spicules ........... 3 Anterior lateral margins of pro-meso and meta- notum with fields of black spicules ................... 4 Anterior lateral margins of pro-meso and meta- notum lacking fields of black spicules Pronotum anteriorly with a pale transverse band. First antennal segment proximally pale, remain- der black. Connexivum and tergites dark brown- Black... en ee 5 Pronotum more or less uniformly coloured, pale anterior band obsolete, antennae pale, connexi- vann amel nensites pelle onsassonossosersoscens fulvus sp. n. Sternites 2-5 with scattered black spicules, male connexivum either slightly reflexed outwards or almost erect (fig. 51). Female connexivum sinu- ate EA) Re browni sp. n. First two visible sternites with black spicules, male connexivum anteriorly flattened, distally re- flexed (fig. 39). Female connexivum reflexed over res 5-7 (fig 47) cecconscconcecooaoes amnicus sp. n. Males 2.88-3.48 mm long, male 7th sternite de- pressed (fig. 34). Paramere truncate (fig. 34). Female connexivum sinuate, distally convergent (MIVD priori sp. n. Males 3.48-3.76 mm long, male 7th sternite evenly rounded (fig. 18). Paramere reniform (fig. 16). Female connexivum not convergent distally. pirata papuensis Lundblad Proepisternum, pro-meso and metasternum with black spiculest tt Sie e 8 Proepisternum, pro-meso and metasternum lack- ing black spicules Pronotum anteriorly transversely orange, not reaching inner margin of eyes. Eye length 1.5x median pronotal length ............... biroi Lundblad Pronotum anteriorly transversely orange, surpass- ing inner eye margin. Eye length subequal to me- dianspronorall Ener hirsuta sp. n. Hind femora of both sexes lacking prominent projections, not incrassate (figs. 141, 148). Body size sexually dimorphic (males 2.32-2.52 mm long, females 3.28-3.36 mm long. ..caesius sp. n. Hind femora of both sexes variably incrassate with one or more prominent ventral projections. Males and females not sexually dimorphic ... .10 . Dark areas of dorsum with prominent fields of adpressedheoldispieulesen ne re 11 Dark areas of dorsum lacking noticeable fields of adpressed soldispicul Sp een 12 Figs. 1-11. Rhagovelia australica Kirkaldy. — 1, male dorsal habitus; 2, ibid female; 3, male external genitalia; 4, ibid female; 5-7 male, 8-10 female: 5, 9, hind leg; 6, 8, front leg; 7, 10, antennae; 11, male genital capsule. Figs. 1-4, scale line 1 mm, figs. 5-10 scale line 1 mm, fig. 11 scale line 0.5 mm. 24 LANsBURY: Papuan Rhagovelia it U) 25 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 11. Male front tibia not explanate medianly, inner margin uniformly fringed with short hairs (fig. 102). Female connexivum not sinuate (fig. 95)... II A arn treat aureospicata sp. n. — Male front tibia explanate medianly, inner mar- gin with a cluster of hairs/spines (fig. 119). Female connexivum sinuate (fig. 118) ................ D tte Net a RE MS ce crinita sp. n. 12. Anterior pronotal margin transversely pale yellow (figs. 106, 107). Male front tibia not explanate, inner margin fringed with short hairs (fig. 109). Female connexivum converging at 7th tergite (Hp 107) ee herzogensis sp. n. — Anterior pronotal margin transversely pale yellow not reaching lateral margins (figs. 126, 132). Male front tibia medianly explanate, inner mar- gin with a cluster of hairs/spines (fig. 129). Female connexivum sinuate, distally fringed with Venyglomestiains| (hel 2)E rar. thysanotos sp. n. SYSTEMATICS Rhagovelia australica Kirkaldy (figs. 1-11, tabs. 1-2) Rhagovelia australica Kirkaldy, 1908: 783-784; Hale 1925: 5; Hale 1926: 203; Lundblad 1933: 286; Lundblad 1936: 25-26; Polhemus & Polhemus 1988: 165. Material. — Australia: Queensland, Mt. Lewis NW of Rumula, 1200 m., 6-15.vi.1966, G.A. & S.L. Samuelson (BPBM) 32 6, 16 £ and 15 immature, all apterous. Redescription Adult apterous. — Males (fig. 1) 4-4.2 mm long, maximum width 1.6-1.72 mm, females (fig. 2) 4.04- 4.4 mm long, maximum width 1.68-1.88 mm. Coloration. — Male (fig. 1): dorsum bright yellow and dark reddish brown-black. Head yellow to dark reddish brown, basally almost invariably yellow. Pronotum anteriorly narrowly black, mesially with a broad dark brown-black transverse bar, usually nar- rowly divided by a yellowish brown longitudinal stripe. Anterior and posterior margins with scattered brown pits. Adjacent to posterior pronotal margin, a narrow black shining bar which may be the mesono- tum. Metanotum yellow, mesially reddish brown- black, posterior margin black. Tergites 1-4 either uni- formly black or with varying areas reddish brown. Tergites 5-7 yellowish brown, marginally variably black. Eigth tergite black. Connexivum bright yellow with margins dark reddish brown. Jugum, pro, meso and metaepisterna, coxae, trochanters and part of fe- Table 1. Proportion of antennal segments of Rhagovelia australica Kirkaldy. I Il II IV Apterous male 60 35 37 29) Apterous female 60 36 40 DIS Table 2. Proportions of leg segments of Rhagovelia australi- ca Kirkaldy. Femur Tibia Tarsus I Tarsus II Tarsus III d front leg 665) 6) 55 16 = 9 front leg 71 71 35 16 = 3 middle leg 113 86 6 37.5 45.5 2 middle leg 111 90 5 40 49 3 hind leg 88 96 4 8.5 18 2 hind leg 97 101 4 5 20 mora pale yellowish brown. Metasternum and 2nd sternite black, sternites 3-8 reddish brown, distally pale yellow. Pleural margins of sternites narrowly margined black. Front femora (fig. 6) distally and middle legs dark brown-black. Hind leg (fig. 5) broadly dark reddish brown-black. Antennae (fig. 7) dark reddish brown, Ist segment basally pale yellow, ventrally more broadly pale. Structure. — Male: Median pronotal length 2.23- 2.4x eye length. Pronotal width 1.35-1.44x median length, posteriorly covering all but a narrow band of ‘mesonotum and between 6-9x length of metano- tum. Tergites 1-6 subequal, 7th longer, 8th shorther imam Jins 1-3 are 12.5 ¢ 152 De 128 2 12,1 es 12,11 = 13.7 : 22.7 : 18.5. Distal margins of 5-7 fringed with short hairs, 8th pilose, genitalia distinctly hairy (fig. 3). Connexivum slightly raised, distal segment fringed with short conspicuous hairs. Jugum clearly distinct from antenniferous tubercles. Propleura, pro- episternum and mesopleura without fields of minute black spicules. Distal margins of propleura with a row of black punctures, lateral margins with sparse semi- erect hairs, upper margins of mesopleura with a field of semi-erect brown hairs. Front femora with a vari- able number of fine black spines along ventral mar- gin, tibia sinuate. Hind trochanter with four or more blunt nodules. Femora variably incrassate, proximal ventral margin with a row of even-sized setae, medial- ly with prominent acuminate and blunt projections, distally becoming smaller. Inner margin of tibia with a row of irregular spaced triangular setae. Male geni- talia (fig. 11) capsule distally produced. Figs. 12-25. Rhagovelia papuensis Lundblad. — 12-18, 23-24 male, 19-22, 25 female. 12, 13, 19, hind leg; 14, 20, front leg; 15, mid-femora; 16, genital capsule; 17, 22, antennae; 18, 21, external genitalia; 23, male macropterous; 24, male apterous; 25, female apterous. Figs. 12-15, 17-22 scale line 1 mm, fig. 16 scale line 0.5 mm, figs. 23-25 scale line 1 mm. 26 LANSBURY: Papuan Rhagovelia 27 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Coloration. — Female (fig. 2): Tending to resemble male, dark brown-black pigmentation of tergites sometimes more extensive than in males. Front femo- ra (fig. 8) anterior dorsal third pale yellow, ventrally pale coloration extending about two thirds femoral length. Coxae and trochanters of all legs pale yellow- ish brown. Structure. — Female: Median pronotal length 2.27- 2.52x eye length. Pronotal width 1.37-1.46x median length, posteriorly covering all but a narrow band of ‘mesonotum’. Pronotum at midline greatly exceeding length of ‘mesonotum’ and between 8-10x length of metanotum. Tergites 1-4 more or less subequal, 5-7 progressively slightly longer, 8th shorter than pro- ceeding. Seventh tergite slightly depressed, proctiger reaching beyond Ist gonocoxa (fig. 4). 1-8 are 12.6: 12,8 2 12,7 : 13352 1593 18:12 20.1 8 15.1, Nerziess 2-8 with conspicuous slightly flattened shining areas. Pronotum, metanotum and tergites 1-6 with ad- pressed dark brown hairs, especially noticeable on pale areas of metanotum. Eighth tergite laterally pi- lose. Connexivum anteriorly slightly reflexed out- wards, converging and becoming more erect and pi- lose posteriorly, upper margins shining black. Venter similar to male. Front femora (fig. 8) with a variable number of fine spines ventrally. Tibia slightly sinuate. Hind femora (fig. 9) moderately incrassate, spinose distad ventrally. Tibia straight, not spinose along in- ner margin. Antennae (fig. 10) Ist segment curved, remaining segments straight. Remarks. — Described from Queensland, Kuranda, apterous female type in BPBM. Hale (1925) lists Malanda, Lundblad (1936) briefly commented on the macropterous form from Kuranda. This species has so far only been recorded from the Atherton Tablelands region near Cairns, Queensland. The apterous form is immediately recognisable by its striking dorsal bright yellow-brown-black pattern which is rather variable, especially the spread of the dark pigmentation. The male genital capsule is distal- ly turned upwards, parameres elongate almost par- allel-sided (fig. 11). Female with two (1+1) clusters of black hairs along the distal margin of the 8th seg- ment. Rhagovelia papuensis Lundblad (figs. 12-25, tabs. 3-4) Rhagovelia papuensis Lundblad, 1936: 28-30; Polhemus & Polhemus 1988: 170-173. Material. — Papua New Guinea: West Britain: Tamari Creek, 300 m., 14.vii.1989, R. N. B. Prior, 126, 129 ap- terous, 12 macropterous; Von River, Banaule, torrential stream, 21.iii.1990, I. Lansbury 1 ® apterous; Bialla, Balima River, 3.111.1989, in dense shade, R.N.B.P. 83, 132 apter- 28 Table 3. Proportions of antennal segments of Rhagovelia papuensis Lundblad. I II III IV Apterous male 42 26 27 26.5 Apterous female 41 29 31 31 Macropterous male 42.5 25 30 27 Macropterous female 43 26 30 26 Table 4 Proportions of leg segments of Rhagovelia papuensis Lundblad. Femur Tibia Tarsus I Tarsus II Tarsus III Apterous d frontleg 47 49 3.5 13.5 - 2 frontleg 48 50 3 14 = 3 middle leg 77 55 3 23 36 9 middle leg 81 57 4 23 36 3 3 3 hind leg 70 66 ® hind leg 70 71 6 16 Macropterous 3 frontleg 50 54 3 12.5 = 9 frontleg 51 52 4 15 = 3 middle leg 83 60 3 22 38 9 middle leg 81 55 4 23 39 d hind leg 76 72, 3 7 16.5 2 molles 70. 70 3 7 16 ous, 1d 19 macropterous. Redescription Adult apterous. — Males (fig. 24) 3.32-3.48 mm long, maximum width 1.2-1.28 mm, females (fig. 25) 3.48-3.76 mm long, maximum width 1.2-1.4 mm. Adult macropterous. Male (fig. 23) 4.12 mm, maxi- mum width 1.52 mm, female 3.88 mm long, maxi- mum width 1.48 mm. Coloration. — Male (fig. 24): Dorsum head, most of pronotum, meso, metanotum and basal tergites black. Remainder of tergites dark brown, 7th tergite basally faintly shining. Anterior margin of pronotum broadly orange and tending to widen laterally. Propleura faintly bluish black over orange coloration. Posterior pronotal margin narrowly orange. Con- nexivum broadly margined pale brown. Mesopleura and metapleura bluish black evanescent. Metaste- rnum and sternites 2-6 laterally black, ventrally pitchy red, 7th sternite pale yellow. Coxae and tro- chanters of all legs pale yellow. Front femora (fig. 14) pale yellow, apically dark brown, tibia and tarsi of front leg and all middle leg greenish black iridescent. Hind femora (fig. 12) dark brown-black dorsally, ventrally pale yellow, tibia and tarsi dark brown- black. Antennae (fig. 17) basal segment broadly pale yellow, remaining segments black. Structure. — Male: Median pronotal length just over 2x eye length and covering most of ‘mesono- Figs. 26-27. Rhagovelia spp. dorsal habitus. — 26-27 priori sp. n., 26, male apterous; 27, female apterous. Scale as for fig. 25. tum’. Pronotal width 1.4x median eye length. Tergites 1-5 subequal, 6th slightly longer, 7th longer and shining, 8th shorter:10 : 11:12:11:11:13:18: 14.5. Tergites and connexivum with adpressed brown hairs, denser and longer along margins of 6th tergite and tip of connexivum. Genital capsule (fig. 16). Jugum barely separated from antenniferous tubercles. Propleura, proepisternum, meso and metasternum and jugum with fields of black spicules. Ventrites 2-4 carinate, laterally sternites bare, 6th with scattered brown adpressed hairs. Front femora (fig. 14) finely spinose, middle femora (fig. 15) dorso-ventrally flat- tened, finely spinose. Hind femora (fig. 12) ventrad- distally spinose, tibia sinuate inner margin distally with prominent triangular projections. Coloration. — Female (fig.25): Head and thoracic coloration similar to male. Connexivum more broad- ly margined pale yellowish brown. Antennae, front and middle legs similar to male, hind leg (fig. 19) dark brown on all surfaces. Structure. — Female: Black denticules on propleu- ra, pro-meso and metasternum more prominent. Tergites 1-3 reflexed downwards, 1-4 pilose, 5-8 with large shining areas, 10): 10:9): 11 : 14:15:16: 13. Proctiger truncate and slightly depressed (fig. 21). Connexivum partially raised converging along mar- gins of tergites 1-3, more elevated distally and con- verging at tergite 8. Ventrites with fields of shining LANSBURY: Papuan Rhagovelia semi-erect hairs covering median areas of sternites 2- 3 gradually decreasing in length and density distally, lateral margins of sternites with similar hairs increas- ing in length and density distally. First visible sternite obtusely raised. Hind femora moderately incrassate with large spine ventrad, tibia straight, not spinose along inner margin. Macropterous male (fig. 23). — Similar to apterous male, pronotum lengthened caudad, posterior margin orange yellow. Wings extending well beyond the end of abdomen with four closed cells, two in distal half. Wings brown, venation slightly darker. Hind femur (fig. 13) not as incrassate as apterous male. Macropterous female. — Similar to male. Venation more conspicuous especially subcosta and radius, in- ner margins near anal vein much paler. Remarks. — Described from Papua New Guinea, Astrolabe Bay, (Stephensort and Erima) and Ralum in New Britain. Type series in HNHM. The enlarged pronotum, ventral conspicuous fields of black spic- ules and shape of the male parameres shows a close re- lationship with species from the Solomon Islands. The female connexivum is more or less elevated its entire length (fig. 25) and the shining distal tergites are visible. Rhagovelia papuensis males are consistently smaller than À fulvus sp. n. and browni sp. n. (Solomon Islands), papuensis is superficially similar to priori sp. n. Rhagovelia priori sp. n. (figs. 26-38, tabs. 5-6) Type material. — Holotype male: Papua New Guinea, Oro Province, Popondetta, Ambogo Plantation, small stream, 29.iv.1989, R. N. B. Prior (OXUM). Paratypes 36, 149, same data; 128 99 P. N. G., Oro Prov. Popondetta, Iseveni, small stream, 2.v.1989, R. N. B. P.; 12 P. N. G. Oro Prov. Popondetta, Sambogo River, 30.iv.1989, R. N. B. P. (OXUM). Description Adult apterous. — Males (fig. 26) 2.88-3.48 mm long, maximum width 1-1.6 mm, females (fig. 27) 3.2-3.48 mm long, maximum width 1.2-1.24 mm. Coloration. — Male (fig. 26): Head most of thorax dorsally black. Tergites greyish black, 7th tergite and lateral margins of connexivum shining pale brown. Anterior margin of pronotum pale yellow, laterally rather wider. Propleura dark grey merging into red- dish brown. Meso and metapleura greyish black. Pro, meso and metasternum and sternites 2-3 bluish grey, coxal cavities tending to be tinged reddish brown. Sternites 3-5 reddish brown, 6th paler. Coxae and trochanters pale yellow. Front femora with extensive pale yellow coloration especially along inner margin. Remainder of front and all middle legs dark shining 29 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 brown. Hind femur shining dark brown with lateral pale areas same colour as trochanter along ventral margin. Antennae (fig. 33) 1st segment basally pale yellow, ventrally surface more broadly yellow, re- maining segments black. Structure. — Male: Median pronotal length 2x eye length and covering almost all mesonotum. Pronotal width 1.44x median length and 5-6x length of meta- notum. Tergites 1-5 subequal, 6 and 8 slightly long- er, 7th longest with a variably shaped shining area; 9 : OPO O EDS ERD Nhoracıerdorsum covered with fine golden hairs. Shining lateral con- nexival margins and tergites 1-6 with longer ad- pressed hairs. Connexivum either flat or slightly raised distally with tufts of hairs. Jugum very close to antenniferous tubercles. Propleura, pro, metaepister- num and jugum with fields of minute black spicules. Sternites anteriorly with a median keel with pale yel- low hairs overlying fine adpressed hairs. Seventh ster- nite depressed posteriorly, lateral margins obtusely carinate and pilose, 8th with a conspicuous keel (fig. 34). Front leg (fig. 30) mid femora ventrally with two rows of fine hairs (fig. 31). Hind femora variably in- crassate (figs. 28, 29). Ventral margin variably spinose Table 5. Proportions of antennal segments of Rhagovelia priori sp. n. I II III IV Apterous male 371 22 26 26 Apterous female 37 DI 24 24 Table 6. Proportions of leg segments of Rhagovelia priori sp. n. Apterous Femur Tibia Tarsus I Tarsus II Tarsus III 3 frontleg 40 45 3 11 = 9 frontleg 41 44 3 12 = 3 middle leg 71 53 4 Dil 33 9 middle leg 72 Si, A 27 35 d hindleg 67 COS 5 14 ® hindleg 59 60 2 5 14 with many fine setae, distally with triangular setae. Tibia either straight or slightly sinuate with triangular setae along inner margin, length increasing distally. Genital capsule (figs. 32, 33), distal lateral aspect of 7-8th segments (fig. 34). Figs. 28-38. Rhagovelia priori sp. n. — 28-35 male, 36-38 female. 28, 29, 36 hind leg; 30, front leg; 31, mid-femora; 32, 33, genitalia; 34, 37, external genitalia; 35, 38, antennae. Figs. 28-31, 34-38 scale line 1 mm, figs. 32-33 scale line 0.5 mm. 30 Coloration. — Female (fig. 27): Head and pronotal coloration similar to male. Mesonotum postero-late- rad sometimes with brown shining areas. Tergites 5-7 sometimes shining tending to resemble À. papuensis (fig. 25). Upper margins of connexivum narrowly shining brown. Legs and antennae similar to male oth- er than hind femora which tend to be paler ventrally. Structure. — Female: Median pronotal length about 2x eye length. Mesonotal length variable, pronotum about 5-7x length of mesonotum. Metanotum slight- ly longer than mesonotum. Tergites 1-4 subequal, 5- 7 progressively longer, 8th short; 1-8 are: 10: 10: 10 : 10: 12:17:20: 8. Connexivum anteriorly reflexed outwards with fields of hairs on inner margins. From 4th tergite abruptly turned inwards and converging distally. Upper distal margins pilose and 8th tergite densely pilose. Underside of thorax similar to male. Anterior sternites with clusters of golden hairs me- dianly. Proctiger reflexed downwards (fig. 37). Hind femur (fig. 36) moderately incrassate with a promi- nent curved spine in distal third. Hind tibia straight without prominent setae along inner margin. Antennae (fig. 38) first segment slightly more curved than male (fig. 33). Etymology. — A noun in apposition. This species is dedicated to Mr. R. N. B. Prior who has collected aquatic Hemiptera-Heteroptera from various local- ities in Papua New Guinea, especially West New Britain. Remarks. — Rhagovelia priori sp. n. is superficially much like papuensis. The apterous females of former are easily distinguished by stronger convergent con- nexiva with fields of hairs along anterior inner mar- gins and prominent distal clusters of hairs. Males of priori tend to be smaller than papuensis (2.88-3.48 : 3.32-3.48 mm). Genital capsule of priori distally acu- minate, of papuensis distally convergent and rounded. Parameres of priori slightly sinuate, of papuensis renif- orm. Rhagovelia amnicus sp. n. (figs. 39-50, tabs. 7-8) Type material. — Holotype male: Solomon Islands, Malaita, Raiako Nafinua, in forest stream c 3 yards wide, much shaded, fast running with shingle bottom, 27.v.1955, E. S. Brown (BMNH). — Paratypes: same data as holotype, 186 409; same data, except ‘larger stream with a few slow flowing areas’, 3d 72 (BMNH, OXUM). Additional material. — Solomon Islands: Malaita: River Aluta just west of Maelegwasu, in forest. A rather large rap- id river with shingle bottom, 26.v.1955, E. S. B., 136 122 apterous, 1d 19 macropterous (BMNH, OXUM); same, in- land east of Kwailasi. Small running stream shaded in forest, 25.v.1955, E. S. B., 196 132 (BMNH, OXUM); same, near Kwailasi, large rocky stream, 25.v.1955, E. S. B., 106 62 (- BMNH, OXUM). LansBury: Papuan Rhagovelia Description Adult apterous. — Males (fig. 39) 3.32-3.96 mm long, maximum width 1.16-1.44 mm, females (fig. 47) 3.16-3.84 mm long, maximum width 1.22-1.44 mm. Adult macropterous. Male (fig. 46) 4.12 mm long, maximum width 1.58 mm, female 4.12 mm long, maximum width 1.68 mm. Coloration. — Male (fig. 39): Ground colour black, anterior pronotal margin pale yellow, not reaching lateral margins. Tergites dark brown-black, 7th ter- gite shining. Connexivum narrowly shining pale brown anteriorly. Legs mostly black. Coxae, tro- chanters, proximal margins of front tibia and inner proximal margin of-hind femur pale yellow. Jugum brown, pro-meso and metasternum black. Sternites dark reddish brown, laterally black. Antennae black, proximal 0.2-0.25 of first segment pale yellow. Structure. — Male: Median pronotal length 2.2x eye length, posteriorly converging most of mesono- tum. Pronotal width 1.47x median length. Metanotum 2x length of mesonotum. First tergite shorter than remainder, tergites 2-6 subequal, 7th longest, 8th shorter and densely hairy. 1-8 are 10: 14: SENS EME SE 2507 Wereitese2-o1and! basal margin of connexivum with many pale hairs, 7th sparsely pilose, distal margin fringed with longer hairs. Connexivum anteriorly slightly flattened, be- coming more reflexed upwards distally. Anterior lat- eral pronotal margins, lateral margins metanotum, and first two tergites with fields of black denticles. Jugum barely separated from antenniferous tubercles. Propleura, proepisternum and mesopleura with a sin- gle row of deep black punctures, upper margins with prominent erect hairs. Meso and metasternum with scattered black denticles, overlying them long semi- erect fine yellowish hairs. First two sternites with black denticles. Anterior sternite bluntly acuminate. Anterior sternites with long fine hairs, distal sternites with short hairs which increase in density distally. Seventh sternite depressed posteriorly, lateral margins with tufts of hairs (fig. 39). Genital segment ventrally carinate with a narrow fringe of fine hairs, laterally smooth and shining (fig. 40) parameres different as- pects (figs. 44-45). Front femora with a variable num- ber of fine spines along ventral margin, tibia straight (fig. 41). Hind trochanter with circa 5 blunt nodules. Femur incrassate, proximal ventral margin with a row of even-sized setae, medially with a prominent spine, distally with irregular smaller spines in two rows. Tibia sinuate, inner margin armed with triangular se- tae, distally with a prominent bifurcate projection (fig. 42). Coloration. — Female (fig. 47): Resembling male, pronotal anterior yellow band more intense. Tergites 5-8 with shining elongate areas. Propleura dark yel- lowish brown. External connexival margins anteriorly 31 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 39-50. Rhagovelia amnicus sp. n. — 39-46 male, 47-50 female. 39, 47, dorsal habitus; 40, genital capsule; 41, tront leg; 42, 49, hind leg; 43, 50, antennae; 44, 45, parameres; 46, macropterous dorsal habitus; 48, external genitalia. Figs. 39-43, 46, 47, 49-50 scale line 1 mm, figs. 44-45 scale line 0.1 mm, fig. 48 scale line 0.5 mm. broadly yellow, upper margins shining brown, distal- Structure. — Female: Median pronotal length 2.5x ly dark brown. Sternites covered with bluish silvery eye length, pronotal width 1.4x median length, poste- pubescence. Legs and antennae similar to male, but riorly covering most of mesonotum, the latter 0.1 1x front tibia are more extensively yellow. pronotal length. Metanotum 2.2x length of mesono- 32 Table 7. Proportions of antennal segments of Rhagovelia amnicus sp. n. I II Il IV Apterous male 43 28.5 29 27 Apterous female 43 26 27.5 25 Macropterous male 42 26 28 Dif Macropterous female 43 26 28 26 Table 8 Proportions of leg segments of Rhagovelia amnicus sp. n. Femur Tibia Tarsus Tarsus Tarsus Apterous d front leg 53 53 3 13 = 2 front leg 49 49 4 14 = 3 middle leg 88 63 5 26 38 9 middle leg 79 57 4 25 37 d hind leg 81 64 4 5) 16 9 hind leg 70 65 DIS) 6 16 Macropterous d front leg 52 56 4 ? front leg 54 57 5} 3 middle leg 90 65 >) DS) 40 9 middle leg 85 66 5 3 hind leg 89 70 4 9 hind leg 75 75 3 tum. Lateral margins of pronotum, meso and meta- notum with fields of black denticles. Tergites 1-3 de- pressed, tergites 4-7 darker than preceeding, tergites 5-7 mostly shining. Tergites 1-2 with scattered long black semi-erect hairs. 1-8 are: 11:9:8.5:12:11: 13 : 26 : 10. Connexivum anteriorly raised almost erect, thickened and shining and sharply converging along 3rd tergal margin, curving and strongly reflexed over tergites 5-7, sometimes posteriorly completely covering distal tergites. Upper distal margins densely hairy, posteriorly with a prominent tuft of hairs, proctiger reflexed downwards (fig. 48). Lower margin of jugum, proepisternum and mesosternum with fields of black denticles. Sternites faintly rugose, 7th shining, laterally and distally fine yellowish pilose. Hind femur moderately incrassate, ventrally spinose, tibia not armed along inner margin (fig. 49). Macropterous male (fig. 46). — Pronotum at widest subequal to median length. Caudad depressed, hu- meral angles slightly elevated. Forewings brown, ve- nation black with four cells. Membrane of elytra just surpassing genital segment. Hind femur incrassate. Macropterous female. — Similar to male, mem- brane rather longer than in male exceeding proctiger and 8th segment. Etymology. — Name derived from ‘amnis’ stream of water, amnicus ‘inhabitant of stream’. LANSBURY: Papuan Rhagovelia Remarks. — This species is similar to À priori; males differing by not having lateral margins of 7th sternite laterally obtusely carinate. Genital capsule truncate distally. Females have connexivum and distal abdominal segments are much more pilose. The ante- rior pronotal yellow band not usually reaching lateral margins. Rhagovelia browni sp. n. (figs. 51-62, tabs. 9-10) Type material. — Holotype d: Solomon Islands, Guadacanal, Sorvohio River near Gold Ridge, rapid water over boulders with calm pools, small shaded river, 20.11.1955, E. S. Brown (BMNH). — Paratypes: same data as holotype, 64, 62 apterous; same data except, Gold Ridge, 1000-1500 feet, clean fast running water with boulders mainly shaded by forest, 25.vi.1956, E. S. B., 174 69 ap- terous, 14 29 macropterous; same data as preceeding, 30.vi.1956, E. S. B., 36 52 (BMNH, OXUM). Additional material. — Solomon Islands, Guadacanal: Suta, Sutakiki River, c. 2300 feet, 26.vi.1956, 46 99 apter- ous, 1 2 macropterous; Sangava River near Suta. Strong sto- ny trickling stream, mostly rather shaded, 27.vi.1956, 4d 5; Kokumbona River 13.v.1956, 148 59 apterous, 1d macropterous; Tsarivonga River, rapid stony river with quiet backwaters, 1000 feet, 22.iii.1955, 198 139 apter- ous, 1d 19 macropterous; Tenaru River. Between Mission and Tenaru Falls, mostly from comparatively calm water along margin and in backwaters, 7.xi.1954, 4d 42 apter- ous, 26 12 macropterous; Matanikao, pools in rocks downstrem of Matanikao Caves, shaded 31.x.1954, 154 6% apterous; Honiara, Matanikoa River, 6d 29 apterous, 16 macropterous (all leg. E. S. Brown) (BMNH, OXUM); Guadacanal, Gold Ridge 600 m. 22.vi.1956, J.L. Gressitt, 1d apterous; Santa Ysabel, Tamatahi-Molao Maringe Distr. 28.vi.1960, Jungle track C.W. O’Brien, 18 49 apterous, 2% macropterous (BPBM). Description Adult apterous. — Male (fig. 51) 3.88-4.12 mm long, maximum width 1.36-1.44 mm, females (fig. 59) 4.12-4.24 mm long, maximum width 1.48-1.52 mm. Adult macropterous. Males (fig. 52) 4.64 mm long, maximum width 1.72 mm, females 4.52-4.8 mm long, maximum width 1.72-1.76 mm. Coloration. — Male (fig. 51): General dorsal color- ation dark brown-black. Head black, anterior margin of pronotum yellow, lateral margins bluish black or reddish brown. Seventh tergite and genital segment shining black. Jugum, coxae and trochanters yellow- ish brown. Sternites dark reddish brown, 7-8th ven- trites pale yellow. Pro, meso and metapleura reddish brown. Lateral margins of sternites black. Antennae black, basal fifth of lst segment pale yellow, dorsally dark brown. Middle and hind legs dark brown-black. Structure. — Male: Median pronotal length 2.36x eye length. Pronotal width 1.44x median length, pos- teriorly covering almost all mesonotum, metanotum 0.2x length of pronotum. Tergites 1-6 more or less 33 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 34 Table 9 Proportions of antennal segments of Rhagovelia browni sp. n. I Il II IV Apterous male 52 31 35 30 Apterous female 47.5 28 32 28 Macropterous male 44 ITS 30 26 Macropterous female 48 30 32 30 Table 10. Proportions of leg segments of Rhagovelia browni sp. n. Femur Tibia Tarsus I Tarsus II Tarsus III Apterous d front leg 60 67 4 16.5 — ironies DISS 5 18 - 3 middle leg 105 76 5 31 42 9 middle leg 91 67 5 28 41 3 hind leg OSG 4 8.5 20 9 hind leg 82 83 4 8 20 Macropterous d front leg I Dee 19:5 = 2 front leg SMS 5 17 = d middle leg 88 64 5 DI 40 9 middle leg 92 70 5 28 4] 3 hind leg DT Te 7 19 9 hind leg 84 86 4 9 20 subequal, dull, lateral margins finely pilose and distal margins fringed with hairs. Seventh longest, shining, transversely rugose, posterior margin fringed with fine hairs. Genital segment (8th) shining, covered with fine hairs, denser along lateral margins. 1-8 are 13:13:14.5:14:15:12: 23 : 20. Anterior lateral margins of pronotum, meso and metanotum with fields of black spicules. Connexivum either slightly reflexed outwards or almost erect, upper margins shining dark brown. Propleura anteriorly with a scat- tered field of black spicules, posterior margin with several short continguous rows of punctures. Meso and metapleura with short black hairs directed cau- dad. Jugum almost touching antenniferous tubercles. Lateral margins of jugum, proepisternum, meso and metasternum with prominent fields of black spicules. Sternites 2-5 with scattered black spicules, 2nd ven- trite obtusely carinate. Seventh sternite depressed to- wards gentital segment. Latero-distal margins pilose. Genital segment carinate and fringed with short hairs. Genital capsule (fig. 53) parameres different aspects (- LANSBURY: Papuan Rhagovelia figs. 56-57). Front leg (fig. 54). Hind leg (fig. 55) fe- mur incrassate, trochanter with 3-4 blunt nodules. Femoral ventral margin anteriorly with two rows of uniform sized setae, posteriorly with larger spinose projections. Tibia sinuate with two rows of blunt spines, medially with a large spine, distally with a large spine. Coloration. — Female (fig. 59): Resembling male, pronotum black other than yellowish brown anterior margin reaching inner margin of eyes. Tergites 5-8 shining. Upper margins of connexivum shining red- dish brown. Pro-meso and metapleura varying between dark reddish brown and bluish black. Jugum, coxae and trochanters pale yellowish brown. Meso and metaepisternum and sternites 2-6 dark red- dish brown, lateral margins dark brown-black, 7th sternite shining pale reddish brown. Antennae black, Ist segment basally and ventrally pale yellow. Front femur ventrally pale yellow, tibia black. Middle leg dark reddish brown. Hind leg (fig. 61) femur proxi- mally pale yellow, remainder of leg reddish brown. Structure. — Female: Median pronotal length 2.4x eye length. Pronotal width 1.5x median length, pos- teriorly covering all but a narrow strip of mesonotum. Metanotum 2x length of mesonotum. Tergites 1-3 subequal, 4-7 increasing in length, 8th shorter. 1-8 TR RS oS) o ASIN 5) 2 OE SOS hinne areas of tergites 5-8 increasing in size distally and faintly transversely rugose. Distal margin of 8th ter- gite fringed with long hairs. Anteriorly connexivum reflexed outwards, along margins of tergites 3-4 more erect, distally slightly reflexed outwards forming a sin- uate line, connexivum distally pilose. Lateral margins of mesonotum with scattered black spicules, metano- tum with larger fields. Propleura and thorax ventrally similar to male. Sternites with black spicules, not car- inate. Seventh sternite shining, laterally with fine yel- lowish hairs. Genital segment more pilose, connexival hairs almost reaching end of genital segment (fig. 62). Hind femur moderately incrassate, spinose posterior- ly, tibia straight, inner margin not spinose. Macropterous male (fig. 52.). — Pronotum anteri- orly yellow, lateral margins bluish black. Pronotal width subequal to median length, humeral angles and posterior tip depressed with scattered black punc- tures, pronotal disc raised. Forewings brown, vena- tion black with four closed cells, anteriorly inner mar- gin of anal vein narrowly pale yellow. Membrane of elytra surpassing end of abdomen and genitalia. Coloration of legs and antennae similar to apterous Figs. 51-62. Rhagovelia browni sp. n. — 51-58 male, 59-62 female. 51, dorsal habitus; 52, dorsal habitus macropterous; 53, genital capsule; 54, front leg; 55, hind leg apterous; 56-57, parameres; 58, antenna; 59, dorsal habitus apterous; 60, hind leg macropterous; 61, hind leg apterous; 62, external genitalia. Figs. 51-55, 58-61 scale line 1 mm, figs. 56-57 scale line 0.1 mm, fig. 62 scale line 0.5 mm. 55 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 form. Hind leg (fig. 60) femur moderately incrassate, tibia straight, inner margin with triangular setae. Macropterous female. — Similar to macropterous male, membrane slightly shorter just surpassing the Ist gonocoxa. Etymology. — This species is dedicated to the late E. S. Brown, an enthusiastic hemipterist who collect- ed extensively in the Solomon Islands. Remarks. - Comparing females of À. browni sp. n. it is rather similar to Rhagovelia fulvus sp. n., browni has a sinuate connexivum whereas the connexivum of fulvus is more or less straight. Rhagovelia browni is a (ANN AAN A | much darker species; parameres elongate compared with the moderately reniform parameres of fulvus. Rhagovelia fulvus sp. n. (figs. 63-72, tabs. 11-12) Type material. — Holotype d : Solomon Islands, Malaita, Fulisano-Maelegwasu, 26.v.1955. Small spring near its source, just west of main water-shed of island, E. S. Brown (BMNH). — Paratypes: same data as holotype, 76 5 (BMNH, OXUM). Description Adult apterous. — Male (fig. 63) 3.64-3.8 mm long, Figs. 63-72. Rhagovelia fulvus sp. n. — 63-68 male, 69-72 female. 63, 69, dorsal habitus apterous; 64, genital capsule; 65 par- amere; 66, front leg; 67, 70, hind leg; 68, 71, antennae; 72, external genitalia. Figs. 63, 66-71 scale line 1 mm, fig. 64, 72, scale line 0.5 mm, fig. 65 scale line 0.1 mm. 36 Table 11. Proportions of antennal segments of Rhagovelia fulvus sp. n. I Il III IV Apterous male 50 29.5 33 29 Apterous female 50 31 32.5 29 Table 12. Proportions of leg segments of Rhagovelia fulvus sp. n. Femur Tibia Tarsus I Tarsus II Tarsus III Apterous 6 frontleg 55 63 3 16 - 9 frontleg 55 58 3.5 15 = 3 middle leg 94 70 5 26 39 2 middle leg 90 64 5 26 40 3 hind leg 92 85 3 8 19 Sihindiles 79786 3 9 16 maximum width 1.28-1.4 mm, female (fig. 69) 3.76- 4.04 mm long, maximum width 1.36-1.44 mm. Coloration. — Male (fig. 63): Head dorsally dark reddish brown. Pronotum either pale orange yellow or pinkish brown. Anterior lateral and medianly rath- er paler. Meso, metanotum and tergites 1-6 varying between dark yellowish and reddish brown. Connexivum and tergites 7-8 pale yellow. Propleura pale reddish brown, meso and metapleura slightly darker. Sternites dark reddish brown, 7th sternite faintly shining, 7th and 8th sternites pale yellowish brown. Coxae and trochanter pale yellow. Femur and tibia of all legs pale yellow, tarsi tending to be dark yellowish brown. First and 2nd antennal segments pale yellow, remainder brown. Structure. — Male: Median pronotal length 2.56x eye length, covering almost all mesonotum. Pronotal width 1.46x median length. Anteriorly pronotum with a transverse row of punctures, lateral margins with a small field of black spicules, sometimes spic- ules much paler and difficult to see. Metanotum about 2x length of mesonotum. Tergites gradually in- creasing in length distally, 1-8 are 10 : 13:12:12: 13 : 14:19:17. Connexivum more or less uniform- ly semi-erect. Lateral and posterior margins of prono- tum with fine silvery adpressed hairs, remainder of pronotum covered with shorter hairs. Posterior mar- gin of metanotum, tergites and lower margin of con- nexivum fringed long pale yellowish hairs. Lateral margin propleura, margins of jugum, proepisternum, meso and metasternum with fields of black spicules. Jugum barely separated from antenniferous tubercles. Sternites 2-6 with scattered black spicules and with long fine adpressed silvery hairs becoming denser on 7th sternite. Front leg (fig. 66) moderately finely spi- nose. Hind leg (fig. 67) femur incrassate, trochanter LansBury: Papuan Rhagovelia with a variable number of blunt nodules. Femur ven- tro-anteriorly with short regular setae, medially with 2 prominent spines, distally. Tibia either straight or slightly sinuate, inner margin armed with a regular row of triangular setae. Antennae (fig. 68). Genital capsule (fig. 64) paramere slightly reniform, paramere from different aspect (fig. 65). Coloration. — Female (fig. 69): Very similar to male. Pronotum orange brown, anterior margin sometimes paler. Pleura with a pinkish tinge. Tergites dark yellowish brown with straggly yellow hairs, ter- gites 5-8 with shining areas increasing in size distally. Connexivum narrowly dark brown adjacent to ter- gites, upper margins paler. Sternites laterally dark brown, ventrally paler. Apices of femur narrowly an- nulated black. First antennal segment pale yellow, segments 2-4 pale brown. Structure. — Female: Median prontal length 2.75x eye length, covering all but narrow strip of mesono- tum. Connexivum reflexed outwards, distally not reaching distal margin of 8th tergite and densely pi- lose. Pleural and ventral black spicules similar to males second sternite with a denser field of black spic- ules. Proctiger slightly raised, gonocoxal plates large (fig. 72). Hind leg (fig. 70) femur not incrassate, ven- tral margin with a prominent distal spine, tibia straight, not armed along inner margin. Antennae (fig. 71) first segment sometimes infuscated pale brown, likewise segments 2-4. Etymology. — Name refers to the tawny, reddish yellow coloration. Remarks. — Rhagovelia fulvus sp. n. is a very pale species compared with amnicus and browni. The fe- murs are almost entirely pale. The connexivum of the female is shorter than in other species not reaching the end of tergite 8. Rhagovelia biroi Lundblad (figs. 73-83, tabs. 13-14) Rhagovelia biroi Lundblad, 1936: 13-14; Polhemus & Polhemus 1988: 165. Material. — Papua New Guinea: West Britain: Bialla, Balima River, 3.iii.1989, R. N. B. Prior, 54 12 apterous; Bialla, River Inu, 10.ii.1989, R. N. B. P., 154 172 apter- ous; Bialla, River Tiaru, 10.1.1989, R. N. B. P., 106 79 ap- terous; same data, near log in river, 25.1.1989, 3d 19 apter- ous; Tamari Creek, 300 m., 14.vii.1989, R. N. B. P., 216 159 apterous; Dami near Kimbe, vii.1989, R. N. B. P., 2d apterous; Bilomi River 9.iv.1989, R. N. B. P., 7d apterous; Von River, Banaule Village near Kimbe, 21.11.1990, I. Lansbury, 174 92 apterous, 1 ? macropterous (OXUM). Redescription Adult apterous. — Male (fig. 73) 3.2-3.48 mm long, maximum width 1.16-1.28 mm, female (fig. 79) SH TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1992 Figs. 73-83. Rhagovelia biroi Lundblad. — 73-78 male, 79-83 female. 73, 79, dorsal habitus apterous; 74, genital capsule; 75, ventral aspect genital plate; 76, 82, antennae; 77, front leg; 78, 81, hind leg; 80, dorsal habitus macropterous; 83, external genitalia. Figs. 73, 76-83 scale line 1 mm, figs. 74-75 scale line 0.5 mm. 3.24-3.48 mm long, maximum width 1.2-1.56 mm. Adult macropterous. Female (fig. 80) 3.92 mm long, maximum width 1.52 mm. Coloration. — Male (fig. 73): Dorsum black, 7th tergite with a triangular iridescent greenish black patch. Pronotum anteriorly with an orange band not 38 reaching inner margin of eyes. Venter greyish black. Coxal cavaties pale yellow. Antennae black other than basal third of Ist segment. Basal third of front femur pale yellow. Trochanters of front and hind legs pale yellow, remainder of all legs greenish black iridescent, most apparent on middle and hind femora. Table 13. Proportions of antennal segments of Rhagovelia biroi sp. n. I II III IV Apterous male 43 19 21 23 Apterous female 42 22 20 22 Macropterous female 46 22 17 DD) Table 14. Proportions of leg segments of Rhagovelia biroi sp. n. Femur Tibia Tarsus] Tarsus II Tarsus III Apterous 3 frontleg 48 53 4.5 13 - 9 frontleg 48 50 2.5 13 - 3 middle leg 83 56, 2 26 34 ? middle leg 78 S20" 94 26 57 d hindleg 72 68 3 5 15 9 hindleg 72 75 2 6 16 Macropterous 9 frontleg 47 48 3 13 - ? middle leg 78 53. 1 4 26 37 9 hindleg 72 69 3 5) 16 Structure. — Male: Median eye length 1.5x median prontal length. Pronotal width 4.8x median length. Mesonotum large, 4.3x pronotal length. Metanotum short, 0.3x pronotal length. Tergites 1-7 increasing in length distally, 8th shorter. Shining area of 7th and all of 8th with long brown hairs. 1-8 are7:9:9: 10.5: 11: 12:22:15. Dorsum clothed in fine pubescence. Pronotum, meso-metanotum and Ist tergite with short recumbent black setae. Remaining tergites with longer greyish hairs. Connexivum slightly reflexed outwards, upper margin narrowly shining brown. Jugum clearly distinct from antenniferous tubercles. Proepisternum with sparse minute black spicules dif- ficult to see against background. Pro, meso and met- asternum with scattered black spicules. Lateral mar- gins of sternites 3-6 with two groups (1 + 1) of black spicules either side of midline. Second sternite with a dense area of black spicules. Venter clothed with long recumbent greyish hairs, 7th ventrite more pilose, medianly carinate with a fringe of semi-erect hairs. Front leg (fig. 77) tibia flattened. Legs with self coloured hairs and spines. Coxae with groups of curved hairs. Hind leg (fig. 78) femora with a promi- nent spine midway ventrad, further projections dis- tally. Tibia slightly curved, inner margin armed with small setae. Antennae (fig. 76) 1st segment with many spines. Genital capsule (fig. 74) parameres elongate, apically acuminate, Lundblad (1936) figure shows apex more truncate. Ventral aspect of genital plate (fig. 75). Coloration. — Female (fig. 79): Very much like the LANSBURY: Papuan Rhagovelia male. None of tergites shining. Connexivum, upper margin narrowly yellowish brown. Structure. — Female: Median eye length 1.85x pro- notal length. Pronotal width 5x median length. Mesonotum large, 4x pronotal length. Metanotum short 0.38x pronotal length. Tergites 1-7 increasing in length distally, 8th shorter, 1-8 are 7:9: 10.5: 11 > 11.5: 14:18:10. Eighth tergite depressed, posteri- or margin of 7th with a cluster of hairs. Connexivum distally produced beyond 8th tergite and densely pi- lose. Mesonotum and metanotum with minute black shining spicules overlaid by long black hairs extend- ing to Ist tergite along margins. Pro-meso and meta- sternum with scattered black spicules. Seventh ster- nite with dense areas of long golden hairs either side of midline. Hind leg (fig. 81) femora ventrad-distally with a bifurcate spinose projection, tibia straight, in- ner margin unarmed. Antennae (fig. 82). Gonocoxa and connexivum (fig. 83). Macropterous female (fig. 80). — Coloration and structure similar to apterous form. Pronotal width subequal to median length, posteriorly slightly de- pressed and transversely rugose. Pronotal margins ad- jacent to prosternum narrowly yellow. Wings dark reddish brown, membrane clearly surpassing the proctiger. Veins not prominent reaching just beyond end of connexivum. Subcostal, medial and anal veins with minute black setae; basally with two closed cells extending onto distal half. Remarks. — Described from Papua New Guinea, Astrolabe Bay, (Stephansort and Erima) and Raium in New Britain. Type series in HNHM. Both sexes of biroi are very similar, the hind femora are much alike. The projecting distal connexival segments of the fe- male are distinctive. The male parameres are long and apically acuminate unlike other Rhagovelia so far re- corded from ‘New Guinea’. The greatly reduced pro- notal length compared with the longer mesonotum distinguish biroi from species of the papuensis group (priori, amnicus and browni etc.). Rhagovelia hirsuta sp. n. (figs. 84-93, tabs. 15-16) Type material. — Holotype d: Papua New Guinea, Oro Province, Kokoda, Mambare River, Mambe Estate, 12.ix.1988, sheltered pools at side of river. R. N. B. Prior (OXUM). — Paratypes: Same data as holotype, 3d 59; East Highlands Province, Isodenu Yonki, 1700 m. 13.x.1989, Prior, 16 12 (OXUM). Additional material. — Papua New Guinea, Oro Province, Popondetta, Iseveni, small stream, 2.v.1989, Prior, 1d (OX- UM); ‘Papua, Mafulu, 4000 feet, December 1933, L.E. Cheesman’, 38 32 (BMNH). Description Adult apterous. — Male (fig. 84) 3.92-4.04 mm 59 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 84-93. Rhagovelia hirsuta sp. n. — 84-89 male, 90-93 female. 84, 90, dorsal habitus apterous; 85, front leg; 86, 91, hind leg; 87, genital capsule; 88, mid-femora; 89, 92, antennae; 93, external genitalia. Figs. 84-86, 88-93 scale line 1 mm, fig. 87 scale line 0.5 mm. long, maximum width 1.4-1.48 mm, female (fig. 90) 4-4.4 mm long, maximum width 1.52-1.64 mm. Coloration. — Male (fig. 84): Dorsum generally black. Tergites and most of connexivum black, upper margins narrowly shining dark brown. Seventh ter- gite with an elongate triangular shining area. Anterior margin of pronotum orange brown. Lateral margins 40 of pronotum, prosternum, mesosternum and ster- nites greyish blue tomentose. Proepisternum pale red- dish brown, meso and metasternum dark grey mar- gined reddish brown. Trochanters yellowish brown. Basal 0.2 of front femur pale yellow, remainder of front leg, middle and hind femora dark iridescent green, remainder of legs black. Antennae black, basal Table 15. Proportions of antennal segments of Rhagovelia hirsuta sp. n. I Il Ul IV Apterous male 48 27 32 29 Apterous female 49 25 34 28.5 Table 16. Proportions of leg segments of Rhagovelia hirsuta sp. n. Femur Tibia Tarsus I Tarsus II Tarsus III d frontleg 56 62 5 16 = 9 frontleg 57 62 5 17 - 3 middle leg 95 73 6 32 40 ? middle leg 92 72 5 32 41 d hindleg 85 84 5 8 19 9 hindleg 75 88 4 7 19 0.2-0.3 of Ist pale yellow. Structure. — Male: Maximum eye length clearly longer than pronotum (18: 16). Pronotal width 3.4- 3.5x median length. Mesonotum 2.3x pronotal length, metanotum 0.3x length of pronotum. Tergites variable length, 1-8 are 12 : 12.5: 10.5: 10: 10 : 11.5 : 22 : 21. Eighth tergite densely pilose. Pronotum laterally, meso and metaepimeron, lateral margins of metanotum, tergites and lower margins of connexivum with very long prominent semi-erect hairs. Proepisternum, prosternum and lateral margins of pronotum with fields of black spicules. Jugum al- most touching the antenniferous tubercles. Second sternite bluntly keeled, 7th slightly flattened, covered with adpressed golden hairs. Eighth segment ventral- ly concave and distally reflexed downwards. Legs, all coxae with clusters of long golden hairs, hairs and spines of all legs self coloured. Front leg (fig. 85). Mid femora (fig. 88) slightly flattened and with fine hairs. Hind leg (fig. 86) trochanter unarmed. Femus mod- erately incrassate, antero-ventrad with a row of even- sized setae, medianly and posteriorly with larger irreg- ular sized spines. Tibia straight armed with small setae along inner margin. Antennae (fig. 89) Ist seg- ment not very spinose. Genital capsule (fig. 87) dis- tally bluntly acuminate. Parameres large almost pad- dle-like. Coloration. — Female (fig. 90): Vertex, posterior margin of pronotum, mesonotum, metanotum and most of Ist tergite black. Anterior margin of pro- notum pale orange band reaching inner margin of eyes. Lateral margins of pronotum, visible tergites and lateral margins of visible sternites evanescent bluish grey. Connexivum apically narrowly dark brown margined narrowly pale yellow. Prosternum pale brown. Remainder of coloration similar to male. LANSBURY: Papuan Rhago velia Structure. — Female: Eye length subequal to me- dian pronotal length (17:18). Pronotal width 3.22x median length. Mesonotum 2x median pronotal length, metanotum about 0.5x pronotal length. Nereitesyl-8rare 12.5: 12291071 3:17:19, 222014. Eighth tergite and proctiger strongly reflexed down- wards (fig. 93). Pronotum, lateral margins of meso and metanotum and posterior margin of Ist tergite, pro and mesosternum with scattered prominent black hairs. Posterior margin of pronotum, all of meso- metanotum and Ist tergite with adpressed short gold- en hairs. Tergites 3-6 bluntly raised medianly with discrete clusters of golden hairs increasing in length and density on 6th tergite. Connexivum anteriorly re- flexed outwards, from 3rd tergite converging and meeting across 7th. Connexival margins of Gth with a cluster of long hairs. Lateral margins of 8th segment with two (1+1) clusters of curled hairs. Sternites cov- ered with fine adpressed golden hairs, very dense on 7th sternite. Proctiger and ancilliary structures pilose. Hind femur with a single prominent spine ventrad, tibia straight, not armed along inner margin (fig. 91). Antennae (fig. 92). Etymology. — hirsuta refers to the hairy appearance of this species. Remarks. — This species tends to resemble R. aure- ospicata sp. n. The female hirsuta differs in having the connexivum converging posteriorly. The eye/prono- tal ratio is subequal whereas the pronotum of aureos- picata sp n. is longer than eye and the male middle fe- mur has several stout setae proximo-ventrally whereas male middle femur of hirsuta sp. n. is fringed with fine hairs ventrally. Rhagovelia aureospicata sp. n. (figs. 94-104, tabs. 17-18) Type material. — Holotype d : Papua New Guinea, Oro Province, Kokoda, Mambare river, 24.ix.1989, R. N. B. Prior (OXUM). — Paratypes, same data, 49 (OXUM). Description Adult apterous. — Male (fig. 94) 4.24 mm long, maximum width 1.56 mm, female (fig. 95) 4.40-4.44 mm long, maximum width 1.60-1.72 mm. Coloration. — Male (fig. 94): Dorsal ground colour black. Anterior pronotal margin pale yellow. Connexivum dark brown with a pale brownish yellow band near upper margin. Prosternum pale yellow. Mesosternum upper margins evanescent blue remain- der pale yellowish brown. Metasternum evanescent blue graduating into pale yellowish brown. Coxae and trochanters yellowish brown. Inner proximal half of front femur pale yellow, outer margin narrowly pale yellow. Front tibia and middle legs iridescent 4] TIDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 greenish brown. Hind femur mostly pale brown, dis- tal and dorsal surfaces and hind tibia and tarsi dark brown. Sternites 1-3 black, remainder yellowish brown tending to merge into black along connexival line. Antennae black, proximal third of Ist segment pale yellow graduating into black. Structure. — Male: Pronotum longer than eye (25:18) pronotal width 2.44x median length. Mesonotum 1.2x length of pronotum and 4.2x length of metanotum. Lateral margins of prothorax fringed with long hairs. Tergites 1-6 subequal, 1-8 are PAPAS 2 12-3). 28): 50. Black dorsal ar- eas covered with fine golden spicules other than 7th tergite which is slightly flattened and dull shining, posterior margin fringed with long hairs. Distal con- nexival segment with very long black hairs which are carried round posterior margin of 7th sternite. Proepisternum without black spicules, pro-meso and metasternum with scattered black semi-erect hairs. Jugum broad almost touching antenniferous tuber- cles. Seventh sternite shallowly depressed, covered with fine brown hairs, ventrally segment depressed adjacent to 7th sternite, segment sharply reflexed downwards. Front leg (fig. 99) tibia slightly explanate distally. Middle femur (fig. 100) proximo-ventrad with four spines. Hind leg (fig. 98) femur moderately incrassate, proximo-ventrad with a row of seven even sized setae, medially with four large projections and a variable number of smaller iregular setae. Tibia sinu- ate, inner margin armed with stout projection distal- ly and a row of small setae. Antennae (fig. 101) Ist and 2nd segments moderately hairy. Genital capsule (fig. 102), distal end of capsule curved upwards. Parameres elongate, distally rounded. Coloration. — Female (fig. 95): Head and thorax similar to male. Tergites 5-7 with variable shining ar- eas. Pale yellowish brown connexival band tending to be slightly wider than of male. Metasternum black, upper lateral margins of sternites black decreasing in area to 7th sternite. Basal half of front femur pale yel- low. Ventrally front femur, middle and hind femura pale brown, dorsal surface of middle and hind femo- ra iridescent green. Structure. — Female: Pronotum longer than eye (25:18) pronotal width 2.6x median length. Mesonotum 1.2x length of pronotum and 6x length of metanotum. Tergites variable in length, 1-8 are 13 : SE1527145171725%: W522 2 WOsStiucturally the thorax is same as male, the fringe of long hairs not present. The density of gold spicules on pronotum is not as dense as on mesonotum which has the spicules arranged in a semi circular pattern. The spicules on LANSBURY: Papuan Rhagovelia Tabel 17. Proportions of antennal segments of Rhagovelia aureospicata sp. n. I Il III IV Apterous male 50 34 42 34 Apterous female 50 32 38 3] Tabel 18. Proportions of leg segments of Rhagovelia aureos- picata sp. n. Femur Tibia TarsusI Tarsus II Tarsus III d front leg 63 64 5 119% = 9 front leg 60 Gl 5 17 - 3 middle leg 108 84 5 37 46 9 middle leg 93 15 5 35 42 d hindleg 105 055 10 DD 9 hind leg 79 95 4 9 20 the metanotum and tergites 1-2 are denser than on male. Seventh tergite flattened and shining. Connexivum anteriorly reflexed outwards becoming more erect posteriorly. Proctiger and gonocoxal plates strongly reflexed downwards (fig. 97). Hind leg (fig. 103) femur with a prominent spine distad ventrally, tibia straight, inner margin unarmed. Antennae (fig. 104). Etymology. — Aureospicata alludes to the golden spicules on the dorsum. Remark. — This species belongs to the group which have the pronotum longer than the eye length, the former being shorter than the mesonotum. The pro- thorax is densely clothed with golden spicules and the male mid-femur with proximo-ventrad group of setae suggests a close relationship with herzogensis sp. n. Rhagovelia herzogensis sp. n. (figs. 105-116, tabs. 19-20) Type material. — Holotype d apterous: New Guinea, Morobe District, Herzog Mountains, Vagau c. 4000 feet, 4- 17.1.1965, M. E. Bacchus (BMNH). — Paratypes, same data, 36 39 apterous, 29 macropterous (BMNH, OXUM). Description Adult apterous. — Male (fig. 106) 4.08-4.2 mm long, maximum width 1.4-1.48 mm, female (fig. 107) 4 mm long, maximum width 1.56 mm. Adult macropterous. Female (fig. 105) 4.8 mm long, maxi- mum width 1.72 mm. Coloration. — Male (fig. 106): Head, thorax and abdomen dorsally black. Pronotum anteriorly broad- Figs. 94-104. Rhagovelia aureospicata sp. n. — 94, dorsal habitus apterous male; 95, ibid female; 96, external genitalia male; 97, ibid female; 98-102 male, 103-104 female. 98, 103, hind leg; 99, front leg; 100, mid-femora; 101, 104, antennae; 102, genital capsule. Figs. 94-101, 103-104 scale line 1 mm, fig. 102 scale line 0.5 mm. 43 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 ly yellow. Inner lateral margins of connexivum with yellow stripe. Seventh tergite slightly shining. Anterior margin of mesonotum variably dark reddish brown. Pro and mesopleura pale yellow, upper lateral margins of metapleura black, remainder yellow. Jugum and prosternum pale yellow, meso and meta- sternum becoming progressively dark brown. Sternites 2-4 dark reddish brown, 5-8 pale yellowish brown. Lateral margins of sternites black, tapering distally to 7. Coxae and trochanters pale yellow. Front femur proximally yellowish brown, distally fe- mur, tibia, middle and hind legs dorsally greenish black shining. Underside of front femur and hind fe- mur yellowish brown, remainder of legs dark brown on ventral surfaces. Structure. — Male: Median pronotal length longer than eye length (20 : 18). Pronotal width 3x median length. Mesonotum 1.6x median pronotal length and 6.4x length of metanotum. Tergites, 1-8 are 13 : 10: Ile Wil 2 ils Ws D7 3 By, Whoyoer merrein Or 7dn were gite slightly shining, rugose, 8th very pilose. Lateral margins of pronotum, thorax, upper margins of con- nexivum and tergites with scattered long black hairs. Posterior margin of pronotum, meso and metanotum and tergites 1-6 covered with silvery short hairs. Connexivum reflexed slightly outwards with sparse silvery pubescense. Distal connexival segment with cluster of long curved hairs. Proepisternum without black spicules, pro-meso and metasternum with scat- tered brownish-black semi-erect spines. Jugum al- most touching antenniferous tubercles. Second ster- nite obtusely carinate, 3rd less so fringed with two (1+1) rounded pilose elongated ridges. Eighth ster- nite sharply depressed adjacent to posterior margin of 7th sternite. Front leg (fig. 109). Middle femur (fig. 110) proximo-ventrad with several stout setae, tiba flattened. Hind leg (fig. 108) femur moderately in- crassate, ventrally with two prominent projections and several smaller setae, tibia slightly sinuate, inner margin armed with setae. Antennae (fig. 111) seg- ments 3-4 missing. Genital capsule (fig. 116) capsule distally acuminate, parameres elongate. External gen- italia and abdomen (fig. 112). Coloration. — Female (fig. 107): Similar in general appearance to male. Anterior yellow band on prono- tum broader and tending to cover lateral margins pos- teriorly. Reddish brown area of mesonotum more prominent. Tergites 5-7 with shining areas. Front fe- mur basal third pale yellow dorsally, ventrally pale colour extending along femur. Middle and hind legs dull greenish black ventrally pale yellowish brown. Abdominal coloration similar to male. Table 19. Proportions of antennal segment of Rhagovelia herzogensis sp. n. I II III IV Apterous male 45 28 = = Apterous female 45 30 39 31 Macropterous female 45 31 39 32.5 Table 20. Proportions of leg segments of Rhagovelia herzo- gensis sp. n. Femur Tibia TarsusI Tarsus II Tarsus III Apterous d front leg 61 67 4 16 = 9 front leg 60 60 5 16 > 3 middle leg 100 78 6 33 41 9 middle leg 94 68 5 32 42 d hind leg SOS 4.5 9 19.5 9 hind leg 74 88 3 7 20 Macropterous 9 front leg 55 60 4 15 = 9 middle leg 91 69 5 34 43 © hind leg 17 92 3.5 8 20 Structure. — Female: Median pronotal length long- er than eye length (23:19.5). Pronotal width almost 3x median length. Mesonotum 1.17x median prono- tal length and 6.75x length of metanotum. Tergites Nes} eves W725) 2 WAS) MS ss 19224105, Tei 1-2, sometimes 3 with a slight median ridge, distal shining tergites slightly flattened. Connexivum erect anteriorly, converging posteriorly, either reflexed in- wards slightly or curved over and almost touching posteriorly obscuring shining area of 7th tergite. Posterior half of pronotum with scattered silvery hairs, mesonotum more densely pilose with hairs ar- ranged in semi-circular pattern either side of midline. Lateral margins of metanotum and tergites 1-4 with decreasing areas of silvery hairs. Sternites medianly clothed with fine semi-erect golden hairs. Hind leg (fig. 113) femur slightly incrassate with a prominent projection distally, tibia straight, inner margin un- armed. Proctiger small and sharply reflexed down- wards (fig. 114). Macropterous female (fig. 105). — Similar to apter- ous form, except pronotum greatly lengthened cau- dad, anterior margin narrowly pale yellow, posterior margin reddish brown. Wings with four closed cells, wings brown, venation black. Wings extending well beyond end of abdomen. Figs. 105-115. Rhagovelia herzogensis sp. n. — 105, dorsal habitus female macropterous; 106, dorsal habitus male apterous; 107, ibid female; 108-112 male, 113-115 female. 108, 113, hind leg; 109, front leg; 110, mid-femora; 111, 115, antennae; 112, 114, external genitalia. Scale line 1 mm for figs 105-107, and 108-115 respectively. 44 LANSBURY: Papuan Rhagovelia 45 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 116. Rhagovelia herzo- gensis sp. n. — Male genital capsule. Scale line 0.5 mm. Ss SS 4, S NN N N N 116 Etymology. — Specific name alludes to the type lo- cality Herzog Mountains. Remarks. — This species is closely allied to aureospic- ata sp. n., male mid femura of both species bearing a short row of small spines proximo-ventrally. Male gen- ital capsules similar. Lateral pilosity is much denser of aureospicata (fig. 94) compared with herzogensis (fig. 106). Female herzogensis has the connexivum converg- ing posteriorly with clusters of long hairs (fig. 107), connexivum of female aureospicata not convergent dis- tally and lacking the clusters of long hairs (fig. 95). Rhagovelia crinita sp. n. (figs. 117-125, tabs. 21-22) Type material. — Holotype d apterous: New Guinea, Madang District, Finisterre Mts., c. 5500 feet, 30.x.- 15.xi.1964, M. E. Bacchus (BMNH). — Paratype: same data, 54 119 (BMNH, OXUM). Additional material. — New Guinea: NE. Wau, Morobe District, Mt. Missim, 1200 m. 19.iv.1966, G. Lippert, 16 apterous (BPBM). Description Adult apterous. — Male (fig. 117) 3.92-4.32 mm long, maximum width 1.42-1.62 mm, female (fig. 118) 3.92-4.32 mm long, maximum width 1.42-1.66 mm. Coloration. — Male (fig. 117): Head velvety black. Pronotum anteriorly bright yellowish orange, pro- pleura, proepisternum and jugum pale yellow. Posterior half or more of pronotum, meso-metano- tum, tergites 1-6 and inner lower margin of connexi- vum greyish black. Tergites anteriorly narrowly black. Dorsum covered with adpressed golden spicules which tend to be denser on median areas of tergites. Seventh tergite shining, 8th dark brown. Slightly ten- 46 NS I eral specimens tend to have anterior half of mesono- tum reddish brown. Connexivum broadly yellowish orange, apically narrowly black. Meso and metaepim- eron dark reddish brown. Sternites 2-5 reddish brown, black laterally, 6-8 tending to be pale yellow. Proximal antennal segment narrowly pale yellow, re- mainder of segments black. Coxae pale yellow. Inner margin front femur and tibia pale yellow, distal outer margins dark brown. Ventral margins of middle legs pale brown, dorsally brown-black. Hind femur pale yellow, dorsally dark brown, tibia and tarsi reddish brown. Structure. — Male: Median pronotal length sub- equal to eye length (20:19). Pronotal width 3x me- dian length. Mesonotum 1.6x pronotal length and 6.4x metanotum at midline. Tergites 1-8 are 13: 13: 13 : 12.5 : 14 : 16 : 25.5 : 20. Seventh tergite broad- ly faintly shining appearing slightly transversely ras- trate, 8th tergite densely pilose. Connexivum almost erect, inner margins fringed with fine brown hairs, distal lateral margins fringed with very long hairs. Pronotum and propleura with scattered black hairs, posterior margin of propleura with a few dark brown pits. Jugum broad almost reaching antenniferous tu- bercles. Thorax ventrally without fields of black spic- ules. Meso and metapleura with scattered black hairs. Second sternite and part of 3rd moderately carinate, 2-5 with scattered adpressed yellow hairs, 6th densely hairy, 7th slightly flattened, laterally with two (1+1) rows of long yellow hairs, 8th proximally acutely de- pressed. Front leg (fig. 119) femur fringed with black hairs, tibia proximally narrow, medially explanate with a prominent fringe of stout setae/hairs decreas- ing in length distally, inner margin distally acumi- nate. Middle femur slightly flattened. Hind leg (fig. 120) femur moderately incrassate, ventral margin LANSBURY: Papuan Rhagovelia Figs. 117-125. Rhagovelia crinita sp. n. — 117, 119-122 male, 118, 123-125 female. 117-118, dorsal habitus apterous; 119, front leg; 120, 123, hind leg; 121, 124, antennae; 122, genital capsule; 125, external genitalia. Figs. 117-121, 123-125 scale line 1 mm, fig. 122 scale line 0.5 mm. 47 “TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 proximally with a row of regular sized setae, distally with two prominent projections and several larger se- tae. Tibia almost straight, inner margin armed with a row of irregularly spaced setae. Antennae (fig. 121). Genital capsule (fig. 122) capsule distally truncate, parameres elongate, finely pilose. Coloration. — Female (fig. 118): Broadly resembling male. Mesonotum tending to be more widely anterior- ly dark reddish brown merging to black laterally and posteriorly. Tergites 3-8 becoming more yellowish brown medianly, 5-7 medianly pale brown and shining areas increasing in size distally. Some females have 3-8 becoming progressively more yellowish brown distally, 8 entirely yellowish brown. Connexivum and under- side of thorax usually similar to male, metasternum sometimes very dark reddish brown. Sternite 2-7 red- dish brown, distally tending to be pale brown. Black areas viewed from different angles appearing to bluish- black. Antennae and legs similar to male. Structure. — Female: Pronotum slightly longer than eye (20:18). Pronotal width 3.7x median length. Mesonotum 1.57x median pronotal length and 5.7x metanotum at midline. Dark areas of pro-meso and metanotum with scattered long black hairs. Tergites 128 are ASIN 2 1620102029 15. Terres 1-2 raised above following tergites. Connexivum an- teriorly erect, reflexed over lateral margins of 3-7, converging slightly posteriorly. Distally connexivum appearing acuminate with a tuft of black hairs. Sternites 1-6 evenly rounded. Distal margin of 7th transversely depressed (fig. 125). Eighth segment re- flexed downwards, proctiger small. Antennae (fig. 124). Front femur densely pilose, tibia laterally with long hairs subequal to tibial width, distally with lon- gitudinal depression. Middle femur slightly flattened. Hind leg (fig. 123) femur not incrassate, distally with a prominent projection, tibia straight, unarmed along inner margin. Etymology. — crinita alludes to the fringe of stout hairs and spines along inner margin of male front tibia. Remarks. — Rhagovelia crinita sp. n. and thysanotos sp. n. are superficially rather similar. The male front tibiae of both have a prominent fringe of stout hairs/- spines along inner margin (figs. 119, 129). Distal connexival segments of female thysanotos densely fringed with long hairs, crinita females have smaller tufts of hairs. The male genitalia of crinita differ from thysanotos being smaller, parameres of crinita distally bluntly acuminate (fig. 122) those of thysanotos are bent over apically. Rhagovelia thysanotos sp n. (figs. 126-135, tabs. 23-24) Type material. — Holotype d apterous: New Guinea, 48 Table 21. Proportions of antennal segments of Rhagovelia crinita sp. n. I II III IV Apterous male 50 33 48 38 Apterous female 50 31 48 38 Table 22. Proportions of leg segments of Rhagovelia crinita sp. n. Femur Tibia Tarsus I Tarsus II Tarsus III d front leg 70 13 17 = 9 front leg 67 68 6 17 = d middle leg 110 83 5 41 43 9 middle leg 105 2 3 40 41 3 hind leg 100 UO 5 14 26 ® hind leg 90 MOSS 13 24 Torricelli Mts., 1700 feet, 11.11.1939, G. P. Moore (BMNH). — Paratypes: same data, 28 19 apterous, 1d macropterous (BMNH, OXUM). Description Adult apterous. — Male (fig. 126) 3.84-3.92 mm long, maximum width 1.4 mm. Female (fig. 132) 3.96 mm long, maximum width 1.52 mm. Adult macropterous. Male 4.08 mm long, maximum width 1.48 mm. Coloration. — Male (fig. 126): Head velvety black. Pronotum anteriorly pale yellow reaching just beyond inner margin of eyes. Lateral margins, propleura and proepisternum bluish-black evanescent. Pronotum posteriorly, meso, metanotum, tergites and most of connexivum black. Upper margins of connexivum narrowly dark brown, apex narrowly blackish-brown. Seventh tergite faintly shining, 8th shining iridescent green. Jugum and coxae pale yellow. Underside of thorax and sternites greyish black, 6th sternite with lateral margins shining brown. Antennae black, prox- imal 0.2 of first segment pale yellow. Front leg, femo- ra proximally pale yellow, remainder with tibia dark iridescent green. Middle femur iridescent green, tibia and tarsi rather darker. Hind femora dark brown with a faint green iridescent patch on outer margin, tibia and tarsi dark brown-black. Structure. — Male: Eye length greater than median pronotal length (18:16). Pronotal width 3.43x me- dian length. Mesonotum 1.9x pronotal length and 4.3x length of metanotum at midline. Lateral bluish margins of pronotum with scattered fields of minute black pits. Posterior margin of propleura with an al- most continuous row of similar pits. Tergites 1-8 are MES 2 13:3 1.529) 75 OR 1322520) hornax and tergites covered with short fine pale brown hairs. Overlying these long black hairs which are particular- ly dense along posterior margin of 7th tergite and LANSBURY: Papuan Rhagovelia i \ N LA —= Figs. 126-135. Rhagovelia thysanotos sp. n. — 126-131 male, 132-134 female. 126, 132, dorsal habitus apterous; 127, 134, hind leg; 128, mid-femora; 129, front leg; 130, antenna; 131, 133, external genitalia; 135 male genital capsule. Figs. 126-134 scale line 1 mm, fig. 135 scale line 0.5 mm. 49 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 whole of 8th. Connexivum anteriorly slightly reflexed outwards, posteriorly erect, inner and outer margins pilose. Distal connexival segment fringed with very long hairs. Eighth tergite distally with two (1+1) clus- ters of shorter hairs. Pro and mesosternum with long black hairs. Jugum broad almost touching antennifer- ous tubercles. Second, 3rd and proximal part of 4th sternites bluntly carinate and with a field of short fine hairs. Fifth and 6th sternites laterally pilose, slightly flattened, 7th more pilose, transversely depressed. Front leg (fig. 129) femur moderately pilose. Tibia proximally narrow, medianly explanate with promi- nent fring of stout hairs decreasing in length distally, inner margin distally acuminate. Middle femur (fig. 128) very slightly flattened. Hind leg (fig. 127) femo- ra moderately incrassate, medianly almost parallel sid- ed. Ventral margin anteriorly and posteriorly with al- most regular rows of triangular setae. Anteriorly with prominent projection, medianly with several smaller projections. Hind tiba slightly sinuate, inner margin armed with setae varying in size and density. Antennae (fig. 130). Genital capsule ventrally pilose, parameres elongate, distally slightly hooked (fig. 135). Entire genital segment very pilose (fig. 131). Coloration. — Female (fig. 132): Broadly resembles male, anterior margin of pronotum more brightly coloured. Tergites appearing dull black with a faint bluish evanescence, 7-8 with small shining bluish- black discs, that of 8 covering most of segment. Connexivum apically broadly dark brown. Thorax ventrally and sternites bluish black. Legs and anten- nae similar to male. Structure. — Female: Eye length less than median pronotal length (16:18). Median pronotal width 3.1x median length. Mesonotum 1.3x median pronotal length, posterior margin deeply emarginate, with a row of black pits along margin. Metanotum 0.24x length of mesonotum. Lateral margins of pronotum with scattered long black hairs. Tergites 1-8 are 13 : SENDER : 17.5 2 22,5% 15: Connie slightly reflexed outwards anteriorly, converging at 4th tergite erect, posteriorly slightly sinuate converg- ing and erect distally, with prominent clusters of long dark brown-black hairs. External genitalia and procti- ger sharply reflexed downwards, proctiger with lateral clusters of hairs (fig. 133). Hind leg (fig. 134) femur slightly incrassate, ventrally with two prominent pro- jections and several smaller setae distally. Hind tibia straight, unarmed along inner margin. Macropterous male. — Elytra damaged one missing, specimen in poor condition. Similar in coloration to apterous male. Pronotum lengthened caudad, anteri- or margin with transverse pale yellow band just reach- ing inner margin of eyes. Wing reaching beyond end of abdomen with four closed cells, basal two small, extending into distal part of wing. Elytra yellowish 50 Table 23. Proportions of antennal segments of Rhagovelia thysanotos sp. n. * = missing. I II III IV Apterous male 49 27 32 30 Apterous female 47 DD + = Macropterous male 4] 21 si % Table 24. Proportions of leg segments of Rhagovelia thysano- tos sp. n. Femur Tibia Tarsus I Tarsus II Tarsus III Apterous 3 front leg 59 60 4 9 front leg 55 55 51 5 - d middle leg 95 70 5 31 4] 9 middle leg 82 61 5) 29 40 3 hind leg 80 80 5 14 18 4 16.5 = © hind leg 73 75 7 17 Macropterous d front leg 52 52 4 13.5 = 3 middle leg 83 53 3 26.5 40 3 hind leg 75 65 4 7 17 brown, veins black, R and Sc with many dark brown hairs. Hind leg similar to apterous male. Etymology. — Thysanotos alludes to the fring of stout hairs along the inner median margin of the male front tibia. Remarks. — Rhagovelia thysanotos sp. n. has features in common with crinita sp. n. Males of both species having a prominent fringe along inner margin of front tibia. The anterior pale band of thysanotos does not reach the lateral margins of pronotum and the shining areas of the distal tergites are much reduced compared with crinita. The male genital capsule of thysanotos is relatively much larger than of crinita and the parameres of the former are somewhat bent over distally, almost spatulate. The condition of the type series of thysanotos is very poor, only one male having complete antennae. All specimens covered with detri- tus/dust with signs of mite damage at some stage. Rhagovelia caesius sp. n. (figs. 136-150, tabs. 25-26) Type material. — Holotype & apterous: Papua New Guinea, Oro Province, Popondetta, Sambogo River, close to stones, 30.iv.1989, R. N. B. Prior (OXUM). — Paratypes: Same data, 648 289 apterous, 1d 19 macropterous (OX- UM); Papua New Guinea, Milne Bay Province, streams near Alotau, 27.iv.1989, Prior, 78 69 apterous (OXUM). Additional material. — Papua: Kokoda, 1200 feet, ix.1933, L. E. Cheesman, 7d 79 apterous; New Guinea, LANSBURY: Papuan Rhagovelia Figs. 136-150. Rhagovelia caesius sp. n. — 136, dorsal habitus macropterous female; 137, dorsal habitus apterous female; 138, ibid male; 139-145 male, 146-150 female. 139, 146, front leg; 140, 147, distal end of tibia and tarsi of front leg; 141, 148, hind leg; 142, 149, distal end of tibia and tarsi of hind leg; 145, 150, antennae. Figs. 136-138 scale line 1 mm, figs. 139, 141, 144-146, 148, 150 scale line 1 mm, figs. 140, 142-143, 147, 149 scale line 0.5 mm. DI TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Table 25. Proportions of antennal segments of Rhagovelia caesius sp. N. I II III IV Apterous male 38 19.5 17 19 Apterous female 48.5 30 23 23 Macropterous male 36 20 18 20 Table 26. Proportions of leg segments of Rhagovelia caesius sp. n. Femur Tibia Tarsus I Tarsus II Tarsus III Apterous 3 frontleg 45 46 3 ? front leg DE) 60 4. 3 middle leg 67 54 À 29 31 9 middle leg 85 67 5 3 hind leg 48 71 2 3 9 hind leg 60 88 9 19 Macropterous d front leg 46 46 3 9 - 3 middle leg 68 23 28 34 3 hind leg 48 70 3 3 11 Morobe District, Lae-Bululo Road, 28.xii.1964, M.E. Bacchus, 1d 29 apterous; Papua New Guinea, Morobe Province, Buso, ix-xi.1979, J. Martin, 3d 32 apterous, 1d 12 macropterous (BMNH). Description Adult apterous. — Male (fig. 138) 2.32-2.52 mm long, maximum width 1-1.08 mm. Female (fig. 137) 3.28-3.36 mm long, maximum width 1.56-1.64 mm. Adult macropterous. Male 2.84-2.92 mm long, maxi- mum width 1.24-1.28 mm. Female (fig. 136) 3.4 mm long, maximum width 1.64 mm. Coloration. — Male (fig. 138): Head velvety black. General dorsal coloration appearing greyish black. Pronotum anteriorly with a rather indistinct orange brown transverse band not reaching the inner margin of eyes. Lateral margins of mesonotum sometimes faintly orange brown. Pro, meso and meta pleura tu- mescent black, viewed from various angles appearing dark evanescent blue. Antenniferous tubercles shin- ing black. Jugum and rostrum shining iridescent greenish black. Median area of 7th sternite shining pale brown. Coxae and trochanters shining yellowish brown. Legs iridescent greenish black. Antennae black, basal third of proximal segment pale yellow. Structure. — Male: Eye length greater than median pronotal length (13:9). Pronotal width 4.4x median length. Mesonotum 3.6x median pronotal length, lat- eral posterior margins emarginate. Metanotum about 0.12x length of mesonotum. Lateral margins of pro- notum, all of mesonotum, metanotum, tergites and connexivum with long semi-erect black hairs overly- 52. ing short greyish hairs. Tergites 1-8 are 5.5 : 5.75 : 5.9:6:6:6:12.5:9. Connexivum usually flattened, from side view tergites visible. Jugum clearly distinct from antenniferous tubercles, proepisternum lacking black spicules. Front coxae with clusters of short se- tae. Front leg (figs. 139-140) femur moderately fine- ly spinose, tibia curved explanate distally with a prominent ridge of fine setae along inner distal mar- gin (fig. 140). Middle and hind trochanters and cox- ae with stout short yellowish hairs. Hind leg similar to middle leg, not incrassate and spinose (figs. 141-142), middle and hind femora slightly flattened, hind tarsi pilose (fig. 142). Antennae (fig. 145) proximal seg- ment with many long fine spines. Seventh abdominal segment rounded, genital segment cylindrical and pi- lose. Genital capsule (fig. 143) distally truncate, para- meres elongate. Coloration. — Female (fig. 137): Ground colour appearing blackish suffused with faint greenish black tumescence, pro-meso and metanotum sometimes suffused pale yellowish brown. Pronotum anteriorly with a faint transverse orange stripe merging into greenish tumescence almost reaching inner margin of eyes. Mesonotum and metanotum laterally pale greenish black, mesonotum and metanotum with field of minute black spicules. Connexivum and ter- gites uniform greenish black. Antenniferous tuber- cles, jugum and rostrum shining dark brown-black. Pro-meso and metaepisternum silvery grey tumes- cent. Venter greyish tumescent, 7th sternite shining dull black. Coxae and trochanters variably pale-dark yellow. Proximal margin of front femur narrowly pale yellow, remainder of all legs greenish black iridescent. Antennae black, proximal third of first segment pale yellow. Structure. — Female: Eye length greater than me- dian pronotal length (17:13). Pronotal width 4.1x median length. Mesonotum 3.3x pronotal length and 7.1x midline of metanotum. Posterior lateral margins of mesonotum slightly emarginate, posterior margin deeply emarginate. Tergites 1-8 are 6.5: 11 : 10 : 8.5: 9:10:11: 10. Connexivum usually flattened, some- times elevated reflexed outwards. Dorsally thorax, ab- domen and connexivum not pilose as in males. Proepisternum without black spicules. Lateral mar- gins of prothorax and abdomen with scattered erect black hairs. Lateral margins of sternites 2-6 with sparse golden short hairs. Seventh sternite black shin- ing, laterally with fields of longer golden hairs increas- ing in density along distal margin. Jugum distinct from antenniferous tubercles. Front coxae and tro- chanters with pale yellow spine-like hairs. Front leg (fig. 146) femur and tibia curved with inner distal margin of tibia slightly expanded (fig. 147). Middle and hind legs similar, hind leg (fig. 148) tarsi elongate and pilose (fig. 149). Antennae (fig. 15) proximal seg- ment with many long fine hairs, 2nd segment sparse- ly pilose. Macropterous male. — Coloration not differing greatly from apterous form. Pronotum long (1.08 mm). Pronotum at its widest raised very prominent- ly. Forewings more or less uniformly brown extend- ing well beyond the end of abdomen. Venation with three closed cells proximally. Macropterous female (fig. 136). — Coloration, re- sembles apterous form. Pronotum about 1.52 mm long, transverse orange band slightly more obvious than in apterous form, posterior lobe transversely stri- ated and finely pitted. Forewings basally greenish grey, distally orange brown reaching just beyond end of abdomen. Three closed cells in basal dark area of wing. Etymology. — Caesius alludes to the bluish grey ap- pearance of the adult especially the female. Remarks. — This species differs from others so far known from New Guinea and Solomon Islands by the absence of prominent spines/projections on the hind femora of either sex. The sexual dimorphic body size indicates phoretic behaviour. A pair from the Sambogo River were taken ‘in cop’, the curved male front femora fit in the lateral divergent margins of the female mesonotum and the curved male tibia encircle the pleural margins. The males also differ from the fe- males in having the dorsum covered with long hairs, whereas the females lack the long hairs dorsally and their coloration is tomentose greyish blue. The fe- males appear to be abnormally flattened compared to the males, this aspect is exagerated by the flat connex- ivum and broad prothorax in the majoriy of speci- mens. DISCUSSION The Rhagovelia of the northern provinces of Papua New Guinea have a distinctive endemic fauna. The species so far recorded from the Oro Province (Popondetta region - Kokoda) see map in Cheesman (1943) includes priori sp. n.; hirsuta sp. n.; aureospi- cata sp. n. and caesius sp. n. representing three species groups, and does suggest that extensive sampling of rain forest creeks and rivers in ‘limited areas’ will gre- atly increase the Rhagovelia species known from Papua New Guinea. Rhagovelia caesius sp. n. is the most aberrant species so far recorded from Papua New Guinea. The materi- al included under this name has a distribution ex- tending from Oro, Kokoda, 1200 feet to sealevel at Alotau, Milne Bay Province. Further studies may show that ‘caesius is a group of closely allied species. Polhemus & Polhemus (1988) discuss at length the zoogeography of Rhagovelia, commenting on the LANSBURY: Papuan Rhagovelia dearth of described species from the Philippines, New Guinea, Solomons and other zoogeographically im- portant areas. The Rhagovelia of New Guinea are thought to have radiated extensively during the late Tertiary and the existing Rhagovelia fauna is stated to be endemic to New Guinea and to have originated primarily since the Oligocene. Sources of New Guinea Rhagovelia are thought to be New Caledonia and relict forms from northern australian refugia. Kikkawa et al. (1981) postulate that the continen- tal margin of Australia extended to southern New Guinea since the Lower Jurassic. Much of northern Australia and New Guinea were subsea suring the ear- ly Cretaceous. The northward movement of Australia continued and the collision of crustal plates in the mid Miocene created the present land mass of New Guinea. By the Pliocene, most of New Guinea had emerged from the sea. The major uplift of the present mountain ranges with major faulting accompanied by widespread volcanic activity in the late Miocene - ear- ly Pliocene (7 Ma) and continues today. Recent stud- ies of the north coast of New Guinea show 32 ter- ranes (micro-tectonic plates), each of which has ‘docked’ to the north coast over the past 20 million years, as the island was pushed northwards through a complex subduction zone. An earthquake on the 15th May, 1992 caused a 10 cm uplift near Sialum (Morobe Province), which was felt very strongly along the Madang coastline. The total uplift may exceed 3000-4000 m. Smith & Briden (1977) commenting on the tectonic folding and uplift in central New Guinea and ‘fragmented plates’ suggest the fragmented plates created the ‘Indonesian islands’ and may have joined the conti- nents of Asia and Australia allowing the first wave of Asian migrants to enter the Australian region. The evidence of palaeogeography, climate and geology present a very confusing set of scenarios to ac- count for the diversity of some groups of water-bugs in New Guinea. During the Pleistocene sea-level changes, New Guinea was linked to Australia until between 6500-8000 B. P. The aridity caused by the lowered sea-levels may have inhibited the movement of Rhagovelia between Australia and New Guinea and species on either land mass may have survived in re- fugic mesic habitats. The record of R. australica Kirkaldy from Mt. Lewis is of more than passing interest, as a number of relict forms have been record- ed from Mt. Lewis. Other examples include the leaf- hopper Myerslopia taylori Evans, a primitive tribe hitherto known from Madagascar, New Zealand, Chile and Juan Fernandez, further /gnambia, a flight- less genus of dung beetles (Coleoptera) also known from New Caledonia, and the stag beetle Sphaeno- gnathus sp. (Coleoptera) unknown outside South America. 53 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 The formation of the mountain ranges with high rainfall has formed a wide variety of isolated lotic hab- itats, which over a long period may have enhanced the biodiversity of pleuston bugs in New Guinea. ACKNOWLEDGEMENTS The generous assistance of Mr. R. N. B. Prior, late- ly residing in West New Britain for the extensive col- lections he made on my behalf, especially from the Oro Province. His help during a short visit to Kimbe, W. N. B. was of great help to me. The British Museum (Natural History), London (BMNH) and the Bernice P. Bishop Museum, Honolulu (BPBM) kindly loaned material for this review. REFERENCES Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera, Gerromorpha). Phylogeny, adaptations, biogeography and classification. — Entomonograph 3: 1-455. Cheesman, L. E., 1943. Japanese Operations in New Guinea. — The Geographical Journal 101 (3); 98-110. Hale, H. M., 1925. Results of Dr. E. Mjoberg’s Swedish Scientific Expedition to Australia 1910-1913. The Aquatic and Semi-Aquatic Hemiptera. — Arkiv för Zoologi 17A (20): 1-19. Hale, H. M., 1926. Studies in Australian Aquatic 54 Hemiptera No. VII. — Records of the South Australian Museum 3: 195-217. Kikkawa, J., Monteith, G. B. & G. Ingrams, 1981. Cape York Peninsula: Major region of faunal interchange. p. 1697-1736. —In: A. Keast (Ed.), Ecological biogeography of Australia. Junk, The Hague. Kirkaldy, G. W., 1901. On some Rhynchota, principally from New Guinea, (Amphibicorisae and Notonectidae). — Annali del Museo Civico di Storia Naturale G. Doria 20: 804-810. Kirkaldy, G. W., 1908. Memoir on a few heteropterous Hemiptera from eastern Australia. — Proceedings of the Linnean Society of New South Wales 32: 768-788. Lundblad, O., 1933. Zur Kenntnis der aquatilien und sem- iaquatilen Hemipteren von Sumatra, Java und Bali. — Arkiv fiir Hydrobiologie, Supplement 12: 1-195, 263- 489, pls. 1-21, figs. 142. Lundblad, ©., 1936. Die altweltichen Arten der Veliidengattungen Rhagovelia und Tetraripis. — Arkiv för Zoologi 28A: 1-63. Polhemus, J. T. & D. A. Polhemus, 1988. Zoogeography, ecology, and systematics of the genus Rhagovelia Mayr (Heteroptera: Veliidae) in Borneo, Celebes, and the Moluccas. — Insecta Mundi 2: 161-230. Smith, A. G. & J. C. Briden, 1977. Mesozoic and Cenozoic Palaeo-continental maps. — Cambridge University Press; Cambridge: 63 pp. Received: 21 October 1992 Revised version accepted: 6 April 1993 WAYNE N. MATHIS' & TADEUSZ ZATWARNICKT ' Smithsonian Institution, Washington, D. C., USA ? Academy of Agriculture, Wroclaw, Poland REVISION OF THE SHORE-FLY GENUS CHLORICHAETA BECKER (DIPTERA: EPHYDRIDAE) Mathis, W. N. & T. Zatwarnicki, 1993. Revision of the shore-fly genus Chlorichaeta Becker (Diptera: Ephydridae). — Tijdschrift voor Entomologie 136: 55-76, figs. 1-53. [issN 0040- 7496]. Published 1 July 1993. Chlorichaeta Becker, an Old-World genus of shore flies of the tribe Gymnomyzini, is revised. Included are seven species of which C. africana (Kenya: Lake Victoria), C. mais (Seychelles: Aldabra), and C. orba (Sri Lanka: Nilaveli) are newly described and C. gioiellae Canzoneri & Rampini is determined to be conspecific and the junior synonym of C. villiersi Canzoneri & Meneghini. Correspondence: Wayne N. Mathis, Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C. 20560, USA. Key words. — Diptera; Ephydridae; Gymnomyzinae; Chlorichaeta; revision; new species; Old World. Shore flies of the genus Chlorichaeta Becker are known only from the Old World, mostly where warmer climates prevail, such as around the Mediterranean Sea or in tropical regions of Africa and the Orient. Only one species, C. albipennis (Loew), is known to occur in the more temperate regions of the Palearctic Region (Cogan 1984). The genus has nev- er been treated on a worldwide basis, and that defi- ciency, in part, prompted the research that resulted in this revision. There are few species in Chlorichaeta, and these are poorly known except to specialists. The history of previous work on the genus, as a consequence, is rela- tively brief and straightforward. Becker (1922) pro- posed Chlorichaeta towards the end of his career as a dipterist, and in the same paper, he described C. tu- berculosa, which became the type species of the genus by monotypy. Becker’s first and finest published work on shore flies was his monograph on Palearctic species (1896), which included Gymnopa albipennis, a species Loew (1848) described in his first paper on shore flies. This species remained in Gymnopa or its senior synonym, Mosillus Latreille, until Cresson (1925) transferred it to Chlorichaeta. The genus com- prised only these two species until the 1940’s when Duda (1942) described Strandilus strandi in a pecu- liar paper in which each new genus- or species-group name, which numbered five and 19 respectively, was named after Embrik Strand (1876-1947) except where a homonym would result. Papp (1979) studied Duda’s collection, now mostly housed in Berlin (ZMHU), and found that the primary types of S. stran- di (a lectotype was then designated) are conspecific with Gymnopa albipennis with the latter being the senior synonym. Cresson (1946), as part of a synopsis of Afrotropical shore flies, described C. aerifer from specimens collected in Angola. After another gap of two decades, Canzoneri & Meneghini (1969a) de- scribed C. villiersi from specimens collected in Senegal and published the first figures of male termi- nalia. In another publication, a well-illustrated faunal treatment of shore flies occurring in Italy, Canzoneri & Meneghini (1983) also provided a synopsis of C. albipennis. Finally, Canzoneri & Rampini (1990) de- scribed C. gioiellae from specimens collected in Sierra Leone. Nothing is known about the immature stages, and what little we know about the natural history comes from brief annotations that were included as part of taxonomic reviews or listings (Cresson 1946). Apart from entries in regional catalogues (Cogan 1980, 1984, Mathis 1989) little other work has been published on the genus. In addition to revising the species of Chlorichaeta, we recharacterize the genus within the context of phy- logenetic studies we are conducting on the tribe Gymnomyzini Latreille. Gymnomyzini include sev- eral genus-group taxa (Mathis et al. 1993), of which we have revised Placopsidella Kertész (Mathis 1986), the western Palaearctic species of Athyroglossa Becker (Mathis & Zatwarnicki 1990), and Mosillus Latreille (Mathis et al. 1993). Other species revisions of gym- nomyzine genera are anticipated. METHODS The methods used generally in this study were ex- 55 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 plained previously (Mathis 1986, Mathis & Zatwar- nicki 1990). Because specimens are small, less than 3.50 mm, study and illustration of the male termina- lia required use of a compound microscope. Although we have followed the terminology for most structures of the male terminalia that other workers in Ephydridae have used (see references in Mathis 1986, and Mathis & Zatwarnicki 1990a, 1990b), Zatwarnicki (1992) advocated usage of the term dististylus rather than surstylus, largely based on an hypothesis advanced by Hennig (1936). The mer- its of Hennig’s hypothesis are still being debated, and here we continue to use surstylus. In papers for which Zatwarnicki is first author, however, dististylus will be used. Three ratios (two venational) are used commonly in the descriptions and are defined here for the con- venience of the user (ratios are averages of three spec- imens): 1. Eye-to-cheek ratio: Genal height / eye height. Measurements are taken from the head in lateral view. 2. Costal vein ratio: the straight line distance between the apices of R,,, and R,,,/ distance between the apices of R, and R,,,. 3. M vein ratio: the straight line distance along M between crossveins (r-m and dm-cu) / distance apicad of crossvein dm-cu. Although this study was based in large part on specimens in the U. S. National Museum of Natural History, numerous others were borrowed, particular- ly type specimens of the species previously described. To our colleagues and their institutions listed below who lent specimens, we express our sincere thanks. Without their cooperation this study could not have been completed. ANSP — Academy of Natural Sciences of Philadelphia, Pennsylvania (J. K. Gelhaus and Donald Azuma); BMNH — British Museum (Natural History), London, England (Brian Pitkin); BBM — Bernice P. Bishop Museum, Honolulu, Hawaii (N. L. Evenhuis); Cas — California Academy of Sciences, San Francisco, California (P. H. Arnaud, Jr.); CANZ — Personal collection of Silvano Canzoneri, Venice, Italy; MNHN — Muséum National d'Histoire Naturelle, Paris, France (Loïc Matile); NMW — Naturhistorisches Museum, Vienna, Austria (Ruth Contreras-Lichtenberg); NMWL — National Museum of Wales, Cardiff, Wales, United Kingdom (J. C. Deeming); TAU — Tel Aviv University, Tel Aviv, Israel (A. Freidberg); USNM — former United States National Museum, collections in the National Museum of Natural History, Smithsonian Institu- tion, Washington, D. C.; ZMHU — Zoologisches Museum, Humboldt Universität, Berlin, Germany (H. Schumann). 56 SYSTEMATIC ACCOUNT Genus Chlorichaeta Becker Chlorichaeta Becker, 1922: 73. — Type species: Chlorichaeta tuberculosa Becker, 1922, monotypy; Cogan and Wirth 1977: 322 [Oriental catalogue], Cogan 1980: 656 [Afrotropical catalogue]; 1984: 127-128 [Palearctic cata- logue], Canzoneri & Meneghini 1983: 227-229 [diagno- sis]. Strandillus Duda, 1942: 10. — Type species: Strandillus strandi Duda, 1942 (= Gymnopa albipennis Loew, 1848), monotypy; Papp 1979: 98 [synonymy]. Diagnosis Small to medium-sized shore flies, length 1.50- 3.10 mm, mostly black, many surfaces subshiny to shiny; dorsum, especially of thorax, somewhat micro- tomentose, appearing subshiny to dull. Head: Setation generally poorly developed; fronto- orbital setae greatly reduced or lacking; ocellar setae short, inserted laterad of anterior ocellus and widely separated; both inner and outer vertical setae present, short. Frons generally smooth, shiny, ocellar triangle of mesofrons sometimes differentiated. Arista com- paratively short, subequal to length of combined first 3 antennal segments, appearing bare or bearing sever- al, very short hairs along dorsal surface, length of hairs subequal with basal aristal width. Face symmetrically pitted around midfacial protuberance, pits with sil- very white microtomentum, making face appear spot- ted; midfacial conical protuberance not prominent; parafacials immediately adjacent to face with a verti- cal row of shallow, horizontal grooves; portion of par- afacial near anteroventral margin of eye with some sil- very white microtomentum; gena generally bare and high, height 2/3 to 3/4 eye height; posterior margin sharply angulate and marginate. Thorax: Mesonotum mostly black, shiny, fre- quently with metallic luster, especially scutellum; mesonotal surface appearing granulose to various de- grees. Notopleuron with | seta inserted near postero- ventral angle; apical scutellar setae arising from tu- bercles; intra-alar seta 1; lacking row of prominent setae between postalar seta and base of scutellum; coloration of halter variable. Wing milky white; vein R,,, long, with length of costal section II not quite twice section III; vein CuA, not reaching wing mar- gin; alula wide alular marginal setulae much shorter than alular width. Fore femur much more swollen than mid or hind femora, produced ventrally to a somewhat pointed ridge near midlength, process bearing a row of 4-5 stout setae along anterior half of posteroventral margin, seta arising from ventral, midlength point conspicuously larger; mid tibia black, somewhat rounded (lacking conspicuously flattened surfaces), lacking silvery-white microto- mentum on anterior surface. Basitarsomere colour variable, apical 2-3 tarsomeres usually blackish brown. Abdomen: Dorsal surface of terga black, shiny, more finely granulose than mesonotum, lacking bands of microtomentum; terga well sclerotized, lat- eral margins continued laterally and ventrally; sterna of male relatively poorly developed, usually as small sclerotized rectangular plates, Ist sternum of male oriented perpendicular to plane of body, sterna 2-4 parallel to plane of body, 5th sternum as 2 sternites, these oriented to form a V, with vertex anteriad, sometimes fused at vertex; 5th tergum exposed but shorter than 4th, usually triangular or trapezoidal, with 2 dorsal pits toward posterior margin. Male ter- minalia: Epandrium in lateral view narrowed dorsal- ly, expanded laterally, in posterior view widest at lev- el of cerci; cerci oval, bearing short setae; hypandrium more or less U or V-shaped with posterolateral por- tion relatively long (representing fused distal portion of pregonite to hypandrium) and anteromedial por- tion forming vertex variously developed; surstyli gen- erally elongate, subequal in length to that of cerci, broadly fused basally with ventral margin of epandri- um, becoming tapered ventrally, pointed apically, ventral portion generally curved anteriorly to some degree; aedeagal apodeme more broadly developed in lateral view toward end that attaches to hypandrium; pregonite apparently fused with hypandrium; post- gonite linear, rod-like, setulose; aedeagus with dis- tinct basiphallus, distiphallus, and a sclerotized rod (ejaculatory apodeme) attached by a membranous duct to the base of the basiphallus; basiphallus well sclerotized, simple, tube-like; distiphallus more com- plex, with 2 sclerotized rods basally, apex membra- nous, in some species bearing 1-2 paired extended lat- eral processes and 1 unpaired ventral process, these sometimes sclerotized, all processes arising near base of distiphallus. Distribution Old World in tropical and warm temperate zones. Afrotropical, Australasian, Oceanian, Oriental and Palearctic (Japan, Lithuania, and Mediterranean) re- gions. Natural history Species of Chlorichaeta apparently feed as general scavengers in concentrated organic material, such as manure of terrestrial vertebrates. Cresson (1945: 53) reported that C. tuberculosa bred in rotten oil cake manure in Madras, India, and that the dissemination of the manure may account for the wide distribution of this species. On atolls of the Aldabra Group, C. mais, a new species described below, occurred in MarHIs & ZATWARNICKI: Revision of Chlorichaeta several habitats. We collected it most commonly on tortoise droppings but also found it around pig pens near the settlement (Picard Island) and occasionally on rotting seaweed that had accumulated at the strand line on beaches. Adults are attracted to moisture on animals and are frequently associated with livestock (camels, cattle, donkeys, pigs). Suppurating sores on livestock are es- pecially attractive (Cresson 1946: 244), and in the Negev Desert of Israel, Amnon Freidberg and W. N. Mathis observed that adults of C. tuberculosa were at- tracted to moisture, especially perspiration on bare legs. Cresson (1946) also noted that a specimen was found on a coccid (Homoptera) on the trunk of a China-tree in Angola (this occurrence may be inci- dental and of little significance). Diagnosis and discussion of phylogenetic relationships Chlorichaeta is similar and closely related to Mosillus Latreille and a group of species related to ‘Gymnopa beckeri Cresson’. The genus is more dis- tantly related to Placopsidella and possibly to Chaetomosillus. The monophyly of Chlorichaeta, Mosillus, and the species related to ; Gymnopa becker? (hereafter called the Mosillus clade) is established by the following synapomorphies: 1. The hypandrium, especially the lateral arms, is fused with the pregonites, a character unique to this clade. 2. The fore femur bears 5-8 stout setae, with the first one or two being the largest. In most genera of Gymnomyzini the fore femur is either unarmed or in a few genera the fore femur is produced posteroven- trally, usually forming a pointed, narrow ridge (Chaetomosillus and Stratiothyrea) or there is 1-2 stout setae (Athyroglossa). 3. The anterior margin of the hypandrium is pro- duced anteriorly to form a Y-shaped sternite with the two lateral arms. In other genera the hypandrium is truncate to broadly rounded, but not produced ante- riorly. 4. The ocellar setae are reduced or lacking. In oth- er genera of Gymnomyzini the ocellar setae are dis- tinct even though varying in size from small to well developed. Chlorichaeta is distinguished from related genera of the tribe Gymnomyzini by the synapomorphies not- ed for the clade and the following characters: body mostly shiny black with reduced setation; face pitted, appearing spotted, and with silvery white microto- mentum in pits; mid facial conical protuberance small; parafacials with a vertical row of furrows; apical scutellar setae usually arising from basal tubercles; wing lacteous; alula of wing broad, auriculate along 5% TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-5. Male genitalia of Chlorichaeta aerifer. — 1, epandrium and cerci, posterior view; 2, idem, lateral view; 3, internal male genitalia, lateral view; 4, idem, ventral view; 5, hypandrium and postgonites, ventral view. 58 inferior margin, strongly convex; fore femur bearing a large ventral seta at ventral prominence near midlength and with 5-6 short but stout setae between largest seta and apex along posteroventral margin; fifth tergum of male and to a lesser degree in female with 2 small, dor- sal depressions towards posterior margin. Key to species groups and species of Chlorichaeta 1. Scutellum with brilliant, golden-bronze luster, distinctly contrasted with blackish scutum; dis- tiphallus short, angulate, lacking long, paired processes at base (the 4erifer group) .................. y) — Scutellum and scutum either essentially concol- orous, mostly blackish, or scutum slightly golden bronze and scutellum blackish; distiphallus elon- gate, mostly straight or gently curved, bearing 2, paired, basal processes (the albipennis group) ...3 2. Parafacials with 3rd silvery-white area narrowly linear (sometimes narrowly connected dorsally with 2nd spot) at anteroventral margin of eye; posteroventral process of postgonite elongate, HTT combo ey (COS TE IONI. C. villiersi Canzoneri & Meneghini — Parafacials with 3rd silvery-white area elongate, lunate or shallowly triangular along anteroventral margin of eye; posteroventral process of postgon- ite relatively short (fig. 3) … … C. aerifer Cresson 3. Hind basitarsomere yellow, sometimes slightly darker basally, mostly concolorous with tarso- mers Dana ee 4 — Hind basitarsomere almost entirely dark colour- edublackishi brown to brown... 5 4. Extended ocellar triangle granulose, contrasted with shiny and smooth parafrons; silvery-white microtomentum on parafacials as 3 patches: 2 spots near level of antenna (dorsal spot at level of ventral margin of pedicel, ventral spot at level of midheight of Ist flagellomere) and a more linear patch at anteroventral margin of eye; mesonotum distinctly microsculptured, appearing granulose, with some metallic bronzish-gold luster ............. RO MI Ra C. mais sp. n. — Extended ocellar triangle shiny and smooth, sim- ilar to parafrons; silvery-white microtomentum on parafacials as 2 patches: 1 spot at level of ped- icel and a long linear patch at anteroventral mar- gin of eye; scutum very lightly microsculptured, scutellum more so, appearing granulose; mesono- tum with some metallic bluish luster .................. RR MR nn C. orba sp. n. 5. Scutellum bearing 3-5 large, marginal setae on each side that arise from distinct tubercles; disc of scutellum distinctly more microsculptured than Seutumappearinetsranulose Renn Matuis & ZATWARNICKI: Revision of Chlorichaeta — Scutellum bearing 2 large, marginal setae on each side that arise from tubercles (1-2 smaller setae usually between larger setae); scutum and scutel- lum about equally microsculptured . 6 6. Surstyli in lateral view distinctly and more or less evenly curved anteroventrally, tapered gradually and evenly to acutely pointed apex...................... BO o Sek SN GM se dors de C. tuberculosa Becker — Surstyli in lateral view more shallowly curved an- teroventrally and with apex slightly recurved and somewhat spatulate … … C. africana sp. n. THE aerifer GROUP Diagnosis This species group is distinguished from other taxa of Chlorichaeta by the following combination of char- acters (synapomorphies, which provide evidence of the group’s monophyly, are indicated by an asterisk): mesonotum appearing granulose, microsculptured, shiny; *scutellum except along base with brilliant, golden-bronze luster, distinctly contrasted with blac- kish scutum, disc of scutellum distinctly rugose, with rugae aligned longitudinally; larger 4-5 setae along each side scutellum arising from tubercles; *distiphal- lus short and lacking long, paired processes at base. Both species of this group occur in the Afrotropical Region, principally in countries along the west coast. Chlorichaeta aerifer Cresson (figs. 1-6) Chlerichaeta aerifer Cresson, 1946: 245. Holotype 9, Angola: Makussi, July 23, 1931, J. Ogilvie (Cresson stat- ed that the holotype is deposited in the BMNH , but on vis- its there, the holotype was not found, nor did we find it at the ANSP); Cogan 1980: 656 [Afrotropical catalogue]. Diagnosis This species and C. villiersi are sister species, as es- tablished by the synapomorphies noted for the species group (see characters noted above and in key). It is distinguished from C. villiersi by the short projection of the postgonite and the more flared apex of the sur- stylus. In specimens of C. villiersi the postgonal pro- jection is long, finger-like, and the apices of the sur- styli are parallel. Description Small to medium-sized shore flies, length 1.85 to 3.10 mm. Head: Mesofrons, especially extended ocellar trian- gle, entirely smooth, shiny, similar to parafrons; para- 59 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 6. Distribution map for Chlorichaeta aerifer (dots) and C. villiersi (diamonds). 20 facials with silvery-white to white microtomentum arranged as 3 spots, Ist at level of pedicel, 2nd at lev- el of first flagellomere, 3rd along anteroventral mar- gin of eye, distinctly wider at middle, lunate or sometime shallowly triangular. Gena very high, eye- to-cheek ratio averaging 0.75. Thorax: Mesonotum finely microsculptured, ap- pearing granulose, scutellum more so; scutum black with faint metallic bluish reflections; scutellum with base 1/8 black, remainder brownish gold with metal- lic reflections; marginal scutellar setae 4-6 on each side that arise from tubercles. Halter white. Costal vein ratio averaging 0.50; vein M ratio averaging 0.48. Fore basitarsomere reddish orange, other fore tarsomeres becoming darker, apical 2 black; hind bas- itarsomere black. Abdomen: Male terminalia (figs. 1-5): Length of epandrium+surstyli in posterior view 1.3 X width; lateral margins of surstyli in posterior view distinctly concave ventrally (fig. 1); surstylus in lateral view di- gitiform, moderately rounded apically, distinctly curved anteroventrally at basal merger with ventral margin of epandrium (fig. 2); hypandrium in lateral view sinuous basally, in posterior view broadly V- shaped, emarginate ventrally; aedeagal apodeme only 60 slightly enlarged toward end that attaches to hypan- drium (fig. 3); postgonite with a posteroventral, short process (fig. 3), in posterior view becoming broader toward apex, and apex concave; distiphallus distinctly angulate, dorsal angle acutely produced, pointed; bearing 1 short, lateral process that is broadly at- tached toward base (figs. 3, 4), basoventral margin of process irregularly serrate (figs. 3, 4). Other specimens examined Afrotropical: Angola. Luanda, 22 Aug 1949, Malkin (18, 39; USNM). Cameroon. Kribi (beach), 28-29 Nov 1987, F. Kaplan (4d, 49; USNM); Kribi, Rt. N7 (beach), 28-29 Nov 1987, A. Freidberg (ld, 29; USNM). Nigeria. Lagos (shore), 15 Dec 1987, A. Freidberg (26; USNM); Lagos, Victoria Island (shore), 15 Dec 1987, F. Kaplan (36, 29; USNM). Sa Tomé and Principe. Sao Tomé (sandy be- ach), G.R. Gradwell, D. Snow (66, 122; BMNH ). Distribution (fig. 6) Afrotropical. Angola (Luanda, Makussi), Came- roon, Nigeria, and Sao Tomé and Principe. MATHIS & ZATWARNICKI: Revision of Chlorichaeta Chlorichaeta villiersi Canzoneri & Meneghini MUSEUM PARIS 2-X-61 [2 Oct 1961] A. Villiers [date and (figs. 6-11) collector handwritten, blue]/sur Alternanthera [hand- written]/HOLOTYPUS d Chlorichaeta villiersi sp. n. [gen- Chlorichaeta villiersi Canzoneri & Meneghini, 1969a: 1481. der, name, and ‘sp. n.’ handwritten, red}. The holotype Holotype d: ‘Sénégal Kayar [Kayar handwritten]/- is glued to a card, is in very poor condition (specimen Figs. 7-11. Male genitalia of Chlorichaeta villiersi. — 7, epandrium and cerci, posterior view; 8, idem, lateral view; 9, internal male genitalia, lateral view; 10, idem, ventral view; 11, hypandrium and postgonites, ventral view. 61 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 glued upside down, obscuring dorsum of head and tho- rax; left wing, forelegs, and abdomen missing), and is de- posited in the MNHN; Cogan 1980: 656 [Afrotropical cat- alogue], Canzoneri 1981: 203 [list, Senegal]; 1982: 57 [list, Sierra Leone]. Chlorichaeta gioiellae Canzoneri & Rampini, 1990: 112. — Holotype d : ‘Sierra Leone[,] W.Area-Spiaggia[,] Sussex - 9.XI.87 [9 Nov 1987] Leg. Rossi/ HOLOTYPUS d Chlorichaeta gioiellae nov. det. Canzoneri S. [gender, spe- cies name, and ‘nov.’ handwritten, red]’. The holotype is double mounted (minuten in a rectangular block of plas- tic foam), is in fair condition (left mid leg missing), and is deposited in CANZ. Syn. n. Diagnosis This species and C. aerifer are closely related and very similar but the former is distinguished by the longer, finger-like projection of the postgonite. Description Moderately small to medium-sized shore flies, length 2.10 to 3.00 mm. Head: Mesofrons, especially extended ocellar trian- gle, entirely smooth, shiny, similar to parafrons; para- facials with silvery-white to white microtomentum arranged as 3 spots, Ist at level of pedicel, 2nd at lev- el of first flagellomere, 3rd along anteroventral mar- gin of eye, distinctly wider at middle, lunate or some- time shallowly triangular. Gena very high, eye-to-cheek ratio averaging 0.65. Thorax: Mesonotum finely microsculptured, ap- pearing granulose, scutellum more so; scutum black with faint metallic bluish reflections; scutellum with base 1/8 black, remainder brownish gold with metal- lic reflections; marginal scutellar setae 4-6 on each side that arise from tubercles. Halter white. Costal vein ratio averaging 0.49; vein M ratio averaging 0.47. Fore basitarsomere reddish orange, other fore tarsomeres becoming darker, apical 2 black; hind bas- itarsomere black. Abdomen: Male terminalia (figs. 7-11): Length of epandrium+surstyli in posterior view 1.5 X width; lateral margins of surstyli only slightly concave ven- trally (fig. 7), in lateral view digitiform, narrowly rounded apically, curved anteroventrally but at same curvature of epandrium (fig. 8); hypandrium in later- al view sinuous basally, in posterior view broadly V- shaped, rounded ventrally; aedeagal apodeme dis- tinctly enlarged on 2/3 portion toward attachment with hypandrium (fig. 9); postgonite with a postero- ventral, relatively long, narrow, somewhat finger-like process (fig. 9), in posterior view curved medially to- ward apex, and apex acutely pointed; distiphallus dis- tinctly angulate, dc sal angle acutely produced, rounded (fig. 9), produced dorsolaterally at base, ap- pearing wing-like in posterior view (fig. 10). 62 Other specimens examined Afrotropical: Gambia. Bakau at tropic bungalow, (me- adow rich in flowers at beach), 16-18 Nov 1977, L. Cederholm, R. Danielsson, Hammarstedt, Hedqvist, Samuelsson (296 , 47 ; BMNH , ZIL, USNM). Senegal. Dakar N’Gor, sabbia nuda e talitr., 24 Jun 1973, A.G. Soika (34, 1; USNM); Dakar N’Gor, spiaggia sabb.-limo, 24 Jun 1973, A.G. Soika (19; USNM). Somone, spiaggia terr. crabs, 6 Jul 1973, A.G. Soika (46, 19; USNM). Distribution (fig. 6) Afrotropical. West Africa: Gambia, Senegal, Sierra Leone. Remarks We directly compared the primary types of C. vil- liersi and C. gioiellae and are of the opinion that they are conspecific. The structures of the male terminalia are virtually identical, and we are thus proposing the synonymy noted above, with C. villiersi as the senior synonym. THE albipennis GROUP Diagnosis This species group is distinguished from other taxa of Chlorichaeta by the following combination of char- acters (synapomorphies, which provide evidence of the group’s monophyly, are indicated by an asterisk): mesonotum appearing granulose, microsculptured, shiny; *distiphallus long and bearing long, paired processes at base. Chlorichaeta africana sp. n. (figs. 12-17) Type material. — Holotype d: “KENYA Lake Victoria 10km S Kisumu 19.X1.1986 [19 Nov 1986] A. Freidberg’; 13 paratypes (78, 6%; TAU, USNM) bear the same label data as the holotype. Other paratypes are as follows: Cameroon. Kribi (beach), 28-29 Nov 1987, F. Kaplan (26, 72; TAU, USNM); Kribi, Rt. N7 (beach), 28-29 Nov 1987, A. Freidberg (16, 49; USNM). Ghana. Kedzi-Keta (on coco- nut), 29 Jul 1959 (Min. Agric. C.I.E. Coll. 16732 8377; 43,62; BMNH). Nigeria. Lagos, Victoria Island (shore), 15 Dec 1987, F. Kaplan (ld; USNM). The holotype is double mounted (minuten in a block of polyporus), is in excellent condition, and is deposited in the USNM. Diagnosis This species is very similar to C. tuberculosa in having a black hind basitarsomere but is distinguis- hed from the latter by characters of the male termi- nalia, especially the shape of the surstylus, which, in lateral view, is shallowly recurved apically and MATHIS & ZATWARNICKI: Revision of Chlorichaeta Figs. 12-16. Male genitalia of Chlorichaeta africana. — 12, epandrium and cerci, posterior view; 13, idem, lateral view; 14, internal male genitalia, lateral view; 15, idem, ventral view; 16, hypandrium and postgonites, ventral view. has a bluntly rounded apex. Description Small to moderately small shore flies, length 1.60- 2.60 mm. Head: Mesofrons, especially extended ocellar trian- gle, smooth, shiny, very similar to parafrons; parafa- cials with silvery-white to white microtomentum ar- ranged as 2 patches, dorsal patch much smaller, a spot laterad of pedicel, ventral patch long, linear, length equal to combined length of pedicel and flagellomere 1. Gena high, eye-to-cheek ratio averaging 0.60. Thorax: Mesonotum microgranulose, scutum with some faint metallic golden luster; scutellum with sim- ilar microsurface as scutum, black, lacking metallic luster, bearing 2 pairs of marginal setae that arise from tubercles. Halter pale, whitish yellow to yellow. Costal vein ratio averaging 0.71; vein M ratio averag- ing 0.58. Fore tarsi black; hind basitarsomere entirely dark coloured, blackish brown to brown. Abdomen: Male terminalia (figs. 12-16): Length of epandrium+surstyli in posterior view 1.65 X width; lateral margins of surstylus in posterior view distinct- ly though shallowly concave ventrally (fig. 12), base of surstylus wide then distinctly narrowed at apical 63 Ty DSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 17. Distribution map for Chlorichaeta africana. 1/3, apex less than greatest width of cercus; surstylus in lateral view digitiform, base in same curvature as epandrium, apex slightly recurved, narrowly spatulate (fig. 13); hypandrium short, L-shaped in lateral view with longer arm slender and more or less parallel with basiphallus, lateral portion (= fused pregonite) more or less as a parallelogram, more than twice width of longer arm (fig. 14); aedeagal apodeme moderately enlarged toward attachment with hypandrium (fig. 14); postgonite evenly lunate in lateral view (fig. 14); distiphallus moderately long, slightly longer than bas- iphallus, gently angulate in lateral view, lateral projec- tions at base of distiphallus long, length equal to nar- rowed portion of aedeagal apodeme (fig. 15), basoventral margin of process distinctly and irregular- ly serrate (figs. 14, 15). Distribution (fig. 17) Afrotropical. Cameroon, Ghana, Kenya, and Nigeria. Etymology The species epithet, africana, refers to the conti- nent on which this species was collected. Chlorichaeta albipennis Loew (figs. 18-23) Gymnopa albipennis Loew, 1848: 14. Syntypes from Italy. Sicily: Messina [ZMHU, apparently lost]. Mosillus albipennis; Schiner 1864: 235 [generic combina- tion]. Chlorichaeta albipennis; Cresson 1925: 235 [generic combi- nation, review]; 1946: 244 [review], Cogan and Wirth 1977: 322 [Oriental catalogue], Cogan 1980: 656 [Afrotropical catalogue]; 1984: 128 [Palearctic cata- logue], Canzoneri & Meneghini 1983: 229-230 [review, Italy], Mathis 1989: 640 [Australasian/Oceanian cata- logue]. Strandillus strandi Duda, 1942: 10. Lectotype d (designat- edi ibys Pappy 197997) sande smc: Duda/[Lithuania] Nidden [= Nida on Kurskaya Kosa Peninsula] 4. 8. 40 [4 Aug 1940]/Lectotypus [red]’. The abdomen of the lectotype has been removed, dissected and is in a plastic microvial attached to the specimen pin. The lectotype is deposited in the ZMHU. — Papp 1979: 97 [synonymy]. Mosillus subsultans of authors, not Fabricius (misidentifica- tion); Miyagi 1977: 27. Diagnosis The species is similar to congeners of the a/bipennis group but may be distinguished from them by the fol- lowing combination of characters: scutellum bearing 3-5 large, marginal setae on each side that arise from distinct tubercles; disc of scutellum distinctly more microsculptured than scutum, appearing more granu- lose; hind basitarsus black; characters of the male ge- nitalia (see description below and figs. 18-22). Figs. 18-22. Male genitalia of Chlorichaeta albipennis. — 18, epandrium and cerci, posterior view; 19, idem, lateral view; 20, internal male genitalia, lateral view; 21, idem, ventral view; 22, hypandrium and postgonites, ventral view; 23, Distribution map for Chlorichaeta albipennis. 64 Maris & ZATWARNICKI: Revision of Chlorichaeta 65 ‘TijDSCH RIET VOOR ENTOMOLOGIE, VOLUME 136, 1993 Description Small to moderately small shore flies, length 1.50 to 2.85 mm. Head: Mesofrons, especially extended ocellar trian- gle, shiny and smooth, similar to parafrons; silvery- white microtomentum of parafacials as 2 patches: dorsal spot much smaller, placed at level of pedicel and a long linear patch at anteroventral margin of eye. Gena high, eye-to-cheek ratio averaging 0.70. Thorax: Mesonotum (colour and texture of scu- tum and scutellum; scutellum distinctly granulose, bearing 3-5 pairs of marginal setae that arise from tu- bercles. Halter dark, brown to blackish brown (pale coloured in specimens from Sri Lanka and Japan). Costal vein ratio averaging 0.60; vein M ratio averag- ing 0.58. Fore tarsi entirely black; hind basitarsomere entirely dark coloured, blackish brown to brown. Abdomen: Male terminalia (figs. 18-22): Length of epandrium+surstyli in posterior view 1.6 X width; lateral margins of surstylus in posterior view only slightly concave ventrally (fig. 18), surstylus narrow, apical 1/3 less than greatest width of cercus; surstylus in lateral view digitiform, bluntly rounded apically, curved anteroventrally but at about same curvature of epandrium (fig. 19); hypandrium in lateral view L- shaped with longer arm slender and more or less par- allel with basiphallus, lateral portion (= fused pregon- ite) more or less as a parallelogram, more than twice width of longer arm (fig. 20); aedeagal apodeme dis- tinctly enlarged toward attachment with hypandrium (fig. 20); postgonite evenly lunate in lateral view (fig. 20); distiphallus short, less than length of basiphallus, gently curved in lateral view, lateral projections at base long, length equal to narrowed portion of aedea- gal apodeme (fig. 20), basoventral margin of process distinctly and irregularly serrate (fig. 21). Other specimens examined Oriental: India. Bombay, Juhu Beach, 7 Mar 1985, K.A. Spencer (1 2; NMWL). Laos. Vientiane, 28 May 1965, P. D. Ashlock (1; BBM). Sri Lanka. Colombo: Colombo, 14 Apr 1980, W. N. Mathis (12; USNM); Colpetty (103 Galle Road), 3 Apr 1968, T. E. Halstead (138, 1725 CAs); Negombo (6 km S), 7 May 1980, L. Jayawickrema, W. N. Mathis, T. Wijesinhe (38, 52; USNM). Mannar: Mannar, 13 Apr 1968, T. F. Halstead (16, 39; CAS). Tricomalee: Mutur, 2 May 1980, L. Jayawickrema, W. N. Mathis, T. Wijesinhe (1d; USNM); Nilaveli (5 km N), 3 May 1980, L. Jayawickrema, W. N. Mathis, T. Wijesinhe (346, 199; USNM). Vietnam. Nha Trang, 17-26 Nov 1960, C.M. Yoshimoto (1 d ; BBM). Palearctic: France. Herault: Etang de Canet, stessa Straz., 27 Jul 1954, A. Giordani Soika (24; BMNH ). Italy. Sicily: Capo San Vito, 29 Aug 1982, A. Freidberg (12; USNM). Veneto: Lido Alberoni (Laguna di Venezia), sentiero dune gulf, 21 Apr-4 Oct 1958, 1961, A. Giordani Soika, Levrini (26, 29; USNM). Japan. Honshu: Osaka, 29 Aug 1954, P. H. Arnaud, Jr. (226, 229; cas, USNM). Okitsu (in cage), 27 Oct 1914, F. Muir (1 d ; BBM). Russia. Primorski Krai (Maritime Territory): Glazkovka (20 km SW Valentin), 1-2 Sep 1986, A. Ozerov (3d, 29; ZM- MU). Ukraine. Charkov: Charkov vicinity, 14 Sep 1883, Jaroshevski (22; ZISP). Distribution (fig. 23) Oriental. India, Laos, Sri Lanka, Vietnam. Palearctic. France, Italy, Japan (Hokkaido, Honshu, Kyushu), Lithuania, Russia (Far East), Ukraine. Remarks Specimens of this species were frequently misiden- tified as C. tuberculosa, and the two species are appa- rently sympatric over some of their range (see records from Italy, Canzoneri & Meneghini 1983). The specimens from Sri Lanka and Japan have pale coloured halteres but are otherwise similar, especially characters of the male terminalia. Thus we attribute the variation to be intraspecific. Chlorichaeta mais sp. n. (figs. 24-29) Mosillus albipennis; Lamb 1912: 319 [misidentification, Seychelles (Mahé)]. Type material. — Holotype d: ‘ALDABRA. Grande Terre: Anse Mais[,] 17 March 1986[,] Wayne N. Mathis. The holotype is double mounted (minuten in a block of plastic), is in good condition, and is deposited in the USNM; 36 para- types (128, 259) bear the same locality label as the holo- type. — Other paratypes are as follows: Aldabra. Grande Terre: Cinq Cases, 12-13 Mar 1986, W. N. Mathis (166, 149; USNM). Malabar: Passe Houareau, 14-18 Mar 1986, W. N. Mathis (2d, 1 2; USNM). Picard: La Gigi, 19-20 Mar 1986, W. N. Mathis (18, 19; USNM); Settlement, 15-21 Mar 1986, W. N. Mathis (76, 21 2; USNM); trail to Jellyfish Pool, 22 Mar 1986, W. N. Mathis (96, 11 9; USNM). Additional specimens. — Cosmoledo. Menai Island: Station, 26 Mar 1986, W. N. Mathis (1 2; USNM). Wizard Island: 6 Mar 1968, B. H. Cogan, A. M. Hutson (ex; BMNH). Astove. around coconut plantation, 5 Mar 1968, B. H. Cogan, A. M. Hutson (4d, 379; BMNH ). Kenya. Mombasa (150 km N), 4 Dec 1989, A. Freidberg, F. Kaplan (1d; USNM). Madagascar. Antseranana: Nosy Be, Ambatoloaka (beach), 4-7 Apr 1991, A. Freidberg, F. Kaplan (56, 42; TAU, USNM); Nosy Tanikely, 6 Apr 1991, Figs. 24-28. Male genitalia of Chlorichaeta mais. — 24, epandrium and cerci, posterior view; 25, idem, lateral view; 26, inter- nal male genitalia, lateral view; 27, idem, ventral view; 28, hypandrium and postgonites, ventral view; 29. Distribution map for Chlorichaeta mais. 66 Marais & ZATWARNICKI: Revision of Chlorichaeta 67 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 A. Freidberg, F. Kaplan (19; USNM); Nosy Komba, 6 Apr 1991, A. Freidberg, F. Kaplan (9d, 42; TAU, USNM); Ramena, 10 Apr 1991, A. Freidberg, F. Kaplan (19; USNM); Sambava Beach, 14 Apr 1991, A. Freidberg, F. Kaplan (49; USNM). Fianarantsoa: Mananjary, 26 Oct 1988, R. Van Epps (19; USNM). Toamasina: Fénérive (on beach), Dec 1955, B. Stuckenberg (1 2; USNM). Toliara: Berenty, 20 Apr 1991, A. Freidberg, F. Kaplan (1 2; USNM); Fort Dauphin (- beach), 18 Apr 1991, A. Freidberg, F. Kaplan (4d, 89; TAU, USNM); Fort Dauphin (50 km W, river), 20 Apr 1991, A. Freidberg, F. Kaplan (16, 19; USNM); Morondava (20°17°S, 44°16°E; sea level), 20 Jan 1985, F. M. Harrington, J. D. Weintraub (49; USNM); Sud-Est Sainte- Luce (10 m), Fort Dauphin, 22-24 Feb 1958, B. Stuckenberg (1d; USNM). Seychelles. Mahé (12; BMNH ). South Africa. Natal: St. Lucia Park, 7-8 Oct 1983, A. Freidberg (1 2; USNM); Umkomaas (beach), 11 Oct-9 Dec 1954, 1983, A. Freidberg, B. Stuckenberg (664, 39 ; USNM); Umhlanga Rocks, 4 Jan 1955, E. Callan (19; USNM); Umhloti (mouth), 19 Jun 1962, B. & P. Stuckenberg (19; USNM); Durban, The Bluff, Oct 1931, A. MacKie (16; ANSP). Diagnosis This species is closely related to C. orba but is dis- tinguished from that species by the granulose exten- ded ocellar triangle and three versus two parafacial microtomentose, silvery spots. Description Small to moderately small shore flies, length 1.60- 2.65 mm. Head: Mesofrons, especially extended ocellar trian- gle, granulose, contrasted with shiny, smooth para- frons; silvery-white microtomentum of parafacials as 3 patches: 2 spots near level of antenna (dorsal spot at level of ventral margin of pedicel, ventral spot at level of midheight of Ist flagellomere) and a more linear patch at anteroventral margin of eye (sometimes 2nd spot and ventral, linear patch narrowly connected). Gena high, eye-to-cheek ratio averaging 0.50. Thorax: Mesonotum with faint to conspicuous metallic bronzish-gold reflections; 2 marginal scutel- lar setae on each side that arise from tubercles, several smaller setulae between. Halter white. Costal vein ra- tio averaging 0.63; vein M ratio averaging 0.45. Basitarsomere of foreleg variable in colour, usually dark, mostly blackish, at most very slightly lighter in colour than other tarsomeres of foreleg, sometimes yellowish; hind basitarsomere yellow, sometimes slightly darker basally, mostly concolorous with tarso- meres 2 and 3. Abdomen: Male terminalia (figs. 24-28): Length of epandrium+surstyli in posterior view 1.65-1.70 X width; lateral margins of surstylus in posterior view convex ventrally (fig. 24), broad, apical 1/3 wider than cercus; surstylus in lateral view narrowly digiti- form, narrowly rounded apically, shallowly curved anteroventrally at about same curvature of epandrium (fig. 25); hypandrium in lateral view very long and sinuous, more or less in same conformation of basi- phallus, lateral portion (= fused pregonite) more or less as a parallelogram, only slightly wider than long arm (fig. 26); aedeagal apodeme distinctly enlarged toward attachment with hypandrium (fig. 26); post- gonite evenly lunate in lateral view (fig. 26); disti- phallus long, more than twice length of basiphallus, sinuous in lateral view, apex recurved posterodorsally, lateral projections at base of distiphallus long, length equal to narrowed portion of aedeagal apodeme (figs. 26, 27), basoventral margin of process not serrate (fig. 27). Distribution (fig. 29) Afrotropical. Aldabra Group (Aldabra, Astove, Cosmoledo), Kenya, Madagascar, Seychelles (Mahé), South Africa (Natal). Natural history This species occurs in several habitats on atolls of the Aldabra Group. We collected it most commonly on tortoise droppings but also found it around pig pens near the settlement (Picard Island) and occa- sionally on rotting seaweed that had accumulated at the high tide mark on the beach. Etymology The specific epithet, mais, alludes to the type loca- lity of this species, Anse Mais, on the atoll of Aldabra (Grande Terre). The species name is a noun in appo- sition. Remarks The specimens from Aldabra, including the holo- type, all have the fore basitarsomere mostly blackish, similar in colour to the other tarsomeres of the fore- leg. The states for this character are variable in speci- mens from Madagascar and South Africa, however. Some specimens have a pale, mostly yellowish basitar- Figs. 30-34. Male genitalia of Chlorichaeta orba. — 30, epandrium and cerci, posterior view; 31, idem, lateral view; 32, inter- nal male genitalia, lateral view; 33, idem, ventral view; 34, hypandrium and postgonites, ventral view; 35. Distribution map for Chlorichaeta orba. The locality in Australia (Northern Territory) could not be precisely located. 68 MATHIS & ZATWARNICKI: Revision of Chlorichaeta 69 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 somere, but in others from the same series this struc- ture is dark coloured, as in specimens from Aldabra. We have dissected the male terminalia of the variable series and see no differences and are thus of the opi- nion that this variation is intraspecific. Chlorichaeta orba sp. n. (figs. 30-35) Type material. — Holotype d': ‘SRI LANKA: TRI[NCOMAL- EE]. Dist[rict]. Nilaveli (5 km N) 3 May 1980/Collectors: W. N. Mathis[,] T. Wijesinhe[,] L. Jayawickrema’. Six par- atypes (26, 49) bear the same label data as the holotype. — Other paratypes are as follows: Sri Lanka. Colombo: Ratmalana, Colombo (9 mi S), 7-13 Jan 1962, H. Andersson, P. Brink, L. Cederholm (29; zit); Kalutara, Colombo (25 mi SSE), 25 Jan 1962, H. Andersson, P. Brinck, L. Cederholm (1 2; zi). Hambantota: Kirinda, 25 Apr 1980, W. N. Mathis (29; USNM). Uva: Heda Oya, Bibile (29 mi SE), 7 Mar 1962, H. Andersson, P. Brinck, L. Cederholm (16; ZIL). Additional specimens. — Australasian Region: Australia. Northern Territory: Wurruwardumanja, 15 Jan 1980, T. Donovan (ld; ANIC). Papua New Guinea. Central Province: Laloki (muddy river bank), 2 Jul 1984, J. W. Ismay (16; USNM). Oriental Region: Philippines. Calicoan (from dead land crab; sand near tent), 27 Jul 1945, F. F. Bibby (83, 72; USNM). Diagnosis This species is similar to C. mais in having the hind basitarsomere yellow but differs from the latter in the following characters: extended ocellar triangle smooth and shiny, similar to parafrons; parafacial with 2 patches of silvery microtomentum; scutum very lightly microsculptured, scutellum slightly more so, appearing granulose; and mesonotum with some metallic bluish luster. Description Small to moderately small shore flies, length 1.80- 2.60 mm. Head: Mesofrons, especially extended ocellar trian- gle, shiny and smooth, similar to parafrons; silvery- white microtomentum of parafacials as 2 patches: dorsal spot much smaller, placed at level of pedicel and a long linear patch at anteroventral margin of eye. Gena high, eye-to-cheek ratio averaging 0.47. Thorax: Mesonotum with some metallic bluish re- flections; scutum very lightly microsculptured, scutel- lum more so, appearing granulose, bearing 2 pairs of marginal setae that arise from tubercles. Halter yellow to yellowish white. Costal vein ratio averaging 0.60; vein M ratio averaging 0.55. Tarsi of foreleg with ba- sal 2 tarsomeres yellow, apical 3 black; basitarsomere of foreleg yellowish, contrasted with blackish apical 70 tarsomeres of foreleg; hind basitarsomere yellow, sometimes slightly darker basally, mostly concolorous with tarsomeres 2 and 3. Abdomen: Male terminalia (figs. 30-34): Length of epandrium+surstyli in posterior view 1.55 X width, lateral margins irregularly tapered toward pointed apex (fig. 30); surstylus in lateral view digitiform, nar- rowly rounded apically, curved anteroventrally but only slightly more curvature of epandrium (fig. 31); hypandrium U-shaped in posterior view (fig. 34), in lateral view L-shaped with anteromedial portion slen- der, more or less parallel with basiphallus, tapered to- ward apex, posterolateral portion (= fused pregonite) as a parallelogram, more than twice width of antero- medial portion (fig. 32); aedeagal apodeme distinctly enlarged toward attachment with hypandrium (fig. 32); postgonite linear, lacking a process (fig. 32); dis- tiphallus slightly longer than basiphallus, distinctly angulate, dorsal angle acutely produced, rounded (fig. 32), bearing 1 paired slender, lateral processes at base with anterodorsal orientation (fig. 33) and a single medial process (fig. 32). Distribution (fig. 35) Australasian: Australia (NT), Papua New Guinea. Oriental: Philippines, Sri Lanka. Etymology The specific epithet, orba, is derived from Latin and means bereft of parents or an orphan. Chlorichaeta tuberculosa Becker (figs. 36-48) Chlorichaeta tuberculosa Becker, 1922: 74. Lectotype 9 (designated by Cresson 1925: 234): ‘[Sudan. Khartoum region] Sennar 18.-24.II. [handwritten] / Aegypt. Sudan Ebner.1914/TYPE [red] / Ch. tuberculosa Beck. [handwrit- ten] det. Becker’. The lectotype is double mounted (min- ute nadel in plastic foam block), is in poor condition (mesonotum damaged, right mid leg and Ist flagello- meres missing), and is deposited in the NMW. — Cresson 1925: 234 [review]; 1945: 53 [review]; 1946: 245 [re- view], Wirth 1955: 49 [list, Tanzania. Mugango (on Lake Victoria)], Canzoneri & Meneghini 1969a: 1481 [list, Senegal, figures of & genitalia]; 1969b: 167 [list, Zaire], Cogan and Wirth 1977: 322 [Oriental catalogue], Cogan 1980: 656 [Afrotropical catalogue], Mathis 1989: 640 [Australasian/Oceanian catalogue]. Chlorichaeta albipennis of authors, not Loew (misidentifica- tion). — Collin 1949: 202 [list, Egypt, Siwa], Tenorio 1980: 267-268 [list, Hawaii]. Diagnosis This species is very similar to C. africana but is dis- tinguished from the latter and other congeners of the Maruis & Zarwarnickt: Revision of Chlorichaeta b, Figs. 36-41. Scanning electron micrographs of Chlorichaeta tuberculosa (Israel; scale length in parenthesis; bar scale for all pho- tographs = fig. 36). 36, head, lateral view (0.27 mm); 37, idem, anterior view (0.30 mm); 38, frons, anterodorsal view (176 um); 39, antennae, anterior view (136 um): 40, face, anterior view (176 lim); 41, parafacial, anterior view (120 um). TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 42-47. Scanning electron micrographs of Chlorichaeta tuberculosa (Israel; scale length in parenthesis; bar scale for all pho- tographs = fig. 44). 42, mesonotum, dorsal view (0.40 mm); 43, scutellum, dorsal view (176 um); 44, left foreleg, anterior view (150 um); 45, pleuron, lateral view (176 um); 46, notopleuron, lateral view (136 um); 47, 5th tergum of abdomen, dor- sal view (75 Um). 72 albipennis group by the following combination of characters: scutellum bearing 2 large, marginal setae on each side that arise from tubercles; scutum and scutellum about equally microsculptured and colou- red; hind basitarsomere almostly entirely brownish black to black; and surstylus in lateral view distinctly and more or less evenly curved anteroventrally, tape- red gradually and evenly to acutely pointed apex. Description Small to moderately small shore flies, length 1.65- 2.70 mm. Head (figs. 36-41): Mesofrons, especially extended ocellar triangle, smooth, shiny, very similar to para- frons; parafacials with silvery-white to white microto- mentum arranged as 2 patches, dorsal patch much smaller, a spot laterad of pedicel, ventral patch long, linear, length equal to combined length of pedicel and flagellomere 1. Gena high, eye-to-cheek ratio av- eraging 0.52. Thorax (figs. 42-46): Mesonotum mostly black, shiny, some specimens with faint, bronzish luster on scutum; scutum and scutellum with similar texture, both at most lightly granulose, bearing 2 pairs of mar- ginal setae that arise from tubercles. Halter pale, whit- ish yellow to yellow. Costal vein ratio averaging 0.64; vein M ratio averaging 0.58. Fore tarsi black; hind basitarsomere entirely dark coloured, blackish brown to brown. Abdomen (fig. 47): Male terminalia (figs. 48-52): Length of epandrium+surstyli in posterior view 1.5 X width; lateral margins of surstylus straight to concave towards apex in posterior view (fig. 48), in lateral view distinctly and evenly tapered to narrowly round- ed apex, curvature in lateral view anteroventrad, dis- tinctly more so than curvature of epandrium (fig. 49); hypandrium short, curved in lateral view with antero- medial portion more or less parallel with basiphallus and tapered toward apex, dorsal portion (= fused pregonite) as a parallelogram, more than twice width of anteromedial portion (figs. 50, 52); aedeagal apo- deme distinctly enlarged toward attachment with hy- pandrium (fig. 50); postgonite evenly lunate in later- al view (fig. 50); distiphallus longer than basiphallus, gently curved along basal 2/3 in lateral view, recurved apically, lateral projections from base long, length equal to narrowed portion of aedeagal apodeme (figs. 50, 51), at base of process with a short membranous lobe that bears a few finger-like serrations. Other specimens examined Afrotropical: Angola. Lobito Bay (on trunk of Melia aze- darach), 19 Jul-10 Aug 1931, T. D. A. Cockerall, J. Ogilvie (12, lex; ANSP). Ethiopia. Lake Langano (7°27'N, Maruis & ZATWARNICKI: Revision of Chlorichaeta 39°11’E), 8 Feb-13 Dec 1973, 1989, R. Baker, A. Freidberg, F. Kaplan (23; BMNH , USNM). Ghana. Cape St. Paul, 22 May 1958 (19; BMNH ). Kenya. Tsavo National Park, 4-7 Apr 1968, B. Cogan, A. Hutson (1d; BMNH ). Malawi. Ruo ChiKonje (on cattle), 16 Dec 1915, R. C. Wood (136, 69; BMNH , ANSP). Mozambique. Pafuri (hip- po carcass), 21 May 1983, L. Braack (24, 49; USNM). Nigeria. Cotonon, in Dahomey (70 mi W Lagos), 5 Jun 1914, W. A. Lamborn (19; ANsP). South Africa. Kruger National Park (22°27°S, 31°17’E, across river from Pafuri, on impala carcass), L. Braack (2d, 19; USNM). Sudan. Erkowit, 17 May 1943, D.J. Lewis (16, 19; BMNH ). Khartoum (sores on the backs of camels), 21 Feb-1 Mar 1932, 1933, H. W. Bedford (23, 59; BMNH , ANSP). Tuti Island (attracted to human sweat), 20 Mar 1932, H. W. Bedford (1d, 42; BMNH ). Wad Medani (on dead goat), 27 Mar 1940, D. J. Lewis (19; BMNH ). Yemen. Aden, 3 Dec 1951 (16; USNM); Kirsh, Wadi ‘Aqqan (3000 ft), 17 Mar 1940, P. W. R. Petrie (1d; BMNH ). Socotra Island: Ras Shóab, Feb 1899, O. Simony (1d; ANSP). Australasian/ Oceanian: Guam. Agana (sea cucumber), 3 Aug 1945, G. E. Bohart, J.L. Gressitt (19; ANSP); Antigua, 2 Aug 1945, G. E. Bohart, J. L. Gressitt (1 2; USNM); SE coast, 9 May 1945, G. E. Bohart, J. L. Gressitt (1d, 19; USNM); Pt Oca, May 1945, G. E. Bohart, J. L. Gressitt (19; USNM). Hawaii. Hawaii: N Kona, Makalawena Tract, 12 Feb 1970, D. E. Hardy (1d, 19; USNM). Maui: Lahaina (2 km S; on herbs and grass along sea shore), 23 Oct 1981, L. & I. Cederholm (26,39; BMNH ). Oahu: Fort Kam, 4 Jul 1944, D. G. Hall (22; USNM). Oriental: India. Madras, Sidhout (bred in rot- ting oilcake manure), 9 Mar 1934, C. S. B (32; BMNH , ANSP). Sri Lanka. W. Province: Ratmalana, Colombo (9 mi S), swept on dry grassland, or on herbs on sandy beach, 7-13 Jan 1962, H. Andersson, P. Brinck, L. Cederholm (64,89; ZIL); Kalutara, Colombo (25 mi SSE), swept on dry grass on sandy beach, 25 Jan 1962, H. Andersson, P. Brinck, L. Cederholm (16, 29; ZIL); Beruwala, Colombo (34 mi SSE), 25 Jan 1962, H. Andersson, P. Brinck, L. Cederholm (16; ZIL). Thailand. Thonburi Province: Mueng District, Manop, 7 May 1959 (12; USNM). Rajburi, Banpongnoung, 29 Jun-3 Jul 1952, R. E. Elbel (12; USNM). Vietnam. Saigon, Cho Binh Tay, 15 May 1973, R.E. Parsons (1d, 12; USNM). Palearctic: Afghanistan. Khourmalik (between Farah and Dilaram, 800 m, under stones), 6 Sep 1957, K. Lindberg (2; USNM). Algeria. Kairouan, 7 Aug, F. Santchi (1d; ANSP). Egypt. Egypt (1d; ANSP). Giza Province: Manshiet Radwan, 28 Apr 1965, K.V. Krombein (16; USNM); Siwa, 15 Jul 1935, J. Omer-Cooper (7d, 92; BMNH ); Wadi el Natrun (N Cairo; sores on donkey), 20 Jul 1950, C. W. Sabrosky (34, 52; USNM); Wadi Ghoderat Kosaima (tamarix), 1934, Priesner (16; USNM). Saqgra, 10 May 1966, P. M. Marsh (16, 22; USNM). Sinai: Bir Tmade, 8 Sep 1977, D. Simon (2d; TAU); Dahab, 7 Apr-23 May 1973, 1981, A. Freidberg, M. Kaplan, W. N. Mathis (2d, 39; TAU, USNM); ‘En Qsaib, 15 Mar 1982, I. Yarom (19; TAU); Wadi Firan (oasis), 9 Apr-29 July 1950, 1971, 1973, A. Freidberg, C. W. Sabrosky (1d, 42; TAU, USNM); 1950, C. W. Sabrosky (19; USNM); Mt. Abas, 14 Aug 1971, J. Kugler (1d, 19; Tau); Muyat-Hara, 12 Aug 1971, J. Kugler (1d, 22; TAU); Ofira (sewage area), 22 Mar-21 May 1981, A. Freidberg, W. N. Mathis (49d, 169 ; TAU, USNM); Neviot, 14 May 1981, T. Furman (1d; TAU); Ras Burka, 20 May-5 Sep 1976, 1981, A. Freidberg (1d, 12; TAU); Sinai Mountains, St. Katharina, 18 Jul 1974, F. Kaplan (1°; TAU); Sinai Mountains, Wadi Tlach (1500 m), 15 Jul 1974, Me TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 48-52. Male genitalia of Chlorichaeta tuberculosa. — 48, epandrium and cerci, posterior view; 49, idem, lateral view; 50, internal male genitalia, lateral view; 51, idem, ventral view; 52, hypandrium and postgonites, ventral view. A. Freidberg, F. Kaplan (29; TAU); Wadi Mesech, 14 Apr 1981, T. Furman (2d; TAU); Wadi Watir, 6 Apr-5 Aug 1973, 1975, A. Freidberg, M. Kaplan (15d, 129; TAU). Khor Gwob, 25 Nov 1930, R. C. M. Darling (16, 29; BMNH ). Greece. Macedonia, 1933, R. C. Shannon (1%; USNM); Larissa, Platamon, 5-9 Sep 1965, B. H. Cogan (46, 1®; BMNH ). Israel. Arad, 21 Apr 1981, F. Kaplan, A. Freidberg (35, 19; TAU); Dead Sea, 11-18 Oct 1939, E. Rivnay (56,39; ANSP, TAU); ‘En ‘Avedar, 16-29 Mar 1980, 1988, A. Freidberg, W. N. Mathis (66, 99; TAU, USNM); ‘En Bogéq, 30 Jul-20 Sep 1971, 1977, A. Freidberg, J. Kugler (16, 19; TAU); ‘Ein Fashkha (Dead Sea at ‘Enot Zugim), 2 Jul 1946, O. Theodor (58, 22; AU); ‘En Gedi, 21 Mar 1980, A. Freidberg, W. N. Mathis (89 ; USNM); ‘En Mor, 31 Mar-30 Oct 1981, 1984, 1986, A. Freidberg (7, 74 59; AU); Hameshar, 16 Mar 1988, A. Freidberg (2d; AU); Hazeva, 21 Apr 1981, A. Freidberg (ld; TAU); Jerusalem (Mt. Scopus), 9 Sep 1935, O. Theodor (2d; TAU); Kefar Yeroham, 22 Jul 1962, J. Kugler (3d, 12; TAU); Mikhmoret, 28 Aug 1980, C. Nagy (1 2; TAU); Nahal ‘Iddan, 22 Mar 1980, A. Freidberg, W. N. Mathis (20,19; USNM); Nahal Ramon, 15-30 Oct 1975, 1984, A. Freidberg (48,6%; TAU); Nahal Zibor, 17 Mar 1988, A. Freidberg, F. Kaplan (4d, 39; rau); Ne’ot ha Kikkar, 21 Mar 1980, A. Freidberg, W. N. Mathis (1d; USNM); Nizzanim, 4-12 Oct 1983, I. Nussbaum (4d; TAU); Qalya, 8 Mar 1976, A. Freidberg (1 2; AU); Taba, 28 Apr 1974, A. Freidberg (1 2; AU); Wadi Yamin, 19 Oct 1983, I. Nussbaum (1 ®; TAU). Italy. Sardinia: Rio Geremeas, 23 Jul 1952, Th.G. Aitken (93, 72; USNM). Oman. Al Khoudh, 11 Nov 1988, M. J. MarHis & ZATWARNICKI: Revision of Chlorichaeta Fig. 53. Distribution map for Chlorichaeta tuberculosa. Ebejer (1d, 1 2; NMWL); Al Seefa, 4 Dec 1987, M. J. Ebejer (19; NMWL); Ghuzayn, date culture, 10 Apr 1985, P. Ardö (18; ZIL); Nizwa-Tanuf road, 12 Apr 1985, P. Ardö (19; ZIL); Wadi Daigah, 23°06 N, 58°52’E, 27 Jan 1989, M. J. Ebejer (22; NMWL). Distribution (fig. 53) Afrotropical. Angola, Cape Verde Islands, Ethio- pia, Ghana, Kenya, Malawi, Mozambique, Nigeria, South Africa, Sudan, Tanzania (no specimens seen), Yemen (including Socotra). Australasian/Oceanian. Guam, Hawaiian Islands. Oriental. India, Sri Lanka, Thailand, Vietnam. Palearctic. Afghanistan, Algeria, Bulgaria (no specimens seen), Canary Islands (no specimens seen), Egypt, Greece, Israel, Italy, Oman. ACKNOWLEDGEMENTS We are grateful to the curators and their respective institutions for the loan of specimens that were bor- rowed in conjunction with this study (a listing of acronyms, collections, and curators are noted in the introduction of this paper). H. B. Williams prepared the distribution maps, and Susann Braden and Victor Krantz assisted with the preparation of the scanning electron micrographs. We are also grateful to David Challinor, former Assistant Secretary for Research, Smithsonian Institution, and Stanwyn G. Shetler, Deputy Director of the National Museum of Natural History, for financial support to conduct field work and study primary types through grants from the Research Opportunity Fund. Curtis W. Sabrosky hand carried the holotype of C. villiersi from MNHN to Washington, D. C. REFERENCES Becker, T. 1896. Dipterologische Studien IV. Ephydridae. — Berliner Entomologische Zeitschrift 41(2): 91-276. Becker, T. 1922. Wissenschaftliche Ergebnisse der mit Unterstiitzung der Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternom- menen Zoologischen Expedition nach dem Anglo- Agyptischen Sudan (Kordofan) 1914. VI. 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A new classification of Ephydridae based on phylogenetic reconstruction (Diptera: Cyclorrhapha). — Genus 3 (2): 65-119. Received: 25 January 1993 Accepted: 8 April 1993 THOMAS PAPE! & HIROSHI SHIMA? ‘Zoological Museum, Copenhagen, Denmark Biological Laboratory, College of General Education, Kyushu University, Japan A NEW GENUS OF TACHINIDAE FROM THE PHILIPPINES (DIPTERA) Pape, T. & H. Shima, 1993. A new genus of Tachinidae from the Philippines (Diptera). — Tijdschrift voor Entomologie 136: 77-81, figs. 1-5. [issN 0040-7496]. Published 1 July 1993. A new genus of Tachinidae, Aporeomyia gen. n., including only the type species Aporeomyia antennalis sp. n., is described from the Philippines. It is placed in the subfamily Tachininae, and possible phylogenetic relationships to other tachinine genera are discussed. Correspondence: Dr. T. Pape, Zoological Museum, Universitetsparken 15, DK - 2100 Copenhagen, Denmark. Key words. — Tachinidae, Philippines, new genus, new species. The present study deals with two male specimens belonging to the Tachinidae family-group and show- ing a tripartition of the first flagellomere. Although possibly sexually dimorphic with the female first fla- gellomere unpartitioned, this feature makes the spe- cies easily recognizable (fig. 3). Tripartition of the first flagellomere occurs within the Tachinidae fami- ly-group in the family Tachinidae and in some unde- scribed Australian—New Guinean taxa of uncertain familial affiliation (see discussion below). However, the configuration of the tripartition is never fully identical to that seen in the present species. We take this opportunity to describe and name the taxon, and as is discussed in detail below the species does not fit very well into any of the named genera, for which rea- son a new genus is described. TAXONOMIC PART Aporeomyia gen. n. Type species: Aporeomyia antennalis sp. n. Etymology. — Gender: Feminine. Composed from the type locality, Mt. Apo of Mindanao, and the Greek, oreos, oros = mountain, and myia = fly. Diagnosis. — Male: Head profile narrowed ventral- ly. Proboscis and palpus short. Antenna very large and with first flagellomere tripartite from base. Arista inserted on upper lobe at a point about 0.3 from base to tip. Antennal scape and pedicel raised above level of lunule. Prosternum bare. Subscutellum slightly swollen. Wing vein R, setulose in distal part, vein M almost straight and withour a bend distal to cross- vein dm-cu, and vein Cu+A, extended to wing mar- gin. Lower calypter narrow and with median margin diverging from scutellum. Hind tibia with postero- ventral apical seta. Aporeomyia antennalis sp. n. (figs. 1-5). Type material. — Holotype d: PHILIPPINES, Mindanao, Mt. Apo, Agko - Lake Binado, 1,350-2,300 m, 5.viii.1978, A. Nakanishi & O. Yata. — Paratype d, data as holotype, terminalia dissected and stored in a microvial pinned with the specimen. Both types are deposited in the Biological Laboratory, Kyushu University, Fukuoka, Japan. Etymology. — From the Latin, antenna = feeler, -is = adjectival suffix meaning with or having. The name refers to the first flagellomere, the peculiar shape of which makes the species easily recognizable. Description. — Male. Head brown in ground colour, without pollinosity; frontal vitta dark brown; face pale yellowish; scape and pedicel reddish, first flagellomere brown; palpus pale yellowish. Head pro- file short and high, strongly narrowed ventrally; frons about 0.4x as long as height of face profile; vertex wide, 0.47-0.50x head width; frontal vitta weakly widened anteriorly, about 3x as wide as fronto-orbital plate at middle; parafacial in profile very narrow; ge- na slightly less than 0.5x eye height, genal dilation obliquely occupying about lower 0.3 of gena; mouth opening reduced, about 0.75x as long as wide. Inner vertical seta strong, slightly less than eye height; 2 fine divergent postocellar setae; ocellar seta fine, procli- nate, subequal in length to postocellar seta; no recli- nate and proclinate orbital setae; 3-4 frontal setae, lowest seta nearly level with base of pedicel; 2-3 irreg- ular rows of short fine hairs on anterior portion of 77 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-5. Aporeomyia antennalis. — 1, right wing, dorsal view; 2, epandrium (in part), cerci and right surstylus; posterior view; 3, male head; 4, epandrium, surstylus and cercus, lateral view; 5, hypandrium, pre- and postgonite and aedeagus, later- al view. fronto-orbital plate descending to upper 0.25 of para- facial; vibrissa fine and short, slightly more than 0.3x length of face; facial ridge with several fine short hairs just above vibrissa; occiput with short black hairs. 78 - Antenna long, distinctly longer than face; first flagel- lomere tripartite, each lobe weakly widened apically, middle lobe slightly longer than others. Arista short, about 0.4x as long as upper lobe of first flagellomere, inserted on basal 0.3 of upper lobe of first flagello- mere; second aristomere about 2x as long as wide. Mouthparts strongly reduced; proboscis very short; la- bella small; palpus short, subequal in length to pedicel. Thorax brown on dorsum except for pale brownish postpronotal lobe, pale brownish on pleura except for brownish posterior 0.5 of anepisternum and anterior 0.3 of katepisternum, without pollinosity. Dorsum with rather sparse short fine hairs except for bare area between rows of dorsocentral and intra-alar setae of postsutural scutum, pleura without hairs other than strong setae; 1 postpronotal seta; 2 notopleural setae; no acr seta; 1-2 + 2-3 dc setae; 0 + 1 ia seta; 1 supra- alar seta; proepisternum bare, with 1 fine proepister- nal seta; 1 fine proepimeral seta; 2 + 2-1 katepisternal setae; katepimeron and katatergite bare; scutellum with 2 pairs of marginal setae, strong subapicals and short apicals; discal scutellar setae absent. Metatho- racic spiracle rather small, anterior and posterior fringes subequal and not standing out from spiracular rim. Wing hyaline, evenly tinged with pale brown; tegu- la and basicosta dark brown; lower calypter pale brownish yellow. Vein R,,, joins costa near wing apex, giving a long third costal sector slightly more than 4x length of second; vein R,,; joins costa slightly beyond wing apex; vein M straight from crossvein dm-cu to wing margin; last section of vein CuA, short, slightly more than 0.25 of crossvein dm-cu. Costa setulose dorsally and ventrally; vein R, setulose dorsally on distal half; vein R,,, ventrally with 1-2 setulae at base, dorsally with several setulae at base as well as setulae from just_proximal to crossvein r-m and almost to wing margin. Halter large, about 0.7x as long as low- er calypter. Legs pale brownish yellow; pulvilli whitish. Fore tibia with 1 fine p seta; mid tibia with 0 ad, 2 pd and 1 v setae. Claws and pulvilli very short. Abdomen brownish in ground colour, without pollinosity; anterior 0.4 of syntergum 1+2, anterior 0.3 of terga 3-4 and anterior 0.25 of tergum 5 pale brownish; mid dorsal stripe dark and narrow; synter- gum 1+2 rather long, only faintly excavated at base; sterna 2-3 exposed. Hairs short, fine, dense and re- cumbent on dorsum, longer and sparser on venter; tergum 2 with 2-3 lateral discal, 2 lateral marginal and 2 median marginal setae, all setae strong; tergum 3 with 2 rather fine lateral discal, 2 strong median and lateral discal setae; terga 4-5 each with 1-2 fine lateral discal setae and a row of strong marginal setae; sterna without strong setae. Male genitalia. Cercal bases fused medially, apical prongs well separated and pointed; surstylus narrow and longer than cerci; epiphallus present; distiphallus rather short, with tapering unpaired dorsomedian process. PAPE & SHIMA : New genus of Tachinidae Length. Body, 3.7-4.5 mm; wing, 3.4-4.3 mm. Female. — Unknown. Biology. — Unknown. Distribution. — Oriental (Mindanao). Region: Philippines SYSTEMATIC DISCUSSION Aporeomyia antennalis is unquestionably a member of the Tachinidae family-group or Oestroidea, the monophyly of which is discussed at length elsewhere (Griffiths 1972, McAlpine 1989, Pape 1992). Families of the Tachinidae family-group are in the main well defined, but many genera showing a com- bination of a slightly swollen subscutellum, oval or tongue-shaped lower calypteres, small metathoracic spiracles, and an obtuse (or missing) bend of vein M have been, and are still being, shuffled around amongst the tachinoid families, e. g., Mimodexia Rohdendorf (currently Rhinophoridae, Tschorsnig 1985a), Bezzimyia Townsend and Malayia Malloch (Tachinidae or Rhinophoridae, Pape 1992), and Morinia Robineau-Desvoidy (currently Calliphori- dae, Rognes 1991). Aporeomyia antennalis possesses these four character states, which may be apomorphic groundplan states of the Rhinophoridae, and it has accordingly a rhinophorid appearance. The structure of the aedeagus, however, provides no support for a rhinophorid affiliation. In the Rhinophoridae, the ae- deagus is generally slender, always equipped with a long epiphallus, and usually with paired dorsolateral processes. In Aporeomyia the aedeagus is rather short with a short and compact epiphallus, and with an un- paired, tapering dorsomedian process (‘Fortsatz des Dorsalsklerits of Tschorsnig [1985a, b]). Pape (1986: 27) recognized a single monophyletic group within the Rhinophoridae, defined by an unpaired dorsomedian process, but this group is part of a more inclusive group sharing a tripartite (and apomorphic) acrophallus not found in Aporeomyia. The ground- plan of the possibly monophyletic Sarcophagidae + Tachinidae is characterized by an unpaired dorsome- dian process (Pape 1992), but Aporeomyia does not share any synapomorphies with any of the sarcophag- id subfamilies and cannot be included in this family. It is our experience that the male terminalia in gener- al provide much more substantial evidence of phylo- genetic relationships than the above-mentioned rhi- nophorid-like features. Thus, we consider an assignment to the Tachinidae as most probable, even though we have no clearcut evidence in support of this. Fitting Aporeomyia antennalis into the subfamilial classification of the Tachinidae with no information on the structure of female terminalia, female repro- ductive system, and immatures (egg, first instar larva) 79 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 has to be preliminary. Of the four subfamilies gener- ally applied, e. g. by Tschorsnig (1985b) and Wood (1987), perhaps only the Dexiinae and the Phasiinae are monophyletic taxa. Indeed, it has been stressed that at least the subfamily Tachininae very probably is not monophyletic (‘sehr wahrscheinlich nicht monophyletisch’; Tschorsnig [1985b: 119]) or even ‘almost certainly polyphyletic’ (Crosskey 1980: 822- 823). Aporeomyia antennalis does not possess the aedeagal hinge characterizing the groundplan of the Dexiinae, nor any of the derived character states listed for the Phasiinae by Tschorsnig (1985b: 121). Also, it does not possess a setose prosternum as found in many (but not all) Exoristinae. Aporeomyia antennalis will key out to subfamily Tachininae in Crosskey’s (1976: 11) comprehensive (given as ‘alternative’) key to sub- families of Oriental Tachinidae, but mainly because it repeatedly fits the entry: “Forms without such combi- nation of characters present simultaneously’. Note that Pape (1992) used a broad concept of the Tachininae, including all species which embryonate their eggs in the uterus. With only males of Aporeomyia available, nothing can be deduced on fe- male reproductive biology. We have chosen to place Aporeomyia in the Tachininae as this does not conflict with the general morphology of this subfamily (whether in the traditional sense or the broader sense used by Pape [1992]). Assigning antennalis to a genus of its own deserves an explanation as monotypy in itself conveys little, if any, information. Often - as here - monotypic genera are erected to accommodate aberrant species which present insufficient and conflicting evidence for in- clusion within already named genera. The tripartite first flagellomere — in so far as this is of very rare occurrence in the Tachinidae — may be taken as evidence of phylogenetic relationship. Trischidocera sauteri Villeneuve is the only other Oriental species with a tripartite first flagellomere, and the head profile of Aporeomyia antennalis more- over resembles that of 7. sauteri to the extent that both have the antennal insertion at or slightly above the level of upper eye margin, an almost horizontal frons, a short lower head margin and very short mouthparts (compare Crosskey 1976: fig. 50 with the present fig. 3). Many of these features, however, may be considered constraints imposed by the very long and tripartite first flagellomere. It is important to note that the first flagellomere of A. antennalis and T. sauteri reveals some structural differences: Firstly, 7. sauteri has the first flagellomere basally bipartite with the lower lobe (i. e., that closest to the facial plate) - further bipartitioned at a point about one fifth from its base, while in A. antennalis the three lobes of the first flagellomere all originate from a common point 80 at the base — or perhaps with the middle and upper lobes sharing a very short common base. Secondly, the arista is terminal in 7. sauteri but inserted at 0.3 from the base in A. antennalis. Thus, for the triparti- tion to be homologous, i.e., for the two configura- tions to be connected in a transformation series, it is necessary to assume some fairly dramatic reorganiza- tions of the first flagellomere. It may thus be at least as parsimonious to put up a transformation series between the bipartite first flagellomere of Diglossocera bifida Wulp (Crosskey 1976: fig. 51) and the tripar- tite flagellomere of A. antennalis. This is not to say that dramatic reorganizations are impossible; only that tripartition per se is difficult to use as a character (state) and consequently as an argument for phyloge- netic relationship. Whether or not Zrischidocera sauteri and T. atratu- la (Malloch) are distinct species (see note by Crosskey 1976: 210) has no relevance for the present discus- sion, but when Chao & Zhou (1987) described 7. yunnanensis, which has a markedly different head pro- file and a simple, and much shorter, first flagellomere, the genus seems defined mainly by the unique posi- tion of the arista apically on the first flagellomere. The displacement of the arista towards the distal part of the first flagellomere in A. antennalis is in itself evidence of phylogenetic relationship with Trischido- cera, but a similar, although less pronounced, dis- placement occurs in the Oriental genera Ger- mariochaeta Villeneuve (Crosskey 1976: fig. 39) and Lophosiosoma Mesnil, which constitute the tribe Germariochaetini. This tribe is interesting as the low- er calypteres of its members are narrow and diverging, as is shown by Villeneuve (1937: fig. 1) for Germariochaeta clavata Villeneuve, and thus repre- sent a possible synapomorphy with Aporeomyia an- tennalis. No further evidence, however, corroborates a sister-group relationship between Aporeomyia an- tennalis and the morphologically very different Germariochaetini, and the latter taxon is often con- sidered closely related to Triarthria Stephens and its relatives, or to the Minthoini (Crosskey 1976). A slightly swollen subscutellum, the small meta- thoracic spiracle, the narrow, diverging lower calyp- teres, the reduced bend of vein M, and the extension of the first anal vein towards the wing margin are character states shared with the Oriental Malayia. This genus was listed by Crosskey (1976: 163) as a tribally unplaced genus within the Tachininae to- gether with Zrischidocera. At least one of the two spe- cies included in Malayia possesses an aedeagus with separate dorsolateral processes, which have the ex- treme tip free of the aedeagal wall (the only male spec- imen of Malayia known is the single M. fuscinervis Malloch reported by Dear & Crosskey [1982: 134]; examined by TP). The aedeagus of Malayia thus pro- vides a character state that, in combination with the rhinophorid-like structure of metathoracic spiracle, subscutellum, wing venation, and lower calypteres, actually is strong indication that Malayia should be transferred to the Rhinophoridae. Similarities also ex- ist between Aporeomyia antennalis and the Australian- New Guinean taxon containing a dozen undescribed species and tentatively put in a separate (unnamed) tachinoid family by Colless & McAlpine (1991). Apart from possessing all the rhinophorid-like fea- tures mentioned above, these undescribed species have the antennal arista displaced towards apex, sometimes inserted very close to the tip (Colless & McAlpine 1991, fig. 39.38: c, d). Bi- and tripartition occurs in some of the undescribed species but is here of a very different appearance. Also, as in Malayia, the aedeagus has paired dorsolateral processes, which here is considered evidence that these species may possibly not belong to the Tachinidae. In conclusion, we have not been able to find evi- dence in the morphology of Aporeomyia antennalis for a reliable inclusion within any named genus. Tripartition of the first flagellomere gives the species a superficial resemblance to the similarly equipped Trischidocera sauteri, but differences in configuration of the lobes weakens a hypothesis of homology. Other character states suggest a hypothesis of a sister-group relationship to the Germariochaetini, but several of the characters discussed are notoriously much prone to evolutionary homoplasy. ACKNOWLEDGEMENTS Dr. H.-P. Tschorsnig, Stuttgart, Mr. Stig Andersen, Copenhagen, and Dr. Verner Michelsen, Copenhagen, kindly read the manuscript and their suggestions are highly appreciated. We are also grate- ful to Mr. A. Nakanishi, Hyogo Natural History Museum, and Dr. O. Yata, Kyushu University, for making the material available. REFERENCES Chao, C.- M. & S. Zhou, 1987. New species of tachinid flies from Hengduan Mountains of China (Diptera: PAPE & SHIMA : New genus of Tachinidae Tachinidae). — Sinozoologia 5: 207-215. [In Chinese with English summary]. Colless, D. H. & D. K. McAlpine, 1991. Diptera (Flies). — Pp. 717-786 in Naumann, I. E. et al. (eds.), The insects of Australia. Vol. 2. Melbourne University Press, vi + 543-1137. Crosskey, R. W., 1976. A taxonomic conspectus of the Tachinidae (Diptera) of the Oriental Region. — Bulletin of the British Museum (Natural History) Entomology Supplements 26: 1-357. Crosskey, R. W., 1980. Family Tachinidae. — Pp. 822-882 in R. W. Crosskey (ed.), Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History), London. 1437 pp. Dear, J. & R. W. Crosskey, 1982. A taxonomic review of the Tachinidae (Insecta, Diptera) of the Philippines. — Steenstrupia 8: 105-155. Griffiths, G. C. D., 1972. The phylogenetic classification of Diptera Cyclorrhapha with special reference to the struc- ture of the male postabdomen. — Junk Publ., The Hague. 340 pp. McAlpine, J. F., 1989. Phylogeny and classification of the Muscomorpha. — Pp. 1397-1505 in J.F. McAlpine (ed.), Manual of Nearctic Diptera. Vol. 3. Research Branch, Agriculture Canada, Monograph No. 32, vi + 1333- 1581. Pape, T., 1986. A phylogenetic analysis of the woodlouse- flies (Diptera, Rhinophoridae). — Tijdschrift voor Entomologie 129: 15-34. Pape, T., 1992. Phylogeny of the Tachinidae family-group (Diptera: Calyptratae). — Tijdschrift voor Entomologie 135: 43-86. Rognes, K., 1991. Blowflies (Diptera, Calliphoridae) of Fennoscandia and Denmark. — Fauna Entomologica Scandinavica 24: 1-272. Tschorsnig, H.-P., 1985a. Die Struktur des männlichen Postabdomens der Rhinophoridae (Diptera). — Stuttgarter Beiträge für Naturkunde (A) 375: 1-18. Tschorsnig, H.-P., 1985b. Taxonomie forstlich wichtiger Parasiten: Untersuchungen zur Struktur des männlichen Postabdomens der Raupenfliegen (Diptera, Tachinidae.). — Stuttgarter Beiträge für Naturkunde (A) 383: 1-137. Villeneuve, J., 1937. Myodaires supérieurs de Chine. — Bulletin de la Musée Royal d'Histoire Naturelle Belgique 13(34): 1-16. Wood, D. M., 1987. Tachinidae. — Pp. 1193-1269 in J. F. McAlpine (ed.), Manual of Nearctic Diptera. Vol. 2. Research Branch, Agriculture Canada, Monograph No. 28, vi + 675-1332. Received: December 1992 Accepted: 26 January 1993 81 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 BOOK REVIEWS Scoble, Malcolm J., 1992. The Lepidoptera: Form, function and diversity. — Natural History Museum Publications, Oxford University Press. 404 pp., 321 figs., 4 colour plates. [ISBN 0 19 854031 0]. Price £ 45. This book is the third volume in a series, issued by the Natural History Museum of London, the former British Museum (Natural History), providing over- views of the insect orders. Unlike the two previous volumes (on Hymenoptera and on Hemiptera), which are primarily based on the British fauna, “The Lepidoptera’ treats the order on a global scale. More than half the book is devoted to general chapters, dealing extensively with morphology, im- mature stages, hearing sound and scent and the eco- logical importance of Lepidoptera. The remaining part of the book deals with the lepidopteran super- families and families. For all families a short descrip- tion of adult and immature stages is given and biolo- gy, phylogenetic relationships and classification are briefly discussed. For some larger families, the sub- families are treated separately. An extensive list of ref- erences and an index conclude the book. The four colour plates show adults in resting pos- ture, various types of larvae, larval habits and eggs. An adult example of most families is shown in 17 black and white plates, with photos of mounted specimens. Scoble has succeeded in bringing together a wealth of information on the whole order, including much recent work, yet the texts remain concise and clear. Especially the general chapters give much detailed information on lepidopteran structure and function. The family treat- ments differ considerably, undoubtedly because for some families much more data is available than for oth- ers. What is missing compared to the earlier volumes are references to identification guides. However, this is quite understandable on the global scale, because the number of references would have been endless. A slight- ly weak point is the lack of illustration of the diversity of Lepidoptera: the systematic chapters are completely without line drawings and some of the black and white photos of adults are of insuficient quality, hardly giving an impression of the enormous diversity. A few more colourplates would have helped a lot. It is tempting to compare this volume to two other recent treatments of the order, both centered on the Australian fauna: I. F. B. Common’s Moths of Australia (1990) and Nielsen and Common’s chapter on Lepidoptera in the new edition of “The insects of Australia’. Undoubtedly both are far better illustrated than the present volume, especially Common’s book with its many colour plates, and in general the family descriptions have more detail. On the other hand, dealing particularly with the australian fauna, these 82 treatments miss some information on non-australian families, and Scoble’s general chapters have much de- tail not found in the other books. One could wonder if it is wise to produce such similar treatments within this short period, with the inevitable amount of overlap. One final point is purely esthetical: I personally do not like the rather coarse design of the cover (a black zygaenid on red), nor the graphic design (typeset in palatino with a slightly too narrow spacing). In conclusion: a valuable overview of Lepidoptera, especially recommended for general and applied en- tomologists, teachers and students. The specialist can find an easy entry into the literature of a specific fam- ily or subject, in addition to the other recent treat- ments of the group. The general chapters give the most extensive recent review of Lepidoptera form and function. [E. J. van Nieukerken] Fibiger, Michael, 1993. Noctuidae Europaeae. Volume 2, Noctuinae II. — Entomological Press, Soro. 230 pp., 11 colour plates, textfigs, many maps. [ISBN 87-89439-02- 6]. Price DKK 680 excl postage; subscribers to volumes 1-12 receive 10% discount. Distributed by Apollo Books, Kirkeby Sand 19, DK-7771 Stenstrup, Denmark. Fax + 4562263780. This is the second volume of this series, which started in 1990, and concludes the treatment of the subfamily Noctuinae. The series Noctuidae Europaeae will provide in 12 volumes a monograph which will enable an accurate identification of all European species of Noctuidae and supply information on bionomics and distribu- tion. Each volume will treat one or more subfamilies, each with their own authorships. The text is in eng- lish and french, printed alongside. In this volume 116 species are dealt with. For each species a short diagnosis is given, together with notes on bionomics, distribution and taxonomy. An area map gives a general impression of the distribution. The adults are superbly illustrated in the colour plates, which were prepared by the well known pho- tographer David Wilson. Usually both sexes, and some variation is shown. It should be mentioned that the genitalia of the species from the volumes | and 2 will be shown in volume 3. Again, Fibiger has prepared a wonderful piece of work: thorough texts, much new information, which is all brought together in an attractive book. We should admire an amateur with a full time job who produces such a fine piece of scientific work. We must hope that the author and his coworkers find time to continue at least with the same speed: if volumes will come out with the same interval of time as between volume one and two, the whole series will be completed in 33 years! [E. J. van Nieukerken] LOUIS M. ROTH Museum of Comparative Zoology, Harvard University, Cambridge, M. A., U. S. A. REVISION OF THE COCKROACH GENUS CTENONEURA HANITSCH (BLATTARIA, POLYPHAGIDAE). Roth, L. M., 1993. Revision of the cockroach genus Ctenoneura Hanitsch (Blattaria, Polyphagidae). — Tijdschrift voor Entomologie 136: 83-109, figs. 1-21. [iss 0040-7496]. Published 1 July 1993. Most of the 15 known species of Ctenoneura are redescribed and 12 new taxa are described. The genus is principally Malaysian, Indonesian, and Asian. A key is given to distinguish the adults. Dr. L. M. Roth, 81 Brush Hill Road, P. O. Box 540, Sherborn, MA 01770, U. S. A. Key words. — Crenoneura; Blattaria, Polyphagidae; cockroaches; taxonomy; new species. There are 15 described species of Ctenoneura (Princis 1963: 101, 1971: 1138). I have seen the types of ten. I have not examined Bey-Bienko’s spe- cies but have included his references and diagnostic features and included them in my key. Twelve new species are described. The following museums and their curators or col- lection managers kindly loaned me specimens, or house types which I have not examined: EASC - Institute of Entomology, Academy of Science, China. HDEO - Hope Department of Entomology, University Museum, Oxford, England; Dr. George C. McGavin & Mr. I. Lansbury. MCZ - Museum of Comparative Zoology, Harvard University, Cambridge, MA, U.S.A. NRSS - Naturhistoriska Riksmuseet, Stockholm, Sweden; Dr. P. Inge Persson. RMNH - National Museum of Natural History (Rijksmuseum van Natuurlijke Historie), Leiden, The Netherlands; Mr. J. van Tol. zits - Zoological Institute, Lund, Sweden; Dr. Roy Danielsson. ZINR - Institute of Zoology, Academy of Science, Russia. SYSTEMATIC PART Genus Ctenoneura Hanitsch Ctenoneura Hanitsch, 1925: 100; Princis (1953) 1954: 207- 208 (key to species); 1963:101. — Type species: Ctenoneura major Hanitsch, selected by Princis 1950: 204. Etymology. — The name refers to the comb-like ar- rangement of the cubitus branches on the hind wing (Hanitsch 1925: 100). Diagnosis. — Head, pronotum and tegmina not pu- bescent. Tegmina and wings fully developed extend- ing well beyond the end of the abdomen, the former horny, venation clearly defined, discoidal sectors usu- ally oblique (fig. 4C), rarely longitudinal or curved anteriorly (figs. 1F, 2B). Hind wings completely over- lapping each other; usually with (fig. 9c) an interca- lary vein (rarely absent; fig. 1E) between the radial and branched media rami; cubitus vein with three to eight parallel, curved, comb-like branches (some of which may be forked) (figs. 1E, 9C), the anal area folded over the rest of the wing, but not fan-like. Anteroventral margin of front femur with a row of minute piliform spinules, terminating with a single large spine (Type C,); pulvilli absent, tarsal claws symmetrical, usually simple, rarely serrated, arolia, if present, minute, fleshy, unsclerotized. Supraanal plate of both sexes transverse, short. Male: Abdomi- nal terga appear to be unspecialized, but many species have a white, membranous area which could be a ter- gal gland on the supraanal plate (slides of the termi- nalia were not prepared). Subgenital plate usually strongly asymmetrical with modified hind margin (e.g., fig. 6B), rarely symmetrical, unmodified (fig. 1B); usually with only one style on the left side (e.g., fig. 5B), rarely with two (fig. 1B), or styles absent (fig. 14E). Female: subgenital plate valvular. I did not study the male genitalia because for most of the species there were very few duplicate specimens to prepare slides. One of the most diagnostic charac- ters is the shape of the male’s subgenital plate and this structure is so convex that it is greatly distorted when flattened on a slide. Polyphagid genitalia are much more complex structures than those of Blattellidae and Blaberidae, and possibly might not show specific 83 Ty DSCHRIET VOOR ENTOMOLOGIE, VOLUME 136, 1993 differences. I did not think it worth mutilating unique specimens for this purpose. Distribution checklist by species of Ctenoneura aberrans Hanitsch: Mentawai; Sumatra ................ 86 Geiten er Bienko Chun 87 annulicornis Princis: Borneo ................................. 88 GUGLIE en 90 binmanıca Paineisy Burman 91 brunnea Hanitsch: Sumatra ................................. 92 Erassısbylaspan ARAN 107 VJ Anes hs aanva kn 94 geantedsprn Derale ven 87 PESA Primal IS Ua 96 kennen PE AVIRA RE 92 CONA DA AIRIS DANS ALAN EEE 104 luma sp -1).2y eral LI sees Anse PO ee 101 VGA SAW 96 niscaAbeBienko Ch Aree E 9D mjoebergi Princis: Kalimantan; Sarawak … … … … … 97 mnd CISSÉ IS arava kn 106 DATASGULIGAIS PRE AS ab a 103 DOTLTLAIS PAM AE RE RE AE 100 Prop IBOATS D> NES aa PEER 89 seutiensprns Sabah AEN ee 102 CINA) BIEN ACI RR 107 SPES outen A EE O ARE RE en 107 DEA SOA GD, es SUN ATOS ER RO 98 rip CES ars p Nn RS AD AE BR EEE PRE 106 tuberculata Princis: Sumatra … nnn 97 DIODES J Oe See SA DATES TI TI 99 PTE PAD |X EINEA OE {Chi bee ene 94 Geographical distribution checklist of Ctenoneura species Borneo (Kalimantan; Sabah; Sarawak): annulicor- nis, fulva, kinabaluana, major, mjoebergi, murudensis, parascutica; poringa; propannulicornis scutica; tripro- cessa; uncata Burma: birmanica China: acuticerca; misera; simulans, yunnanea Java: kemneri Malay Peninsula (Pahang; Perak): crassistyla, gigan- tea; luma; spinastyla Mentawai I.: aberrans Sumatra: aberrans, biguttata; brunnea; hanitschi; tuberculata South Vietnam: sp. Key to species of Ctenoneura le Hind wing without a intercalary vein (figs. IE, DO) wAroliavabsentsamen sets RP ne 2 = Hind wing with an intercalary vein (fig. 9C). 84 8(6) Arolia minute or absent. Male subgenital PIACE ASY MANCHI CAR E 4 Tarsal claws serrated. Cubitus vein of hind wing with three branches (fig. 1E). (male & female). Male subgenital plate symmetrical with two similar styles (fig. 1B) … … aberrans Tarsal claws simple. Cubitus vein of hind wing with four to six branches (figs. 2C, 2D) Pronotal disk reddish brown, lateral zones darker brown, the edges yellow (fig. 2A). (sex TRIO WIN) re N gigantea Pronotal disk yellowish brown, lateral regions yellow-hyaline. (male) ................... acuticerca Subgenital plate with a setal tuft on the left posterolateral corner (figs. 3A, 3C) ............ 5 Subgenital plate without a setal tuft on the le porter COMENT sesame 6 Hind margin of subgenital plate with two U- shaped excavations on the right side (fig. 3A). Antennae with first four segments brown, succeeded by three whitish antenno- meres, remaining segments brown. (male) ... TICE PET ATES, MIE MES TU te bee? annulicornis Hind margin of subgenital plate unmodified on the right side (fig. 3C). Antennae dark brown, distal end with three white antenno- meres succeeded by four dark terminal seg- ments. (male) ...................... propannulicornis Subgenital plate hind margin deeply excised, with a large, round swelling on the left side Ges, WDA, NSD) scocsssnescecocosoaene 7 Subgenital plate hind margin excised or not, without a round swelling on the left side ..8 Left style small, arising near the apex of the left lobe of the subgenital plate, apexes of both lobes not setose (fig. 12B). (male) … … RNN ee ee leo tuberculata Left style elongate, spine-like, arising near the base of the left lobe of the subgenital plate, apexes of both lobes setose (figs. 13A-D). (Male) e ER ANA spinastyla Pronotum shining black medially, dark red- dish brown laterally, and with a pair of orangish maculae on either side of the ante- rior margin (fig. 4A). Right of subgenital plate with a pair of curved, spine-like pro- cesses directed towards the left (figs. 4B, 4E). (MITE eaters biguttata Pronotum and subgenital plate not as above. pia PE es dini Bora Lu 9 Hind margin of subgenital plate not excavat- ed, with an elongate process on the right side; style, if present, not visible from below OL the rear (Gies EMA) (a MEN 10(9). 11 (10). ANDP 1312). 14 (13). 15 (14). 16 (15). 17 (16). Hind margin of subgenital plate excavated, left style, if present, visibile from below or TE on 10 Left side of subgenital plate with a hook- like structure, style absent (figs. 14C-14E). Antennae with four brown basal segments succeeded by yellowish antennomeres to about the middle, those on the distal half brown with some white terminal or preter- minallones. (MAC)... uncata Subgenital plate without a hook-like struc- ture on the left side, style present. Antennae MOMBASA bover me el. ital Subgenital plate with a long, slender process on the left side, its apex directed towards the right, bearing a small cylindrical style near its middle (figs. 11B-11D). (male) .....mjoebergi Subgenital plate not as above Hind margin of subgenital plate widely exca- vated, with a colourless, membranous area bearing a small, cylindrical style on its basal end (figs. 15A-15C). (male) Hind margin of subgenital plate not as above 13 Hind margin of the subgenital plate divided principally into two or three, dissimilar lobes, with a small cylindrical style arising on the left lobe near its middle, or posterolateral corner, or on the inner margin of the excava- UOMO, th occ ome eRe n. 14 above Hind margin of subgenital plate with a deep V-shaped excavation, with a slender style on the left side; a slender, thorn-like filament is located on the dorsal surface of the plate, protrudes beyond the apical margin of the right lobe (figs. 15E-15G). Supraanal plate inigpnali(fiea15D). (male) 222277... luma Subgenital and supraanal plates not as above ee Lal 15 Subgenital plate with two overlapping lobes on the right side, the dorsal one upturned and partially covering the right side of the su- praanal plate, the ventral lobe with minute spicules near its distal margin (figs. 5B, 5C). (Malo) nnt. birmanica Subgenital plate not as above Left lobe of the subgenital plate much nar- rower than the right one, the style originat- ing on the posterolateral corner (fig. SE). (male) tte ea 2 misera Hind margin of subgenital plate not 25 above i iedee ga 17 Hind margin of subgenital plate with a pair of narrowly separated lobes directed posteri- 20 (19). 23(22). 24 (23). Rorn: Revision of Ctenoneura orly, a style originating near the middle of the left one (figs. 8B-8D). Arolia absent. mal a Se Re fulva Hind margin of subgenital plate not as above ANON RIONI CR RO 18 ‚Style on the left lobe of the subgenital plate originating along the inner margin of the ex- cavation (fig. GB). (male) ............... brunnea Style originating near the posterolateral cor- nemoftheleftlobets. al. nnd... 19 Hind margin of the subgenital plate with a small process that curves dorso-sinistrad (fig. 10B-10D). Pronotum subrectangular, the disk with a sublozenge-shaped reddish brown macula narrowly margined with yel- low, broad lateral zones yellow-hyaline (fig. IVA) (male) ee secs hanitschi Subgenital plate and pronotum not as above Ne En 20 Right lobe of subgenital plate narrow, elon- gate, extending well beyond hind margin of the shorter, wider left lobe (figs. ID, 9E). Pronotum suborbicular, reddish brown (fig. 9A). Cubitus of hind wing with eight branches (4e 20) na major Right and left lobes of subgenital plate not as above (figs. 7C, 7F) : Pronotal edging yellow, disk reddish brown, lateral regions brown-hyaline. Cerci brown. (MAO ee er. ur kemneri Pronotal disk brownish black, lateral regions yellow. Cerci yellow. (male) ......... yunnanea Subgenital plate trigonal, hind margin with three slender processes that are curved dorsad, and a long tapering filament that protrudes between the left and middle processes; style ab- sent (figs. 19A-19E). Cubitus vein of hind wing with three branches (fig. 19F) FO ee ae nk, lee rade à triprocessa Subgenital plate not as above. Style present. Cubitus vein of hind wing with more than three branches. Style present Subgenital plate with a broad plate whose distal margin is excavated and upturned; left style is located in a pale, membranous area (figs. 20A-20E). Cubitus vein of hind wing with eight branches. (male) ........ murudensis Subgenital plate not as above. Cubitus vein of hind wing with less than eight branches . A ee ie eenden 24 Hind margin of subgenital plate deeply exca- vated with a very large, stout style in the left corner (figs. 21A-21C). (male) … crassistyla Subgenital plate and left style not as above … 85 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 25 (24). Subgenital plate broadly excavated, with a short, broad, process without close lobes on either side, that extends dorsad, its distal end curving anteriorly towards the supraanal plate (figs. 18A-18D). (male) ............ kinabaluana — Subgenital plate narrowly divided medially, with a dark spine-like process that curves dorsad between the two lobes ................. 26 26 (25). Margins of the excision smooth, practically touching the spine-like process (figs. 16A- KEEN male) hs. ER eee scutica = Margins of the excision are irregular and on the left side the region is unsclerotized (fig. AB) E (MIO DE parascutica REDESCRIPTIONS, AND DESCRIPTIONS OF NEW i SPECIES Ctenoneura aberrans Hanitsch (fig. 1) Ctenoneura aberrans Hanitsch, 1928: 37, pl. 2 figs. 8,9 (2); 1929a: 294, Bruijning 1948: 41, 148, fig. 54; Princis, 1963: 101. Material examined. — Holotype, (2), Mentawelali, Siberut, no. 91, 23.ix.1924, H. H. Karny; Type Orth. 342'/,, in HDEO. — Paratype. Mentawai. HDEO: same locality and collection data as holotype, 1d (reported as ‘sex 2’), no. 80, 22.ix.1924, Type Orth. 342°/,. — SUMATRA. HDEO: Wai Lima, 16, 12, 2 (abdomens missing). Redescription. — Female: Head covered, interocu- lar space distinctly wider than the distance between antennal sockets. Pronotum with lateral and anterior margins convex, hind margin straight (fig. 1A). Tegmina and wings extending beyond end of abdo- men, the former with a few well marked raised veins, few almost longitudinal veins in discoidal sector (fig. 1F). Hind wings broad, mediastinal vein straight, simple, radial vein slightly sinuous, costals not distin- guishable; media vein, simple, cubitus vein with three curved parallel branches; intercalary vein (between the radial and media veins) absent (fig. 1E). Front fe- mur Type C;; pulvilli and arolia absent, tarsal claws symmetrical, distinctly serrated. Supraanal plate trigonal, apex broadly rounded (fig. 1D). Subgenital plate trigonal, valvular, completely hidden under the supraanal plate. E 1mm F 1mm Fig. 1. Ctenoneura aberrans Hanitsch. — A. female (holotype) pronotum; B, C. male (paratype) subgenital (ventral) and su- praanal (dorsal) plates, respectively; D-F. female holotype: D. supraanal plate (dorsal); E, F. hind wing and tegmen respec- tively (a = radius; b = media; c = cubitus). 86 Fig. 2. Ctenoneura gigantea sp. n., hol- otype. — A. prono- tum; B. right teg- men; C, D. left and right wings. Rorn: Revision of Ctenoneura 2mm Male (previously undescribed): Differs from the fe- male as follows: Supraanal plate transverse, very short, shallowly trigonal, apex rounded (fig. 1C). Subgenital plate essentially symmetrical, with two widely separ- ated, similar styles, interstylar margin unmodified, convexly rounded (fig. 1B). Colour. — Head dark chestnut brown, shiny; an- tennae relatively pale with yellowish tinge. Pronotum reddish brown throughout (fig. 1A). Tegmina hyaline amber, humeral and part of costal vein regions darker (fig. 1F). Wings with a dark blotch in the costal vein region (fig. 1E). Abdomen dark brown. Cerci brown. Legs pale testaceous. Measurements (mm) (2 in parentheses). — Length, 3.8-4.3 (3.6-4.0); pronotum length x width,, 1.1-1.3 x 1.5-1.8 (1.1-1.3 x 1.7-1.8); tegmen length, 4.0-4.5 (4.2-4.6); interocular space, 0.8 (0.8). Comments. — This is the smallest species in the ge- nus and is also unique in having a symmetrical sub- genital plate with two styles, and serrated tarsal claws. Ctenoneura acuticerca Bey-Bienko Ctenoneura acuticerca Bey-Bienko, 1957: 896, 903, 912 (ty- pe 2, Mt. Santaishan, Yunnan, China; in EASC) [not exa- mined] (Russian, English summary); Princis 1963: 101. Description (from Bey-Bienko). — Female: Pronotum smooth. Tegmina with five oblique branches of the radius which run into the median third of the costal margin, anal veins not quite dis- tinct. Radius of hind wings with three slightly pro- nounced oblique branches, intercalary vein absent between radius ‘and media, cubitus divided into six branches. Arolia absent. Supraanal plate large, trian- gular, apex emarginate. Subgenital plate with valvular part lozenge shaped. Apex of cerci with a large spine. Colour. — Brownish yellow, pronotal disk yellow- ish brown, lateral parts yellow, transparant. Wings with distal part yellowish. Legs yellow. Cerci brown- ish yellow. Male: Unknown. Measurements (mm). — Body length, 8.5-8.7; pro- notum length, 2.7; tegmen length, 10.2-10.5. Comments. — According to Bey-Bienko, this spe- cies is related to aberrans (probably because of the ab- sence of an intercalary vein, and arolia). Ctenoneura gigantea Sp. n. (fig. 2) Type material. — Holotype, sex unknown (abdomen miss- ing), Gunung Kledang, Perak, 2646’, xi.1916 (with a handwritten label, C. pauciramosa..., ms. name); in RMNH. Description. — Sex unknown: Head exposed, inter- ocular space greater than the distance between ocellar spots and antennal sockets. Pronotum parabolic (fig. 2A). Tegmina and wings fully developed, the former with raised veins, and longitudinal discoidal sectors (fig. 2B). Hind wing with about three poorly defined costal veins, intercalary vein absent, media vein with one branch, cubitus vein with four branches (one forked near the base on the left wing), and six branch- es (two of them joined distad on the right wing) (figs. 87 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 3. Ctenoneura spp. — A-B, C. annulicornis Princis, male holotype, A. subgenital plate (ventral); B. subgenital plate (rear view); C-E. C. propannulicornis sp. n., male holotype: C. subgenital plate (ventral); D. supraanal and subgenital plates (rear view); E. supraanal and subgenital plates (dorsal; the distal region of the supraanal plate is deflexed and the margin is distort- ed) 2C, D). Front femur Type C;; pulvilli absent, tarsal claws very long, simple symmetrical, arolia absent. Abdomen missing. Colour. Head reddish brown, the region from the occiput to ocellar spots darker; antennae blackish. Pronotal disk reddish brown, lateral zones darker brown, lateral edges yellowish (fig. 2A). Measurements (mm). — Length, ?; pronotum length x width, 2.9 x 3.7; tegmen length, 10.4; inter- ocular space, 1.3. Etymology. — The specific name refers to the large size of the species. Comments. — I am describing this species even though its abdomen is missing, because its size is so much greater than any of the known species. Also the parabolic shape of the pronotum, the absence of an intercalary vein on the hind wing (known only in aberrans and acuticerca), and the complete absence of arolia, are good distinctive characters when coupled with its size. 88 Ctenoneura annulicornis Princis (figs. 3A-B) Ctenoneura annulicornis Princis, (1953) 1954; 206, fig. 3. Holotype, d, Borneo, Long Navang, Mjöberg (NRSS) [examined]; Princis 1963: 102. Ctenoneura fulva (nec Hanitsch, 1925); Hanitsch 1933: 235 [in part; only the Long Navang d |; Princis 1963:102. Redescription. — Male: Head hidden, interocular space greater than the distance between ocellar spots and antennal sockets. Pronotum suboval. Tegmina and wings extending well beyond end of abdomen. Tegminal veins not raised. Hind wing with interca- lary vein, media vein with one forked branch, cubitus with six branches. Front femur Type C;; pulvilli and arolia absent, tarsal claws symmetrical, simple. - Abdominal terga unspecialized. Supraanal plate transverse, narrow, symmetrical. Subgenital plate asymmetrical with a setal tuft on the left posterolater- al corner, and with three sclerotized structures and a Fig. 4. Ctenoneura biguttata Hanitsch, male holotype. — A. pronotum; B. su- praanal and subgeni- tal plates (rear view); C, D. left tegmen and hind wing; E. supraanal and sub- genital plates (dor- sal). Rove: Revision of Ctenoneura well developed cylindrical style, also on the left side, the right side with a pair of U-shaped excavations (figs. 3A, B). Colour. — Head dark reddish brown; antennae with first four segments brown, succeeded by three whitish antennomeres, remaining segments brown. Pronotal disk dark reddish brown, lateral zones hya- line. Tegmina hyaline light brown. Adomen brown, broad lateral zones on the sterna pale. Legs light brown. Female: Unknown. Measurements (mm). — Length, 5.7; pronotum length x width, 1.7 x 2.2; tegmen length, 6.3; interoc- ular space, 0.6. Ctenoneura propannulicornis sp. n. (figs. 3C-E) Type material. — Holotype, d, Malaysia, Sabah, Ulu Rurun, 115°40’30”E 4°22’N, 1500 m, 21-22.xii.1986, J. Huisman (RMNH). — Paratype. Sabah, same locality and col- lector as the holotype, ML, 1d, 20.xii.1986 (RMNH). Description. — Male: Head hidden, interocular space greater than the distance between the antennal sockets. Pronotum subelliptical. Tegmina and wings 05 mm fully developed extending well beyond the end of the abdomen, discoidal sectors of the former oblique. Hind wing with distinct costal veins, the proximal ones thickened distad, intercalary vein present, media vein with one branch, cubitus vein with five branches (one of them may be forked). Front femur Type C;; pulvilli absent, tarsal claws simple, symmetrical, aro- lia minute. Abdominal terga unspecialized. Supraanal plate transverse, hind margin broadly, shallowly exca- vated, with a large, dense group of curved setae on the left corner, style absent, right side of plate unmodi- fied (figs. 3C-E). Colour. — Head dark brown; antennae dark brown, the distal end with three white antennomeres, suc- ceeded by four dark terminal segments. Pronotal disk dark brown, the lateral regions infuscated-hyaline. Tegmina dark brown-hyaline, humeral region yel- lowish. Hind wing infuscated, darkest in the costal vein and apical regions, lightest in the mid portion. Abdominal terga brown, lateral edges darker, supraa- nal plate with a white macula anteromedially. Abdominal sterna light brown, subgenital plate dark- er. Cerci brown, apical segment lighter. Coxae, femo- ra, and tibiae brown, tarsi lighter. Female: Unknown. Measurements (mm). — Length, 5.8-6.3; prono- 89 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 5. Ctenoneura spp. — A-D, C. birmanica Princis, males: A. pronotum (holotype); B-D. paratype: B. subgenital plate (ventral); C, supraanal and subgenital plates (dorsal; the dotted line indicates the margin of the supraanal plate that is hid- den by one of the lobes of the subgenital plate); D. hind wing. — E, C. misera Bey-Bienko, male holotype, subgenital plate (ventral) (redrawn from Bey-Bienko, 1970). tum length x width, 1.9 x 2.3-2.4; tegmen length, 7.1-8.3; interocular space, 0.6. Comments. — I place this species near annulicornis (hence the specific name), because both taxa have a dense tuft of setae in the left corner of the subgenital plate. Differences in the shape of the right side of the plate, absence of the left style (in propannulicornis), and antennal colouration, separate the two species. Ctenoneura biguttata Hanitsch (fig. 4) Ctenoneura biguttata Hanitsch, 1932a: 6. Holotype, d, Sumatra, Deli, Fulmek, 1921-26, Brastagi, Urwald, vili.1924, R. Ebner; Type Orth. 375 (HDEO), [exami- ned]; Bruijning 1948: 41, 149; Princis (1953) 1954: 207 (footnote), 1963: 103. Redescription. — Male: Head exposed, eyes wide 90 apart, interocular space greater than distance between antennal sockets. Pronotum subcircular (fig. 4A). Tegmina and wings extending beyond end of abdo- men, veins and cross veins of both distinct, discoidal sectors of the former oblique (fig. 4C). Wing with six costals, the first four with distal ends merging into a marginal dark patch; intercalaray vein present between the radial and median; media vein branched, cubitus with five branches (fig. 4D). Front femur Type C,; pulvilli and arolia absent, tarsal claws sym- metrical, simple (not serrated). Abdominal terga un- specialized. Supraanal plate transverse, very narrow, hind margin entire (fig. 4E). Subgenital plate asym- metrical, with a small cylindrical style on the left side, (Hanitsch stated, ‘styles not observed’), right side with a pair of curved spinelike processes directed to- wards the left (figs. 4B, E). Colour. — Head shining black; antennae pale yel- lowish with darker terminal segments. Pronotum shiny black medially, dark reddish brown laterally, and with a pair of orangish maculae on either side of the anterior margin (fig. 4A). Tegmina uniformly dark brown (fig. 4C). Wings hyaline, brown, with a dark patch along the anterior costal region (fig. 4D). Abdominal terga light brown, sterna dark brown, subgenital plate with an orangish area distad. Legs with coxae and femora dark brown, tibiae and tarsi yellowish. Cerci with brown basal segments, remain- ing cercomeres yellowish. Female: Unknown. Measurements (mm). — Length, 5.7; pronotum length x width, 1.7 x 2.2; tegmen length, 7.3; interoc- ular space, 0.8. Ctenoneura birmanica Princis (figs. SA-D) Ctenoneura birmanica Princis, (1953) 1954: 206, fig. 5. Holotype, d, Kambaiti, N. E. Burma, 7000 ft, Rovu: Revision of Ctenoneura 25.v.1934, R. Malaise (NRSS) [examined]; Princis 1963: 102. Ctenoneura major (nec Hanitsch, 1925); Princis 1950: 203, 204 (8), 1963: 102. Material examined. — Holotype, and paratypes. Burma. NRSS: 1 d, same locality and collector as holotype, 2000 m, 28.v.1934. ZIS: 1d, same data as holotype; 1 d , same local- ity and collector as holotype. 2000 m, 29.v.1934. Redescription. — Male: Head hidden or slightly ex- posed; interocular space greater than distance between ocellar spots and antennal sockets. Pronotum subelliptical (fig. 5A). Tegmina and wings fully developed extending well beyond end of abdo- men, discoidal sectors of former oblique. Hind wing with the distal ends of the proximal costal veins reach- ing only to the dense spot near the costal margin; intercalary vein present, media vein with two branch- es (one forked), cubitus vein with six branches (one forked) (fig. 5D). Front femur Type C;; pulvilli ab- Fig. 6. Crenoneura brunnea Hanitsch, male holotype. — A. pronotum; B. subgenital plate (ventral); C, D. left tegmen and hind wing; E. supraanal and subgenital plates (dorsal). 91 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 sent, tarsal claws symmetrical, simple, arolia minute. Supraanal plate transverse, hind margin convexly rounded, symmetrical, entire (fig. 5C). Subgenital plate asymmetrical, divided by a medial excavation, one lobe on the left side bearing a cylindrical style, two overlapping lobes on the right side, the dorsal one upturned (in the holotype only it partly covers the right side of the supraanal plate), the ventral lobe with minute spicules near its distal margin (figs. 5B, ©) Colour. — Antennae dark brown with pale junc- tions between individual segments. Head reddish brown. Pronotal disk reddish brown, lateral areas dark-hyaline (fig. 5A). Tegmina brownish, hyaline. Hind wings weakly infuscated. Abdomen dark brown, supraanal plate black. Legs brown. Cerci brown, proximal cercomeres darker on both surfaces. Female: Unknown. Measurements (mm). — Body length, 7.7-8.1; pro- notum length x width, 2.1-2.2 x 3.0; tegmen length, 10.0-10.4; interocular space, 0.8-0.9. Ctenoneura misera Bey-Bienko (fig. SE) Ctenoneura misera Bey-Bienko, 1969: 832, fig. 1 (Russian). Holotype d, S. China, Yunnan, Tapa-shan Mountains, near Pin P’ien, 1300 m, 23.vi.1956, Huen K’e-yen; in ZINR. [not examined]; 1970: 528 (English translation). Description (after Bey-Bienko). — Male: Small. Head with flat vertex, almost pendent (overhanging), forming an obtuse angle with the frons; interocular space slightly more than double the distance between antennal sockets. Pronotum one and one half times as wide as long, with broad transparent lateral parts, hind margin with a narrower hyaline border. Hind wings with oblique branches of the radius distinct, vein between radius and media arched backwards; cubitus vein with seven branches. Subgenital plate as in fig. 5E [style in the left posterior corner]. Cerci long, thickened towards the bases, terminal segment not spine-like. Colour. — Rusty yellow. Head chestnut brown, an- tennae brownish yellow. Hind wing just visibly infus- cate, yellowish along costal margin. Female: Unknown. Measurements (mm). — Body length, 6.8; prono- tum length x width, 2.0 x 3.0; tegmen length, 8.0; overall length, 10.0. Comments. — According to Bey-Bienko, this spe- cies is similar to yunnanea from China and kemneri from Java, but clearly distinguished by the colour of head and pronotum, small body, and structural de- tails of the subgenital plate. The subgenital plate (fig. SE) of misera more closely resembles that of birmani- 92 ca (fig. 5B) than it does yunnanea (fig. 7F) or kemne- ri (fig. 7C). Ctenoneura brunnea Hanitsch (fig. 6) Ctenoneura brunnea Hanitsch, 1929a: 266, 292, fig. 5; 1932b: 52, 81. Holotype, d [Hanitsch did not indicate the sex], Gunung Singgalong (Sumatra, Westkust), 1800 m, 1925, E. Jacobson; Type Orth. 362, in HDEO [exam- ined]; Bruijning 1948: 41, 148: Princis (1953) 1954: 208, fig. 8 (d) Princis 1963: 102. Additional material. — Sumatra. RMNH: N. Sumatra, Bivouac One, Mt. Bandahara, 3°43’N 97°41’E, ca. 810 m, 1 6, 25.vi. —5.vii.1972, J. Krikken. Redescription. — Male (Hanitsch did not deter- mine the sex of the type): Head covered, interocular space less than distance between antennal sockets. Pronotum subcircular (fig. GA). Tegmina and wings fully developed extending beyond end of abdomen, veins of the former distinct, not raised, discoidal sec- tors oblique (fig. 6C). Hind wing with a forked media vein, intercalary vein present, cubitus with six branches (fig. 6D). Front femur Type C,; pulvilli ab- sent, tarsal claws symmetrical, simple, arolia subobso- lete. Abdominal terga unspecialized. Supraanal plate strongly transverse, shallowly trigonal (fig. GE). Subgenital plate asymmetrical, hind margin deeply, widely excavated, with a small cylindrical style on the left side along the inner margin of the excavation (fig. 6B). Colour. — Head shiny dark reddish brown; anten- nae dark brownish black. Pronotal disk dark reddish brown, lateral borders broad, hyaline, partially yel- lowish (fig. 6A). Tegmina shining brown. Wings brownish, darker along the distal half of the costal margin and apically (fig. 6D). Abdomen yellowish to reddish brown, lateral borders darker. Cerci dark brown to black. Coxae, femora, and tibiae reddish brown, tarsi yellowish. Female: Unknown. Measurements (mm). — Length, 5.6-6.5; prono- tum length x width, 1.6-1.8 x 2.1-2.3; tegmen length, 7.1-8.5.; interocular space, 0.5-0.7. Ctenoneura kemneri Princis (figs. 7A-E) Ctenoneura kemneri Princis, 1967: 146, fig. 1. Holotype d, Tjitjoroeng, Java, 23.1.1921, N. A. Kemner (ZILS) [exam- ined] ; Princis 1971: 1138. Redescription. — Male: Head hidden, interocular space wrinkled, slightly less than the distance between the antennal sockets. Pronotum strongly transverse, Fig. 7. Ctenoneura spp. A-E. — C. kemneri Princis, male holotype: A. pronotum; B. supraanal and subgenital plates (dorsal); C. subgenital plate (ventral); D. subgenital plate (rear view); E. subgeni- tal plate (left lateral). — F. C. yunnanea Bey-Bienko, male holotype, subgenital plate (ventral) (redrawn from Bey- Bienko, 1957). its greatest width before the middle, anterior margin cucullate above the head (fig. 7A). Tegmina extend- ing well beyond end of abdomen, discoidal sectors of the former oblique. Hind wing with distinct costal veins, not thickened distad; intercalary vein present, media vein with one branch, cubitus vein with six or seven branches (one of the branches may be forked). Front femur Type C,, terminal spine very small; pul- villi absent, tarsal claws symmetrical, simple, arolia minute. (Princis claimed that only the fourth segment of the basitarsus had a pulvillus. However this ‘pulvil- lus’ is spined and is no different from the pulvillar re- gion on the other tarsomeres. Pulvilli normally are fleshy and lack spines and I consider the basitarsus in this species as lacking a pulvillus). Abdominal terga unspecialized; supraanal plate short, transverse, hind margin convexly rounded (fig. 7B). Subgenital plate divided into two dissimilar lobes, the left one bearing a cyclindrical style in the posterolateral corner (figs. 7B-D). Colour. — Head dark brown with lighter mouth- parts; antennae brown. Pronotum with yellow edging (wider laterally than on the anterior and posterior Rorn: Revision of Ctenoneura margins, disk bright red brown, the lateral parts hya- line brown. Tegmina uniformly reddish brown, veins yellowish. Hind wings darkly infuscated, part of co- stal vein area with a yellowish patch. Abdomen brown. Cerci brown, darker ventrally. Coxae, femora, and tibiae reddish brown, tarsi yellowish. Female: Unknown. _ Measuremnts (mm). Body length, 6.5; pronotum length x width, 2.0 x 3.2; tegmen length, 8.2; interoc- ular space, 0.6. Comments. — Princis stated that kemneri to some degree resembled major, chiefly because its subgenital plate is of the same type. However their lobes differ in shape. Actually Princis’s drawing of the subgenital plate of kemneri (Princis, 1967: fig. 1) resembles that of yunnanea (fig. 7F), more closely than it does that of major (fig. IE). My camera lucida drawing of the sub- genital plate of kemneri differs from Princis’s but slight differences in the angle from which the speci- men is drawn can make a marked difference in the shape of the structure. The shapes of the subgenital plate lobes of kemneri (fig. 7C) show some resem- blance to those of birmanica (fig. 5B). 93 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 1mm 0.5 mm 0.5 mm Fig. 8. Ctenoneura fulva Hanitsch, male holotype. — A. pronotum; B. subgenital plate (ventral); C. supraanal and subgenital plates (rear view); D. supraanal and subgenital plates (dorsal); E, F. left tegmen and hind wing. Ctenoneura yunnanea Bey-Bienko (fig. 7F) Ctenoneura yunnanea Bey-Bienko, 1957: 896, 902, 912, fig. 1. Holotype d, Mt. Santaishan, Yunnan, China (EASC) [not examined] (Russian, English summary); Princis 1963:102. Description (from Bey-Bienko). — Male: Prono- tum transverse. Tegmina and wings with typical ve- nation; wings with distinct oblique branches of the radius, intercalary vein between radius and media present, cubitus vein with seven branches. Supraanal plate very short. Subgenital plate with a deep round emargination, right margin thickened, style long, thin, placed on the upper left margin of the plate (fig. 7F). Cerci moniliform. Colour. — Brownish yellow. Vertex distinctly bor- dered above antennal sockets. Pronotal disk brownish black, lateral parts yellow. Cerci yellow. 94 Female: Unknown. Measurements (mm). — Body length, 7.0; prono- tum length, 2.0; tegmen length, 8.5. Comments. — Bey-Bienko stated that this species strongly resembles hanitschi. However, its subgenital plate is similar to that of brunnea and kemneri. Ctenoneura fulva Hanitsch (fig. 8) Ctenoneura fulva Hanitsch, 1925: 78, 101, figs. 13-14 (d & 2). Lectotype (here designated) d , Mt. Murud, Sarawak, 6000 ft., Dr. E. Mjöberg, Sarawak Mus. dd 1925; Type Orth. 300'/,, in HDEO [examined]. [Hanitsch listed six specimens in his original description, three d d from Mt. Murud, and one 6 and two 2 ® from Mt. Dulit; there are only two paralectotypes in the HDEO, and neither one is fulva; see comments, below]; Princis 1933: 303, 329; Bruijning 1948: 41, 148; Princis (1953) 1954: 207, fig. 6 (8); Princis 1963: 101. Redescription. — Male: Head covered by prono- tum, interocular space greater than distance between antennal sockets. Pronotum subelliptical (fig. 8A). Tegmina and wings extending well beyond end of body, the former with proximal costal veins arising from a common trunk, the distal ones singly, discoid- al sectors oblique (fig. 8E). Hind wing with interca- lary vein, media vein with two branches, cubitus with six simple branches (fig. 8F). Front femur Type C; pulvilli and arolia absent, tarsal claws symmetrical, simple. Abdominal terga unspecialized. Supraanal plate short transverse, hind margin shallowly convex (figs. 8C, D). Subgenital plate asymmetrical, hind margin with a pair of weakly dissimilar lobes, and a small cylindrical style on the left side adpressed to the surface, its apex directed anteriorly. The style is hid- den in ventral view but can be seen in end view. The tips of two apically acute structures, probably genital phallomeres, protrude laterally, and dorsally from be- neath the supraanal plate (figs. B-D). Colour. — Head dark reddish brown, shiny, anten- Rotu: Revision of Ctenoneura nae dark brown. Pronotal disk reddish brown, sur- rounding area broad, hyaline (fig. 8A). Tegmina brownish yellow, humeral and costal area lighter (fig. 8E). Wings pale hyaline with a yellowish tinge, ante- rior portion with a dark area along the anterior mar- gin where the ends of the proximal costals merge (fig. 8F). Abdominal terga brown, terminal segments darker, supraanal plate with a pale medial zone. Abdominal sterna light brown, margins darker. Legs brown. Female: Not seen. Measurements (mm). — Length, 6.6; pronotum length x width, 1.9 x 2.7; tegmen length, 9.5; interoc- ular space, 0.7. Comments. — Two of Hanitsch’s paralectotypes are not fulva. One from Mt. Murud (Type Orth. 300'/,) is a new species, murudensis, described below. The other paralectotype, from Mt. Dulit (Type Orth. 300°/,), is possibly new but I am not describing it be- cause its subgenital plate is distorted and it is difficult to see the shape of the hind margin. Fig. 9. Ctenoneura major Hanitsch, male holotype. A. pronotum; B, C. left tegmen and hind wing; a = intercalary vein. D. supraanal and subgenital plates (dorsal); E. subgenital plate (ventral); F. subgenital plate (right lateral). 95 Ty DSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Ctenoneura major Hanitsch (fig. 9) Ctenoneura major Hanitsch, 1925: 78, 102 (d & 9). Holotype d, Mt. Murud, Sarawak, 6500 ft., x.1922- i.1923, Dr. E. Mjôberg, Sarawak Mus., dd 1925; Type Orth. 301, in HDEO [Hanitsch listed a d and 2 from Mt. Murud, but according to the number 301, there is only one type specimen in HDEO] [examined]; Hanitsch 1927: 9, 26, 42 (in part?); Bruijning 1948: 41, 148; Princis (1953) 1954: 207, fig. 7 (8); Princis 1963: 101. Redescription (Hanitsch described both d & 9 in the same paragraph. His description can not be used to distinguish between the sexes). — Male: Head slightly exposed, interocular space less than the dis- tance between ocellar spots and antennal sockets. Pronotum suborbicular (fig. 9A). Tegmina and wings extending beyond end of abdomen, the former with distinct, but not thickened veins, discoidal sectors oblique (fig. 9B). Hind wing with eight costals, the distal ends of the first three merging into a thicken- ing, intercalary vein present, media with two branch- es, cubitus vein with eight rami (fig. 9C). Front femur Type C;; pulvilli absent, tarsal claws simple, symmet- rical, arolia minute. Abdominal terga unspecialized. Supraanal plate transverse, very narrow, hind margin convex, entire (fig. 9D). Subgenital plate asymmetri- cal, a small cylindrical style on the left side near the posterior corner, right side produced, the process di- rected posteriorly (figs. 9D-F). Colour. — Head reddish and dark brown, shining, ocelli pale, antennae dark brown. Pronotal disk dark reddish brown (fig. 9A). Tegmina brown (fig. 9B). Wings brown, thickened costal vein region darker (fig. IC). Abdomen brownish yellow, hind margin of subgenital plate in part yellowish. Cerci fuscous brown. Legs dull brownish yellow. Female. Not seen. Measurements (mm). — Length, 9.7; pronotum length x width, 2.2 x 3.0; tegmen length, 11.1; inter- ocular space, 0.7. Ctenoneura hanitschi Princis (fig. 10) Ctenoneura hanitschi Princis, (1953) 1954: 204, fig. 1 (6d. Holotype, d, Sibajak, Sumatra, 1600 m, Mjöberg; (NRSS) [examined]; 1963: 102. Ctenoneura major (nec Hanitsch, 1925); Hanitsch 1929b: 3, 18 (in part, d). Redescription. — Male: Head hidden, interocular space greater than the distance between antennal sockets. Pronotum subrectangular (fig. 10A). Tegmina and wings fully developed extending well beyond end of abdomen, the former with oblique dis- coidal veins. Hind wing with an intercalary vein. Front femur Type C,; pulvilli absent, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized. Supraanal plate transverse, very short, hind margin shallowly convex, entire (fig. 10C). Subgenital plate asymmetrical, with a cylindrical style on the left side; the right side has a process that is Fig. 10. Ctenoneura hanitschi Princis, male holotype. — A. pronotum; B. subgenital plate (ventral); C. supraanal and subgen- ital plates (dorsal); D. supraanal and subgenital plates (rear view). 96 A | ne RotH: Revision of Ctenoneura TOR Ne Fig. 11. Crenoneura mjoebergi Princis, male holotype. — A. pronotum; B. supraanal and subgenital plates (rear view); C. su- praanal and subgenital plates (dorsal); D. subgenital plate (ventral). curved dorsad towards the left (figs. B-D). Colour. — Head dark reddish brown; antennae dark brown, monochromatic. Pronotal disk with a sublozenge-shaped reddish brown macula narrowly margined with yellow, broad lateral zones yellowish- hyaline (fig. 10A). Tegmina brownish. Hind wing weakly infuscated. Abdomen brown. Legs dark to yel- lowish brown. Female: Unknown. Measurements (mm). — Body length, 7.3; prono- tum length x width, 2.2 x 2.5; tegmen length, 9.4; interocular space, 0.6. Ctenoneura mjoebergi Princis (fig. 11) Ctenoneura mjoebergi Princis, (1953) 1954: 206, fig. 4 (d). Holotype, d, O. Borneo, Pajau River, [Kalimantan], Mjöberg; in (NRSS) [examined]. Ctenoneura fulva (nec Hanitsch, 1925). — Hanitsch 1933: 235 (in part; Mt. Tibang and Pajau River material); Princis 1963: 102. Material examined. — Holotype, and paratype: Kalimantan. 1d, same data as holotype (NRSS). Sarawak. Mt. Tibang, 1400 m, 1 d, Mjöberg (NRSS). Redescription. — Male: Head concealed, interocu- lar space greater than distance between antennal sock- ets. Pronotum subelliptical (fig. 11A). Tegmina and wings fully developed extending well beyond end of abdomen, the former with oblique discoidal sectors. Hind wing with an intercalary vein, media vein with a forked branch, cubitus with five branches (one forked). Front femur Type C,; pulvilli absent, tarsal claws simple, symmetrical, arolia minute. Supraanal plate symmetrical, transverse, shallowly trigonal, hind margin entire (fig. 11C). Subgenital plate asymmetri- cal with several processes, one of which on the left side is elongated and bears a single style (figs. 11B-D). Colour. Head reddish brown; antennae with seg- ments on proximal half yellow (except for the first two brown antennomeres), distal half brownish. Pronotal disk reddish brown with a pair of dark spots on anterior margin, lateral zones hyaline (fig. 114A). Tegmina brownish, subcostal vein yellow. Hind wing lightly infuscated. Abdomen brown, the sterna light- er medially and laterally. Legs and cerci brown. Female: Unknown. Measurements (mm). — Length, 5.5-5.8; prono- tum length x width, 1.5-1.7 x 2.1-2.3; tegmen length, 6.6-6.8; interocular space, 0.6. Ctenoneura tuberculata Princis (fig. 12) Ctenoneura tuberculata Princis, (1953) 1954: 205, fig. 2 (3). Holotype, d, Sumatra, Sibajak, Mjoeberg (NRSS) [examined]; Princis 1963: 101. 97 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 C _ 05mm B 0.5 mm Fig. 12. Ctenoneura tuberculata Princis, male from Mt. Bandahara. A. subgenital plate (left lateral); B. subgenital plate (ven- tral); C. subgenital plate (right lateral). Ctenoneura major (nec Hanitsch, 1925); Hanitsch 1929b: 3, 18 (à, in part); Princis 1963: 101. Additional material. — Sumatra. N. Sumatra, Bivouac One, Mt. Bandahara, 3°43’N 97°41’E, ca. 810 m, 1d, 25.vi. —5.vii.1972, J. Krikken (RMNH). Redescription. — Male: Head hidden, interocular space greater than the distance between antennal sockets. Pronotum subelliptical. Tegmina and wings fully developed extending beyond end of abdomen, the former with oblique discoidal sectors. Hind wing with intercalary vein, media vein with one branch, cubitus vein with six branches. Front femur Type C; pulvilli absent, tarsal claws symmetrical, simple, aro- lia minute. Abdominal terga unspecialized. Supraanal plate shallowly trigonal, apex rounded and medially incised. Subgenital plate strongly asymmetrical, hind margin deeply divided forming two asymmetrical lobes, the one on the left bearing a small cylindrical style below its apex on the inner margin of the exci- sion; the lobe on the right deeply excised; a large round swelling is located along the margin distad on the left side (figs. 12A-C). Colour. — Head reddish brown; antennae dark brown. Pronotal disk reddish brown the surrounding area hyaline and partly yellowish. Tegmina with hu- meral and costal vein areas yellowish, remainder brown. Hind wing with yellowish tinge. Abdomen brown, subgenital plate reddish brown (holotype), or partly yellowish. Legs light brown. Cerci light brown dorsally, darker brown ventrally. Female: Unknown. | Measurements (mm). — Body length, 6.0; prono- tum length x width, 1.7-1.8 x 2.3-2.5; tegmen length, 7.2-9.0; interocular space, 0.5-0.6. 98 Ctenoneura spinastyla sp. n. (fig. 13) Type material. — Holotype, d, Pahang, F.M.S., ‘Cameron’s Highlands’, No. 4 Camp, 4800 ft, 14.- viii.1923, H.M., Pendlebury (incorrectly labelled by Hanitsch, Ctenoneura major Hanitsch); in HDEO. Description. — Male: Head hidden, interocular space greater than the distance between ocellar spots and antennal sockets. Pronotum suboval. Tegmina and wings extending well beyond end of abdomen, discoidal sectors of the former oblique. Hind wing with distinct costal veins, intercalary vein present, media vein forked beyond the middle, each branch again dividing (left wing, fig. 13E), or with two branches (one of them forked, right wing); cubitus vein with five or six branches (two of them branching once or twice) (fig. 13E). Front femur Type C,; pul- villi absent, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized. Supraanal plate strongly transverse, very short, hind margin al- most truncate (fig. 13A). Subgenital plate asymmetri- cal with a pair of large dissimilar processes, the left one with a bulbous swelling near the apex, both lobes setose apically, those on the right much longer; on the left margin near the base of the plate is a long, slender, spine-like style (figs. 13A-D). Colour. — Head reddish brown; antennal segments black (most of the antennae are missing). Pronotal disk reddish brown, becoming yellowish in the poste- rior region, lateral zones yellowish-hyaline. Tegmina with humeral zone yellowish, remainder brownish- hyaline, mediastine vein very dark. Abdomen, cerci, and legs brown. Female: Unknown. Measurements (mm). — Length, 8.2; pronotum Fig. 13. Ctenoneura spinastyla sp. n., male holotype. — A. su- praanal and subgenital plates (dorsal); B. subgenital plate (right lateral); C. subgenital plate (ventral); D. subgenital plate (left lateral); E. left hind wing. length x width, 2.3 x 2.9; tegmen length, 11.1; inter- ocular space, 0.8. Etymology. — The specific name refers to the slen- der, greatly elongated left style. Comments. — The two setose processes, and the spine-like style on the subgenital plate are unique and readily identifies this species. Ctenoneura uncata sp. n. (fig. 14) Type material. — Holotype, 4, Malaysia, S.W. Sabah, nr. Long Pa Sia (East), c. 1000 m, Mal. trap 5, 1-13.iv.1987, C. v. Achterberg; in RMNH . — Paratypes: Sabah. RMNH: The following were collected by C. v. Achterberg: S.W. Sabah, nr. Long Pa Sia (West), c. 1050 m, Mal. trap 3, 1 d, 1- 14.iv.1987; same data as holotype, 1 d, 25.xi. —7.xii.1987; nr. Long Pa Sia (West), 1010 m, Mal. trap 1b, 1 d, Mal. trap 2, 1020 m, 2 dd, 25.xi. -8.xii.1987. The following were collected by J. Huisman: Sg. Malabit, Rd. Long Sia-L Semado, 115°41’E 4°21’N, 1175 m, 1 d, 8.xii.1986; Long Pa Sia, confl. S. Maga and S. Pa Sia, 4°26’30”N 115°40’E, high moist forest, sandsoil, 1350 m, 2 d d, 16-19.x.1986. One d retained in MCZ. Description. — Male: Head hidden, interocular space greater than the distance between the ocellar spots and antennal sockets. Pronotum suboval (fig. 14A). Tegmina and wings fully developed extending RorH: Revision of Ctenoneura beyond end of abdomen. Hind wing with intercalary vein, media vein with one branch, cubitus vein with five or six branches (some may be forked). Front fe- mur Type C,; pulvilli absent, tarsal claws symmetri- cal, simple, arolia minute. Abdominal terga unspe- cialized. Supraanal plate transverse, subtrigonal (fig. 14E). Subgenital plate strongly asymmetrical, the right side with a broad, elongated lobe that curves to- wards the left where there is a broad, apically curved, hook-like process; style absent (figs. 14B-E). Colour. — Head dark reddish brown; basal four an- tennal segments brown, succeeded by pale (yellowish) antennomeres to about the middle, those on the distal half brown, with as many as six terminal (or pretermi- nal) white antennomeres (some lacked, or had only two or three white segments, but these may have been damaged specimens). Pronotal disk with a broad, dark reddish brown macula that extends to both the anteri- or and posterior margins, lateral areas usually hyaline (fig. 14A), sometimes partly opaque yellowish. Tegmina with humeral and proximal half of the costal vein region yellowish-hyaline, the remainder dark hya- line-brown. Hind wing with yellowish tinge in the co- stal vein region. Abdomen brown, supraanal plate with a white area, subgenital plate may be a lighter brown and sometimes partly yellow. Female: Unknown. 99 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 14. Ctenoneura uncata sp. n., male paratype from near Long Pa Sia. — A. pronotum; B. subgenital plate (ventral); C. sub- genital plate (left lateral); D. subgenital plate (rear view); E. supraanal and subgenital plates (dorsal). Measurements (mm). — Length, 5.0-5.8; prono- tum length x width, 1.6-1.7 x 2.1-2.3; tegmen length, 7.0-7.7; interocular space, 0.6. Etymology. — The specific name refers to the hook- shaped structure on the left side of the subgenital plate. Ctenoneura poringa Sp. n. (figs. 15A-C) Type material. — Holotype, d, Malaysia, Sabah, Poring hot spring, Sg. Kepungit, E. of park boundary, c. 116°42°E 6°03’N, 480 m, ML, 26.xi.1986, J. Huisman; in RMNH. Description. — Male: Head hidden under prono- tum, interocular space greater than distance between ocellar spots and antennal sockets. Pronotum sub- oval. Tegmina and wings fully developed extending beyond end of abdomen, the former with oblique dis- coidal sectors. Hind wing intercalary vein present, media vein with one branch, cubitus vein with six ra- 100 mi (one may be branched). Front femur Type C;; pulvilli absent, tarsal claws symmetrical, simple, aro- lia minute. Abdominal terga unspecialized. Supraanal plate transverse. Subgenital plate asymmetrical, hind margin deeply, widely excavated, with a large whitish, membranous area bearing a cylindrical style at its ba- sal end (figs. 15A-C). Colour. — Head dark brown; two basal antennal segments dark brown succeeded by about 25 yellowish antennomeres, then three or four brown and three or four white segments, terminating in one or two brown ones (some terminal segments may have been miss- ing). Pronotal disk dark reddish brown, anterior and posterior areas darker, lateral regions hyaline. Tegmina with humeral region yellow-hyaline, remain- der brown-hyaline. Hind wing infuscated, darker along the costal vein area. Abdominal terga brown, su- praanal plate with a white medial zone. Abdominal sterna light brown, subgenital plate darker. Cerci with four basal segments brown, distal three segments yel- lowish. Coxae and femora brown, tibiae and tarsi pale. Rorn: Revision of Ctenoneura 0.25 mm Fig. 15. Ctenoneura spp. A-C. C. poringa sp. n., male holotype. A. supraanal and subgenital plates (dorsal); B. subgenital plate (ventral); C. subgenital plate (rear view). - D-G. C. /uma Sp. n., male holotype. D. supraanal and subgenital plates (dorsal); E. subgenital plate (left lateral); F. subgenital plate (right lateral); G. subgenital plate (ventral). Female: Unknown. Ctenoneura luma sp. n. Measurements (mm). — Length, 5.0; pronotum (figs. 15D-G) length x width, 1.6 x 2.0; tegmen length, 6.3; interoc- Type material. - Holotype, 4, Malay Penin., Perak, ular space, 0.5. Larut Hills, 4000-4500 ft., ii. —iii.1905, R. Shelford (incor- Etymology. — The species is named for Poring hot rectly labelled C. fulva, by Hanitsch); in HDEO. — Paratype. spring. Perak. HDEO: Perak, F.M.S., Jor Camp, 2000 ft, 16, 101 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 0.5 mm Fig. 16. Ctenoneura scutica sp. n., male holotype. A. supraanal and subgenital plates (dorsal); B. subgenital plate (ventral); C. subgenital plate (rear view). 6.ix.1922, 2d, at light, 27.viii.1922, E. Sumiind (labelled C. pumosa, mss name, by Hanitsch). Description. — Male: Head barely exposed, interoc- ular space greater than the distance between ocellar spots and antennal sockets. Pronotum suboval. Tegmina and wings fully developed, the former with oblique discoidal sectors. Hind wing with distinct co- stal veins, the proximal ones thickened distad, interca- lary vein present, media vein bifurcate (one of the ra- mi also branched), cubitus vein with six branches. Front femur Type C,; pulvilli absent, tarsal claws sym- metrical, simple, arolia minute. abdominal terga un- specialized. Supraanal plate trigonal, entire (fig. 15D). Subgenital plate asymmetrical, deeply excavated medi- ally forming two dissimilar lobes, with a slender style on the left side (the style was accidentally broken off and lost after the drawings were made); on the right side, a slender, thorn-like structure appears to arise from the margin of the plate but actually lies pressed tightly against the dorsal surface, and its apex pro- trudes beyond the the right lobe margin (figs. 15E-G). Colour. Head reddish brown. Pronotal disk red-- dish brown, the macula narrowly margined with yel- low, lateral portions hyaline. Tegmina and wings pale hyaline. Abdominal terga light brown, supraanal 102 plate with a white macula medially near the hind margin, subgenital plate darker except for a pale zone around the medial excavation. Cerci pale. Legs light brown. Female: Unknown. Measurements (mm). — Length, 5.5; pronotum length x width, 1.5 x 1.8; tegmen length, 6.3; interoc- ular space, 0.6. Etymology. — The specific name refers to the thorn- like structure that arises on the dorsal surface of the subgenital plate and protrudes beyond the right lobe. Ctenoneura scutica sp. n. (fig. 16) Type material. — Holotype d, Malaysia, Sabah, one half way Rd. Melligan, Long Pa Sia, 115°42°E 4°35’N, 1200 m, ML, 15.xii.1986, J. Huisman; in RMNH. Description. — Male: Head hidden, interocular space greater than the distance between ocellar spots or antennal sockets. Pronotum suboval. Tegmina and wings extending well beyond end of abdomen, the former with oblique discoidal sectors. Hind wing with intercalary vein, media vein with one bifurcated branch, cubitus with five branches. Front femur Type C,; pulvilli absent, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized. Supraanal plate symmetrical, transverse, short, hind margin entire. Subgenital plate asymmetrical, the left side concavely excavated bearing a small cylindrical style in a white membranous area; near the mid region of the plate is a deep, narrow, oblique (towards the left) excision with a long, stout, tapering whip-like process that curves upwards and anteriorly between the smooth walls of the excision (figs. 16A-C). Colour. — Head dark reddish brown; antennae brown, monochromatic. Pronotal disk reddish brown, darker near the anterior and posterior mar- gins. Tegmina with humeral and proximal part of the costal vein area yellowish-hyaline, remainder slightly darker. Hind wing with a yellowish tinge, costal vein region darker. Abdominal terga brown, lateral edges and terminal segments darker, supraanal plate with a RotH: Revision of Ctenoneura white medial zone. Abdominal sterna light brown, lateral edges darker, distal part of subgenital plate yel- lowish. Cerci brown, terminal segment lighter. Legs light brown. Female: Unknown. Measurements (mm). — Length, 5.6; pronotum length x width, 1.8 x 2.2; tegmen length, 7.6; interoc- ular space 0.6. Etymology. — The specific name refers to the whip- like process between the narrow incision on the sub- genital plate. Ctenoneura parascutica sp. n. (fig. 17) Type material. — Holotype, d, Malaysia, Sabah, base of S. Nologana, 1010 m, Crocker range, 115°55’E 5°10°N, UV, 10-11.x. 1986, J. Huisman (RMNH). Fig. 17. Ctenoneura parascutica sp. n., male holotype. A. pronotum; B. Supraanal and subgenital plates (rear view); C. sub- genital plate (right lateral); D. subgenital plate (left lateral); E. supraanal and subgenital plates (dorsal); F. left hind wing. 103 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 0.25 mm Fig. 18. Crenoneura kinabaluana sp. n., male paratype from Kiau, Mt. Kinabalu. A. supraanal and subgenital plates (dorsal); B. subgenital plate (rear view); C. subgenital plate (left lateral); D. subgenital plate (right lateral). Description. — Male: Head exposed, interocular space greater than the distance between ocellar spots and antennal sockets. Pronotum subelliptical (fig. 17A). Tegmina and wings fully developed extending beyond end of abdomen, the former with oblique dis- coidal sectors. Hind wing with the proximal three or four costal veins clubbed, intercalary vein present, media vein with one branch, cubitus vein with five branches (the right wing has one of the branches forked) (fig. 17F). Front femur Type C;; pulvilli ab- sent, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized. Supraanal plate transverse, hind margin entire (figs. 17B, E). Subgenital plate asymmetrical, hind margin excised, with a large dark, spine-like process that curves dor- sad, its apex hidden under the supraanal plate; the margins of the excision on either side of the spine-like process are irregular, and on the left side the region is unsclerotized; a small cylindrical style is on the left side (figs. 17B-E). Colour. — Head reddish brown; antennae brown (distal halves missing). Pronotal disk reddish brown, anterior and posterior border regions darker, a small yellowish area anteriorly on each side along the edges of the macula, lateral regions hyaline (fig. 17A). Tegmina with humeral and proximal costal vein re- gions yellowish-hyaline, remainder darker. Hind wing with a yellowish tinge in the costal vein region (fig. 17F). Abdominal terga brown, supraanal plate with a white macula posteromedially (fig. 17E). Abdominal sterna brown, subgenital plate darker. 104 Cerci brown, segments on distal half lighter. Legs pale. Female: Unknown. Measurements (mm). — Length, 5.2; pronotum length x width, 1.7 x 2.3; tegmen length, 7.3; interoc- ular space, 0.6. Comments. — I place parascutica close to scutica be- cause both have a large, darkly sclerotized, curved, spine-like process on the hind margin of the subgeni- tal plate. Ctenoneura kinabaluana sp. n. (fig. 18) Material. — Holotype, &, B.N. Borneo [Sabah], Mt. Kinabalu, Lumu Luma, 3500 ft., 16.iv.1929 [labelled C. fulva, by Hanitsch] (HDEO). — Paratypes. Sabah. Same local- ity and date as holotype, 3500 ft, 16; Mt. Kinabalu, Kamboranga, 7000 ft, 2d, 30.iii.1929 (HDEO); Malaysia, Sabah, Mt. Kinabalu Kg. Kiau, S. Kadamaian, 116°31’E 6°02’N, 720 m, UV, 1d 26.viii.1986, J. Huisman (RMNH). Description. — Male: Head hidden, interocular space greater than the distance between ocellar spots and antennal sockets. Pronotum suboval. Tegmina and wings fully developed extending well beyond end of abdomen, discoidal sectors of the former oblique. Hind wing with distinct costal veins, intercalary vein present, media vein with one branch (the branch may be bifurcate), cubitus vein with six branches (some may be forked; also one of the wings has two veins fused a short distance from their bases and then sep- arate). Front femur Type C,; pulvilli absent, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized. Supraanal plate transverse, hind margin convex, entire (fig. 18A). Subgenital plate asymmetrical, with a short, median process that ex- tends postero-obliquely, or dorsad, its distal end cury- ing anteriorly towards the supraanal plate; a cylindri- cal style is located on the left side and the margin to the right of the process is prolonged (figs. 18A-D). The length of the median process varies and is longest in the specimen from Kiau. Colour. — Head reddish brown; antennae brown. Pronotal disk reddish brown, lateral zones hyaline. Tegmina with humeral and costal vein zones yellow- hyaline, remainder brown-hyaline. Hind wing infus- cated, darkest in the costal vein zone. Abdominal ter- Roth: Revision of Ctenoneura ga brown, supraanal plate with a white macula ante- romedially. Abdominal sterna brown, distal margin of subgenital plate yellowish. Cerci and legs brown. The specimen from Kiau is smaller and darker than the others: head dark brown, basal three antenno- meres brown, remainder yellowish (distal segments of the antennae missing); supraanal plate yellow lateral- ly, and with a white area posteromedially (fig. 18A). Cerci with basal segments brown, distal cercomeres yellowish. Coxae, femora, and tibiae brown, tarsi lighter. Female: Unknown. Measurements (mm) (male from Kiau in parenthe- ses). — Length, 6.5-6.6 (5.3); pronotum length x width, 1.8-1.9 (1.5 x 2.1); tegmen length, 9.0 (6.7); interocular space, 0.6-0.7 (0.6). 0.25 mm Fig. 19. Ctenoneura triprocessa sp. n., male holotype. A. subgenital plate (ventral); B. subgenital plate (dorsal); C. subgenital plate (rear view); D. subgenital plate (right lateral); E. subgenital plate (left lateral); F. left hind wing. 105 TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Ctenoneura triprocessa Sp. n. (fig. 19) Type material. — Holotype d, Malaysia, SE. Sabah, Danum Valley Field C., c. 150 m, W.O., Mal. trap 5, 22.xi.-4.xii.1987, C. v. Achterberg & D. Kennedy (RMNH) [The type is fragmented, head and thorax, tegmina and wings, and terminal abdominal segments are mounted on cards]. Description. — Male: Head hidden, interocular space greater than the distance between ocellar spots and antennal sockets. Pronotum suboval. Tegmina and wings fully developed extending beyond end of abdomen, discoidal sectors of former oblique. Hind wing with a intercalary vein, media vein with one branch, cubitus vein with three branches (one of which may or may not be branched near its base) (fig. 19F). Front femora missing; other legs lack pulvilli, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized. Supraanal plate transverse, hind margin convexly rounded, entire. Subgenital plate asymmetrical, trigonal Ge, 194), hind margin with three long processes that curve dor- sad; arising on the dorsal surface of the plate near the left process is a long filament that protrudes between the left and middle processes; style absent (figs. 19B- E). Colour. — Head dark reddish brown; antennae with pale yellowish segments (about half the segments are missing). Pronotal disk reddish brown, slightly lighter on the anterior and posterior regions, lateral areas clear hyaline. Tegmina with humeral and proxi- mal half of the costal vein area yellowish-hyaline, re- mainder light brown-hyaline. Hind wing with yel- lowish tinge mostly in the costal vein zone (fig. 19F). Abdomen brown, distal part of supraanal plate white, subgenital plate yellowish brown. Female: Unknown. Measurements (mm). — Pronotum length x width, 1.5 x 1.8; tegmen length, 5.7; interocular space, 0.5. Etymology. — The specific name refers to the three prominent processes or prolongations on the hind margin of the subgenital plate. Ctenoneura murudensis Sp. n. (fig. 20) Ctenoneura fulva, Hanitsch (in part, 6 paralectotype) 1925: 101. Type material. — Holotype, d paralectotype (Type Orth. 300°/,) of Ctenoneura fulva Hanitsch, Mt. Murud [Sarawak], 6500 ft. (HDEO). Description. — Male: Head hidden, interocular Fig. 20. Crenoneura murudensis sp. n., male holotype. A. supraanal and subgenital plates (rear view); B. supraanal and sub- genital plates (dorsal); C. subgenital plate (ventral); D. subgenital plate (right lateral); E. subgenital plate (left lateral). 106 space greater than the distance between ocellar spots and antennal sockets. Pronotum suboval. Tegmina and wings fully developed extending beyond end of abdomen, the former with oblique discoidal sectors. Hind wing with distinct costal veins, the proximal ones clubbed distally, intercalary vein present, media vein forked about at the middle, cubitus with eight unbranched rami. Front femur Type C,; pulvilli ab- sent, claws simple, symmetrical, arolia small. Abdom- inal terga unspecialized. Supraanal plate transverse, deflexed, hind margin convex, entire (figs. 20A-B). Subgenital plate asymmetrical with a small, dark, cy- lindrical style located in a membranous area on the left side; a darkly sclerotized lobe with a U-shaped ex- cavation on its apical margin arises medially, and on the right side is a small acute prolongation (figs. 20A- B) Colour. — Head light reddish brown; antennae dark brown, monochromatic. Pronotal disk light red- dish brown, surrounding areas hyaline. Tegmina with humeral and proximal half of the costal vein zone yel- lowish-hyaline, remainder darker, hyaline. Wings pale, with a yellowish tinge, darker in the clubbed co- stal vein region. Abdominal terga uniformly light brown, supraanal plate without a medial white spot. Abdominal sterna yellowish-brown, distal part of subgenital plate yellow. Cerci light brown dorsad, darker ventrally. Legs light brown. Female: Unknown. Measurements (mm). — Length, 6.7; pronotum length x width, 1.8 x 2.3; tegmen length, 8.5; interoc- ular space, 0.7. Comments. — Hanitsch incorrectly considered this as fulva, but the subgenital plate of that species (fig. 8B) is distinctly different from murudensis (fig. 20C). Ctenoneura crassistyla sp. n. (figs. 21A-D) Type material. — Holotype, d, Pahang, F. M. S., Cameron’s Highlands, Tanak...., 4800 ft., 22.v.1931, H. M. Pendlebury (HDEO). Description. — Male: Head hidden, interocular space greater than the distance between ocellar spots and antennal sockets; antennae missing. Pronotum suboval. Tegmina and wings fully developed extend- ing beyond end of abdomen, the former with oblique discoidal sectors. Hind wings with distinct costal veins, media vein bifurcate, cubitus vein with six branches. Front femur Type C,; pulvilli absent, tarsal claws symmetrical, simple, arolia minute. Abdominal terga unspecialized, supraanal plate strongly trans- verse, very short, hind margin almost truncate, entire (fig. 21A). Subgenital plate asymmetrical, hind mar- gin deeply, unevenly excavated, with a large, stout Rorn: Revision of Ctenoneura style on the basal corner of the left side of the excava- tion; the apex of the style is split and a small piece of ‘fleshy’ tissue protrudes between the ends; right side with distal region concavely depressed (figs. 21B-D). Colour. — Head reddish brown. Pronotum with narrow yellow edging, disk reddish brown, lateral re- gions hyaline. Tegmina hyaline yellow, darker on the anal field region. Hind wings pale with yellowish mostly in the costal vein region and apical part of the anterior field. Abdominal terga light brown, supraa- nal plate yellowish with a medial white area near the hind margin (fig. 21A). Abdominal sterna brown, subgenital plate darker. Cerci pale dorsally. Legs light brown. Female: Unknown. Measurements (mm). — Length, 6.4; pronotum length x width, 1.9 x 2.5; tegmen length, 8.3; interoc- ular space, 0.7. Etymology. — The specific name refers to the large, stout, left style. Ctenoneura sp. Material. — South Vietnam. HDEO: 1d, S. Annam, Longbian Peaks, 5500-7500 ft, iv.-v.1918, C. B. Kloss (la- belled Ctenoneura major Hanitsch, by Hanitsch). Comments. — This species has a subgenital plate similar to that of crassistyla, with a large, stout, style on the basal corner of the left side (the style is not split at the apex). The media vein of the hind wing has two branches, and the cubitus vein has seven branches, one or two of which may be branched. The tarsal claws are long, simple, symmetrical, arolia ab- sent. The supraanal plate is transverse, hind margin weakly convex. This is the only species recorded from Vietnam. Measurements (mm). — Length, 7.8; pronotum length x width, 2.3 x 3.1; tegmen length, 10.2; inter- ocular space, 0.8. Ctenoneura simulans Bey-Bienko (figs. 21 E-F) Ctenoneura simulans Bey-Bienko, 1969: 833, figs. 2,3, (d) (Russian). Holotype 3, S. China, Yunnan: Hsiamonyang vicinity, 900-1100 m, 6.v.1957, Hung Kuang-ti (ZINR) [not examined]; 1970: 528 (English translation). Description (from Bey-Bienko). — Male: Small. Vertex passing roundly into frons, interocular space two and one half times wider than distance between antennal sockets. Pronotum somewhat less than one and one half times as wide as long. Hind wings with cross branches of the radius vein distinct, intercalary vein between the radius and media, almost straight; 107 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 21. Ctenoneura spp. — A-D, C. crassistyla sp. n., male holotype. A, supraanal and subgenital plates (dorsal); B. subgenital plate (ventral); C. subgenital plate (left lateral); D. subgenital plate (right lateral). — E, F. C. simulans Bey-Bienko; E. sub- genital plate (ventral); F. subgenital plate (rear view) (redrawn from Bey-Bienko, 1969). cubitus vein with six branches, the first one from the wing base bifurcate. Subgenital plate when viewed from below as in fig. 21E; when viewed from behind it has an irregular appendage on the right (fig. 21F). Cerci moderately long, fourth to sixth segments oval, tip of apical segment blunt, rounded. Colour. — Brownish yellow. Head dark chestnut. Pronotum chestnut brown with broad, hyaline lateral parts. Wings just perceptably infuscated. Female. Unknown. Measurements (mm). — Body length, 8.0; prono- tum length x width, 2.2 x 3.0; tegmen length, 9.0; overall length, 11.5. 108 Comments. — According to Bey-Bienko, outward- ly this species resembles misera, but is clearly distin- guished from all known species in the genus by the structure of the subgenital plate. Bey-Bienko does not mention whether or not there is a style (which could be small on the dorsal surface of the plate, and not visible in ventral or rear views). ACKNOWLEDGEMENTS I thank the Australian Biological Resources Survey (ABRS) for partial support, and the museums, cura- tors, and collection managers indicated in the intro- duction, who loaned me specimens. I am most grate- ful to Dr. Horst Bohn, Zoologisches Institut der Universitat Miinchen, for taking the time from his own taxonomic studies to review my manuscript. REFERENCES Bey-Bienko, G. Ya., 1957. Blattoidea. I. The results of the Chinese-Soviet zoologico-botanical expeditions to southwestern China 1955-1956 . — Entomologicheskoe Obozrenie, Moskva 36: 895-915. (Russian; English sum- mary). Bey-Bienko, G. Ya., 1969. New genera and species of cock- roaches (Blattoptera) from tropical and subtropical Asia. — Entomologicheskoe Obozrenie, Moskva 48: 831-862. (Russian). (English translation of this article: Entomological Review, Washington 48 (1970): 528- 548). Bruijning, C. F. A., 1948. Studies on Malayan Blattidae. — Zoologische Mededelingen Leiden, 29: 1-174. Hanitsch, R., 1925. On a collection of Blattidae from northern Sarawak, chiefly Mt. Murud and Mt. Dulit. — Sarawak Museum Journal 3 (1): 75-106. Hanitsch, R., 1927. On a collection of Blattidae from south- ern Annam. — Journal of the Siam Society, Natural History Supplement 7(1): 7-48. Hanitsch, R., 1928. Spolia Mentawiensia: Blattidae. — Bulletin Raffles Museum 1: 1-44. Hanitsch, R., 1929a. Fauna Sumatrensis. Blattidae. — Tijdschrift voor Entomologie 72: 263-302. Rotu: Revision of Ctenoneura Hanitsch, R., 1929b. Dr. E. Mjöberg’s zoological collec- tions from Sumatra. 11. Blattidae. — Arkiv for Zoologi 21A (2): 1-20. Hanitsch, R., 1932a. On a collection of blattids from the east coast of Sumatra. — Miscellanea Zoologica Sumatrana 62: 1-8. Hanitsch, R., 1932b. Beccari and Modigliani’s collection of Sumatran Blattidae in the Museo Civico, Genoa. — Annali del Museo civico di Storia naturale di Genova 56: 48-92. Hanitsch, R., 1933. On a collection of Bornean and other oriental Blattidae from the Stockholm Museum. — Entomologisk Tidskrift 54: 230-245. Princis, K., 1950. Entomological results from the Swedish expedition 1934 to Burma and British India. — Arkiv for Zoologi 1: 203-222. Princis, K., 1954. Kleine Beitrige zur Kenntnis der Blattarien und ihrer Verbreitung. VI. — Entomologisk Tidskrift 74 (1953): 203-213. Princis, K., 1963. Blattariae: Subordo: Polyphagoidea, Fam. Polyphagidae. /n Beier (ed.): Orthopterorum Catalogus 4: 77-172. ’s-Gravenhage. Princis, K., 1967. Kleine Beitrige zur Kenntnis der Blattarien und ihrer Verbreitung. X. — Opuscula Entomologica 32: 141-151. Princis, K., 1971. Blattariae: Subordo Epilamproidea, Fam.: Ectobiidae. /n Beier (ed.): Orthopterorum Catalogus. 14: 1039-1224. ’s-Gravenhage. Received: 12 November 1992 Revised version accepted: 19 February 1993 109 vere ed a UR, Be og smic tif a REET ri +. + > ne - IR Aa) Sei Re ar Pa a ne) AN CA ki i u mt uit sa cet oe rt PO ‘a | Y gta mr n sd ona 1 IR tar Leti ar vs i ques Te 4 Er CES NAE LATE " Lal ankam 1) 44 hj tach lata Lisi nA ' È Ne Gt oly lee at 3 ae En a et ed ne di Ch Calo ei u ULERY An m va ear! EN "cb ai “A iol adi iin Aue j in | N RES mann) Kahn ré > = tere Meis 4 fi N pre | Toth > 16 Clube Long ink CRO ig it ‘ill rat? im tari nad HEE | ace rs dA wary re ae tent EN it, Vigor weit Lsstiquiralt | Eee ide x i FE Are ng + ¥ a dou sa ; fant PSA eho atic) Mines gr cd ’ LS bitte le u ch, zn‘ ey Opty wea): (lui wa De LATES ‘ Fi = au n> Gare aa dee: ie es AH in i (DE ì “lie 4 | Lul af En meni Tuo TICA) | à | i I thi L pi ‚or 7 A lisi pen juli - a, are, sode pean a 5 A i DR 5 à : Le: =. POP En Si 1 IM d a HR nite dl ATS Bie 14 ee oy A Ne ee’ ee glas bj Del d Sy hans 32 ar eon Ae bre . ng 4 3 MAL dl Fi on. > I i D VE Ly f N = ® Lik I With LME = : n È \ È) "a Di MERS : ar TEA Ae ; A = ? n i A », 4, = 7 2 4 A SSR LL 4 tr È u é Du A = ¢ n i > ns = | ay È À . | = 4 j : = | a A er Ù bf) rari i ig bidens, En | 5 pui wire) maa? tel ame di raven A, a “ i la viari vatte à shal rpg rita pres kung) ran nl LOUE | | je 7 we on Te SE = > ad he, wen weicht six Inc the tien ‘one fea the \ ou #, qe loriae: Midagearirst bar hen ee. > rr ded rww re do hi Dex sce vs Hearn din adm hdi biegichy, che Wi sah on “= i "Coca chele LP: Caer) Lo atant$ Fa vs dels i MI vd apa, pation Oued, mara, Os ; nutro Braga serio: N ie Peuren ehem Brawl with ch : E puri Wings past grrceprably dia Lex ds na | E Lai Veoh: | law. > - =| en | pe u Mig e im Ms | HIT, n Publications Netherlands Entomological Society Prices in Dutch guilders excluding VAT and postage. Prices for members in brackets. MONOGRAFIEEN I. 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Price 25.— (10.-) Send your order to: Publishing Department Netherlands Entomological Society, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. The invoice will be sent together with the ordered titles. 111 Sth European Congress of Entomology University of Y ork, UK 29 August - 2 September 1994 e Insect life histories « Habitat management, creation & restorations Population processes & spatial dynamics * Biodiversity: Does taxonomy matter?* eManagement of pests & beneficial insects + Insects as indicators of environmental quality* Themes to be interpreted broadly. Offers of papers & workshops welcome. For details and further mailings contact IFAB Communications Institute for Applied Biology University of York York YO1 5DD, UK Tel: +44 (0)904 432940 Fax: +44 (0)904 432917 Hosted by the Royal Entomological Society Tijdschrift voor Entomologie Volume 136, no. | Articles | J. G. M. Cuppen Distribution and ecology of Hydraena in The Netherlands (Coleoptera: Hydraenidae). II R. 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III Publications Netherlands Entomological Society. 112 Announcement: 5th European Congress of Entomology, University of York, U.K. © Nederlandse Entomologische Vereniging, Amsterdam Published | July 1993 ISSN 0040-7496 Volume 136, no. 2, 1993 ISSN 0040-749 Tijdschrift voor iw Entomologie À journal of systematic and evolutionary entomology since 1858 Published by the Nederlandse Entomologische Vereniene 12 ae Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address 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 Published with index of volume 136 (1993). Graphic design Ontwerpers B.V., Aad Derwort, ’s-Gravenhage GEORGE W. BECCALONI MCZ NERC Centre for Population Biology, Imperial College, UK L IB RA R Y JAN 1 4 1994 A NEW SPECIES OF EXTATOSOMA GRAY (PHASMATODEA: PHASMATIDAE) FROM PAPUA NEW GUINEA, WITH REMARKS ON THE SPECIES IN ime GENUS HARVARD UNIVERSITY Beccaloni, G. W., 1993. A new species of Extatosoma Gray (Phasmatodea: Phasmatidae) from Papua New Guinea, with remarks on the species in the genus. — Tijdschrift voor Entomologie 136: 113-123, figs. 1-16, tabs. 1-2. [ISSN 0040-7496]. Published 10 December 1993. Extatosoma carlbergi sp. n. (Phasmatidae), is described from Papua New Guinea. The biology, ecology and distribution of this and the three other species of Extatosoma Gray is discussed and a hypothesis is advanced to explain the discrete altitudinal ranges of the two New Guinean spe- cies. The phylogenetic relationships of the species are considered and a key to the adult females is presented. G. W. Beccaloni, Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK. Key words. — Phasmatodea, Phasmida, Extatosoma, New Guinea, Australia, biology, ecology, distribution, phylogeny. The genus Extatosoma is widely known due to fa- miliarity with an Australian species, £. tiaratum (MacLeay 1826). This is one of the stick-insects most commonly cultured by amateur enthusiasts (Bragg 1991) and it is also a popular laboratory animal on which many physiological, behavioural and morpho- logical studies have been conducted (for references see Carlberg 1983, 1987a). Two other species are currently recognised as be- longing to the genus : E. bufonium Westwood, 1874 from Australia and £. popa Stal, 1875 from New Guinea. Another nominal species, E. elongatum Froggatt, 1922, was synonomised with E. bufonium by Vickery (1983). The holotype of E. bufonium (in the Hope Entomological Collection, Oxford, UK) is a penultimate (?) instar female nymph, not a male as stated by Vickery (1983), and Froggatt (1922) based E. elongatum on the adult female of this species. A third, as yet undescribed, species may exist in the vi- cinity of Brisbane, Queensland, Australia (P. D. Brock, in prep.). The adult female of this taxon (fig. in Wilson 1991) is superficially similar to E. tiaratum in appearance, while the egg resembles that of £. bu- fonium. Both £. popa and E. bufonium are poorly known and only the former species has been cultured in cap- tivity (Harman 1985). The males of these two species have not been figured or formally described, although Harman briefly noted that the male of £. popa can be distinguished from that of E. tiaratum by its clear wings and a white dot behind its head. From person- al observation of a series of preserved adult male £. popa cultured by Harman and deposited in the BMNH, it is evident that the ‘white dot’ referred to is in fact a pale v-shaped marking on the mesonotum, similar to that possessed by the adult female of this species. The existence of a second New Guinean species of Extatosoma was first suggested in print by Harman (1985) who stated that there is ‘....another species [of Extatosoma] which occurs in the mountainous regions of Papua New Guinea. I have seen a number of spec- imens in the collection at the Wau Ecology Institute Museum’. During July and August of 1990 the au- thor led an Imperial College (University of London) expedition to Papua New Guinea (PNG). For the first four weeks we were based at the Wau Ecology Institute (WEIC) and the author took the opportunity to investigate Harman’s claim. Three adult females of an undescribed Extatosoma were present in the WEIC collection, together with a note stating that the taxon was new. A reward was offered to two Papuan youths to col- lect living specimens of this species and one day later an adult female (fig. 1) was received. This individual was placed in a cylindrical net cage on cut foodplant standing in water and was kept alive for twelve days for observation. Despite further requests and a search of the locality 113 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 1. Extatosoma carlbergi sp. n. — Holotype © resting on twig of Calliandra surinamensis Benth. from which the specimen had been collected, no other individuals were obtained. Several local Papuans who were interviewed stated that they were familiar with this species but that it was only rarely seen (more rarely than E. popa). MATERIALS AND METHODS The terminology used to describe eggs is after Clark (1976a), with the following additions and mo- difications : ‘total egg length’ is the length of the egg capsule plus the length of the operculum ; ‘length of operculum’ is the maximum length of the operculum from its base to its summit ; ‘diameter of operculum’ is the maximum diameter across the base of the oper- culum ; ‘micropylar arm’ is a new term given to the lateral extensions of the micropylar plate located on either side of the micropylar cup region ; ‘length of micropylar arm’ is the length of one micropylar arm measured from the centre of the micropylar cup to the apex of the arm; “distance across micropylar arms’ is the maximum distance between the tips of the mi- cropylar arms ; ‘height of micropylar mound’ is the height with which the micropylar plate is raised abo- ve the surface of the egg capsule and is measured mid- way along the plate’s length; ‘chorionic membrane’ is a new term given to the membrane or layer which co- vers the entire outer surface of the egg. This membra- ne expands and detaches from the egg surface if the egg is immersed in alkali (e.g. sodium hydroxide solu- tion). The membrane is thinner and weaker around 114 the base of the operculum and the portion of mem- brane covering the operculum remains attached to it when it detaches on hatching of the egg. The eggs of all four Extatosoma are covered by a chorionic mem- brane, but it appears to be absent from the eggs of other stick-insect species (Clark 1976b); ‘capitulum’, in light of the recent discovery that this structure has a function analogous to the elaiosome of certain plant seeds, and acts as a handle which enables ants to carry stick-insect eggs back to their nests (Compton & Ware 1991, Hughes & Westoby 1992), the author proposes to restrict Clark’s (1976a) definition to structures attached to and protruding from the sur- face of the operculum. The coloration of £. carlbergi is described from slides of the living holotype taken on ‘Kodachrome 64’ slide film under natural lighting conditions. Specimens of E. carlbergi and E. popa collected by the author were killed by freezing and preserved by injec- tion of embalming fluid (formula in Cogan & Smith 1974). Eggs (fully developed) were removed from the NS i si ik # 4 3 1 ad TA a Fig. 2. Extatosoma carlbergi sp. n. — Holotype 9, dorsal hab- itus. 20 mm Figs. 3-4. Egg of Extatosoma carlbergi sp. n. taken from ovi- positor of holo- type. — 3, dorsal view; 4, lateral view. abdomen of the £. carlbergi holotype by first relaxing the specimen (in a sealed plastic box with damp tissue paper in the base) and then by softening a section of intersegmental membrane between tergites and ster- nites with 10% aqueous ammonia solution, so that an incision could easily be made and the eggs removed with forceps. The chorionic membrane was removed from eggs for study, by placing the eggs for c. 5 min- utes in a 5% aqueous solution of sodium hydroxide at room temperature. Drawings were produced using a drawing tube on a Wild M5 stereomicroscope. All measurements are given in millimetres. Measure- ments of eggs (accurate to 0.01 mm) were made using an ocular micrometer, except for micropylar plate length, which because of the plate’s curvature, was first drawn using a drawing tube and then measured using inelastic fishing line and a ruler. All other meas- urements (accurate to 1 mm) were taken from speci- mens using a pair of dividers and a ruler. Abbreviations of depositories BMNH: The Natural History Museum, London, England; BPBM: Bishop Museum, Hawai'i; TPNG: Central Reference Insect Collection, Department of Agriculture and Livestock, Port Moresby, Papua New Guinea; FICB: National Forest Insect Collection, Forest Research Institute, Lae, Papua New Guinea; NHMN: Nottingham Natural History Museum, Nottingham, England; UPNG: University of Papua New Guinea, Port Moresby, Papua New Guinea; WEIC: Wau Ecology Institute, Wau, Papua New Guinea. BECCALONI: A new species of Extatosoma ‘TAXONOMY Extatosoma carlbergi sp. n. (figs. 1-5, 8, 12-14) Type material. — Holotype 9: Papua New Guinea, Morobe Province, Wau, golf course, c. 1150 m, ex Calliandra surinamensis Benth., 14.vii.1990, G. W. Beccaloni (BMNH). — Paratypes: PAPUA NEW GUINEA: 29, Southern Highlands Province, Tari, 1600 m, ex Casuarina, 4.x.1963, R. Straatman (BPBM); 19 (fragmented), Chimbu Province, Karimui, 4.vi.1961, Gressitt (BPBM); 1 2, Morobe Province, Wantoat, feeds on Casuarina, 1.xii.1957, R. W. Paine (BMNH). Additional material examined. — NEW GUINEA: 19, no other data (NHMN); PAPUA NEW GUINEA: 22, Southern Highlands Province, Pangia, defoliating Casuarina oligodon, 1.vi.1971, J. Lowien (FICB); 19, Southern Highlands Province, Pangia, found on Casuarina trees, 30.iv.1971, J. Wallis (TPNG); 22, no data (WEIC); 12 (nymph, penulti- mate ? instar), Morobe Province, Wau, 1200 m, 14.x.1966, G. A. Samuelson (BPBM); 1 2, Morobe Province, Wau, 1100 m, collected on Casuarina sp., 10.xii.1974, A. D. Hart (WEIC); 1 2, Morobe Province, Wau, Wau Ecology Institute Office, 1150 m, 27.xi.1986, J. Somp (UPNG). Description Based on adult female holotype and four adult fe- male paratypes. Measurements of selected structures given in table 1. Male unknown. Female (adult) . — Head typical for the genus; hy- pognathous with conical occiput. Occipital medial spines large and robust; four pairs anterior to crest and two pairs posterior to crest. A large lamellate compound spine present on either side of occipital 115 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 5-6. Lateral views of ab- domens of adult Q Extatosoma spp. showing spi- nation on tergites 3-10. — 5, Extatosoma carlbergi sp. n. (holotype); 6, Extatosoma tia- ratum. crest. Eyes small and globular, situated anterolateral- ly. Ocelli indicated by slight swellings covered by in- tegument. Antennae simple, with 25 segments (n=2). A large spine present between eyes and antennal sock- ets. Pronotum elongate-rectangular, slightly rounded posteriorly. Spines around margin variable in size and number; a pair of large medial lamellate spines usual- ly present on posterior third of segment. Mesonotum with a well developed pair of pre-me- dian spines and a large pair of compound lamellate spines situated between tegminal bases. Lateral bor- ders with 3-6 large spines, the two directly above the tegminal bases often confluent basally. Tegmina re- duced, lobose, extending to posterior margin of meta- notum. Metanotum with a pair of well developed lamellae above posterior margin. Wings rudimentary, extend- ing into anterior third of first tergite. Tergites one to nine bearing a pair of medial com- pound lamellate spines on the posterior third, with a well developed simple spine posterior and to either side of these. The lamellate spines on tergites five and six especially well developed (fig. 5) ; those on tergites seven to nine are sometimes simple. The posterolater- al margins of tergites two to nine expanded laterally 116 and strongly serrated or spined. The anterolateral margin of the tenth tergite is similarly produced. Lateral expansions of tergites five to seven are c. twice the size of those on the other tergites. Tenth tergite usually with a pair of medial spines on posterior half and the hind margin has large posteriorly directed spines (fig. 5). Supra-anal plate with large pair of posteriorly di- rected medial spines. Cercus 2.6 mm in length, bi- lobed, setose (fig. 14). Both structures concealed be- neath tenth tergite. Operculum keel-shaped, the anterior third ventral- ly expanded with a serrated edge (fig. 5). Sternites two to seven with many long to short irregularly arranged robust spines. A pair of double weakly confluent me- dial spines may be present posteriorly. Spination of sternites four to six reduced in comparison with that of sternites two to three. Meso- and metathoracic ba- sisterna with few irregular short robust spines. Ventral margins of meso- and metathoracic episterna with long to short laterally or ventrally directed spines. Fore femur with strongly serrated dorsal and later- al expansions. Fore tibia with dorsal expansion arcu- ate and ventral expansion denticulate (fig. 8). Proximal tarsomere of fore leg expanded into lobe BECCALONI: A new species of Extatosoma Table 1. Measurements of selected structures of type specimens and female (? penultimate instar) nymph of £. carlbergi. All measurements in mm (accurate to 1 mm). “excluding occipital spines; “n = 1. Structures measured Holotype Paratypes Nymph (Lengths) (n=4) x+SD Range Body 128 IEN) 116 - 140 70 Antenna 27 - — 11 Head ' 19 195=#13 18-21 11 Pronotum 9 10.5 + 1.3 Se 2 5 Mesonotum 15 16.0 + 0.8 15-17 10 Metanotum 12 11.5 + 0.6 lil = 12 5 Fore femur 25 DD SEA DAT 15 Median femur 20 20.3 + 1.0 19-21 11 Hind femur 30 28.8 + 1.0 28 - 30 15 Fore tibia 23 23.5 + 0.6 23 - 24 13 Median tibia 21 Mil) ze 12 VO = 22 12 Hind tibia 32 DAB ae 17 3135 16 Operculum 26 24.3 + 0.5 VA = WS) 7 dorsally towards distal end. Dorsal surfaces of median and hind femora (figs. 12-13) with serrated expansion distally. Outer ventral margin serrated and expanded ventrally. Inner ventral margin spined and expanded distally. Dorsal expansions of hind tibia usually strongly arcuate (fig. 12), rarely weakly so (fig. 13); median tibia likewise. Inner and outer ventral mar- gins of median and hind tibiae with large ventrally di- rected spines. Legs and body densely setose. Coloration in life: Mesonotum with conspicuous v-shaped marking which is paler than rest of segment (this marking is usually white or cream in preserved specimens). Arms of ‘v’ extend from anterolateral cor- ners of segment, with base of ‘v below middle of seg- ment. Area between arms usually black or dark brown. Dorsal and ventral surfaces of body (includ- ing spines, tegmina and wings) pale green with dark- er green, white and black mottling; mirrored on each side of body. Intersegmental membranes cream to light brown. Legs with similar coloration to rest of body; dorsal surfaces with a higher incidence of larger black and dark green marks often arranged as trans- verse stripes; ventral surfaces (less so of fore legs) apos- ematically patterned with thick irregular black, ochre- ous and white transverse bands. Median and hind tibiae often with black predominating. Nymph (female, penultimate ? instar). — Measurements given in table 1. Morphology similar to that of adult, except that wings not developed (small, c. 1.5 mm long wing buds present) and oper- culum is relatively shorter; not extending beyond pos- terior margin of ninth tergite and not covering the su- pra-anal plate or cerci. Coloration in life presumed similar to that of adult, as coloration is similar to that of the preserved adult specimens examined (i.e. with light to dark brown replacing pale green ground colour). Egg (figs. 3-4). — Measurements given in table 2. Operculum conical. Chorionic membrane transpar- ent, not thickened and modified into a capitulum on the operculum (see below). Capsule smooth and glossy, marked with mottled pattern of black and grey on cream. Micropylar plate cream, slightly raised above surface of capsule on micropylar mound, with lateral arm on either side of the micropylar cup region. Etymology This species is named in honour of Dr. Ulf Table 2. Means, sample standard deviations and ranges of dimensions of 17 eggs from E. carlbergi holotype (1 egg tak- en from ovipositor, 16 extracted from abdomen). All meas- urements in mm (accurate to 0.01 mm). Egg dimensions x + SD Range Total egg length 6.54+0.12 6.40 - 6.85 Capsule length 5.170072 5062531 Capsule height 4.49 +0.03 4.42 - 4.56 Capsule width (maximum) SDE 3771025284 Length of operculum 1.37220.12 NAR = 1154 Diameter of operculum (maximum) 2.32+0.07 2.20 - 2.43 Micropylar plate length 6.09+0.28 5.70 - 6.73 Micropylar plate width (at middle) 0.67 +0.03 0.61 - 0.70 Length of micropylar arm 1.74+0.06 1.60 - 1.79 Distance across micropylar arms 3.00+0.07 2.88 - 3.07 Height of micropylar mound 0.15+0.03 0.13 - 0.19 iy TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 L L Syn 118 Carlberg, in recognition of his work on and passion for Extatosoma. Biology and behaviour In common with the other members of the genus, E. carlbergi is arboreal and when not feeding hangs motionless upside down under thin branches of the host tree (fig. 1). The holotype was collected on Calliandra surinamensis (Leguminosae), an introduc- tion from South America commonly grown in PNG gardens (Verdcourt 1979). In captivity the specimen accepted Leucaena sp. (Leguminosae) and guava, Psidium sp. (Myrtaceae). Both originate from the Neotropics and the former is cultivated in Wau as a shade tree for coffee, while the latter is grown for its fruit. J. Somp (pers. comm.) informed the author that he has collected E. carlbergi on Sterculia sp. (Sterculia- ceae), and Casuarina sp. (Casuarinaceae) is the only foodplant recorded on the data labels of this taxon examined during the course of this study. Both £. tiaratum and E. popa are also polyphagous. The latter has been collected on Casuarina, Diospyros ferrea (Ebenaceae), C. surinamensis and Hibiscus sp. (Malvaceae) (taken from data labels listed in Appendix) and it will also feed on Leucaena and Psidium in captivity (pers. observation). For a list of the foodplants of E. tiaratum see Carlberg (1987b). The only foodplant known for E. bufonium is Japanese plum, ? Prunus cerasifera (Rosaceae) (from data label of adult female in Queensland Museum, Brisbane, Australia; collected at Mt. Tamborine, near Brisbane, December 1924). When disturbed the £. carlbergi holotype was ob- served to exhibit a defensive behaviour similar to that shown by adult females of £. tiaratum (Carlberg, 1981) and E. popa (pers. observation). The abdomen is curled over the body, presenting the dense spina- tion on the sternites towards the potential predator. The hind legs are raised and extended to reveal the aposematic pattern of black, white and ochreous bands on the ventral surfaces of the femur and tibia. This threat posture is maintained until any possible danger has passed. Should a predator approach too close, the insect will attempt to grasp it between the femur and tibia of the hind legs; retracting and ex- tending the legs with a rapid, synchronous scissor-like action. The ventral margins of these appendages are furnished with sharp downwardly directed spines, which are capable of inflicting injury to human skin if caught between them. The egg of E. carlbergi (figs. 3-4), which resembles BECCALONI: A new species of Extatosoma a seed, is held in the ovipositor for several hours and is then catapulted to the ground with a vigorous flick of the abdomen. Eggs of E. popa (A. J. E. Harman, pers. comm.) and £. tiaratum (Carlberg, 1984) are deposited in a similar manner. ECOLOGY AND DISTRIBUTION OF THE EXTATOSOMA SPECIES The known distributions of the two New Guinean Extatosoma species is shown in fig. 16. The locality re- cords plotted on this map were taken from data labels of specimens examined by the author or from the li- terature. Records for E. popa are listed in the Appendix while those for £. carlbergi are listed under the species description. Dubious records or possible misidentifications have been omitted. From fig. 16 it can be seen that £. popa has a wider recorded geographical distribution than £. carlbergi, ranging from Jayapura in Irian Jaya to Mt. Obree in the Owen Stanley Range, PNG. Altitude appears to be the most important factor in determining the dis- tributions of these two species. Giinther (1929) placed E. popa in an altitude band of 600-1500 m above sea level. However, the highest that this species has been recorded is c. 1200 m at Budemu, PNG. In contrast, all known specimens of Æ. carlbergi have been collected in the range 1100 m (Wau, PNG) to 1600 m (Tari, PNG). Wau (1100-1200 m) and Wantoat, PNG, (1200 m), where the species are sym- patric, are located within the c. 100 m altitudinal overlap of the two species ranges. The altitudinal ranges of E. popa and E. carlbergi match the distributions of two discrete vegetation belts, lower montane rain forest and mid-montane rain forest respectively, which differ from each other both in terms of species composition and in the struc- ture and physiognomy of the vegetation. Lower mon- tane rain forest has a mean canopy height of 30-40 m and is distinctly drier in aspect than the mid-mon- tane, with few species of ferns and mosses. In con- trast, the mid-montane rain forest has a mean canopy height of 20-30 m and its main characteristic is the abundance and high species diversity of terrestrial and epiphytic ferns, mosses and lichens (Johns 1982). It is proposed that the gross differences in colour pattern and morphology seen between E. popaand E. carlbergi (nymphs and adult females only; adult males of Extatosoma are macropterous and differ from nymphs and adult females in both body form and be- haviour) are the result of these species evolving cryp- Figs. 7-15. Extatosoma spp. adult © appendages. — 7-8, lateral views of left fore tibia and proximal tarsomere: 7, tiaratum, 8, carlbergi sp. n. (holotype). 9-13, lateral views of left hind femur, tibia and proximal tarsomere: 9, tiaratum; 10, popa; 11, bu- fonium; 12, carlbergi sp. n. (holotype); 13, carlbergi sp. n. (paratype from Tari, PNG). 14 - 15, ventral views of left cercus: 14, carlbergi sp. n. (holotype); 15, tiaratum. 119 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 16. Sketch map of Papua New Guinea and eastern Irian Jaya, showing © E. carlbergi OS a À. popa == known distri- butions of E. popa (13 records) and E. carlbergi (11 records). PAS AURA: NEW GAU RENDER < > < = £ < [ad PER eee oord PORT MORESBY © sis adapted to different habitat types. Once evolved this would tend to restrict the species to the habitat in which it is best concealed from predators. The body coloration of £. popa is primarily brown and when the insect is hanging under a branch at rest the foliose expansions of the legs (fig. 10) and abdo- men give it a resemblance to a bunch of dried leaves. In the lower montane rain forests, where this species is found, dry brown leaves are presumably more com- mon than in the wetter mid-montane forests occu- pied by £. carlbergi. In these latter forests, the bark of trees and other plants is covered by dense growths of lichens, and mosses are three times as abundant as in the lower montane (Johns 1982). Thus the dark green, white and black mottling of £. carlbergi, to- gether with the arcuate flanges of the tibiae (figs. 8, 12) and small lateral expansions of the tergites (fig. 2), are probably adaptations for resembling foliose epi- phytic lichens, such as the abundant Usnea, or leaves covered with epiphyllous growth. The crypsis of the Australian Extatosoma mirrors that of the New Guinean species, with one, £. tiara- tum, a leaf mimic (Brock 1992) and the other, £. bu- fonium, a probable lichen mimic. Both taxa are re- stricted to the coastal forests of eastern Queensland and New South Wales (NSW), although the former species has doubtfully been recorded from Lord Howe Island (Redtenbacher 1908, Giinther 1931). E. tiaratum is distributed from Daintree in north-east 120 Queensland (data from a final instar female nymph in BMNH) to Kiama, near Sydney, NSW (Froggatt 1922) and appears to range further inland than Æ bufonium (Beccaloni, unpublished). It is possible that literature records of ‘E. tiaratum from Brisbane southward rep- resent the undescribed taxon referred to earlier. E. bufonium ranges from Moreton Bay, near Brisbane, south-east Queensland (data from a female nymph in BMNH) to Camden, near Sydney, NSW (Froggatt, loc. cit.). This species may also occur in north-east Queensland, as a female nymph probably belonging to this taxon was collected in Atherton in 1989 (P. D. Brock, pers. comm.). A patchwork of different forest types (ranging from rain forest to dry eucalypt wood- land) is found in eastern Queensland and NSW and without further detailed study it is not possible to de- termine in which habitats the Australian Extatosoma live. However, considering the types of crypsis exhib- ited by these species, it is hypothesised that E. bufoni- um will be found to inhabit a habitat type with a higher representation of lichens and other epiphytes than that in which £. tiaratum occurs. COMPARATIVE MORPHOLOGY AND PHYLOGENETIC RELATIONSHIPS The characters discussed below are taken from adult females and eggs, as only males of E. popa and E. tiaratum are known and they do not exhibit suffi- cient characters to make comparisons worthwhile. Both £. popa and E. tiaratum range from light to dark brown in colour, although the latter species is known to have a green form which is not uncommon. The dorsal foliose expansions of the femur and tibia of the legs (figs. 7, 9-10) are broad and well developed in both species, as are the lateral expansions of the fifth to seventh tergites (as in fig. 6). The lateral ex- pansions of tergites two to four and eight to nine are reduced (£. popa) or obsolete (E. tzaratum). The coloration of E. carlbergi and E. bufonium is again similar: both are green, with white and black mottling in varying proportions. The dorsal expan- sions of the tibiae (figs. 8, 11-13) are arcuate in both taxa and the lateral expansions of the fifth to seventh tergites (as in figs. 2, 5) are much reduced relative to those of E. popa and E. tiaratum. Lateral expansions of tergites two to four and eight to nine are relatively well developed. The superficial similarities in coloration and body form (listed above) between £. popa and E. tiaratum and between £. carlbergi and E. bufonium, are prob- ably due to convergence upon analogous cryptic mor- phologies and thus do not reflect the phylogenetic re- lationships of the group. Characters not obviously related to crypsis (see below) are incongruent with those characters involved in crypsis and suggest a grouping reflecting the biogeographical division of the taxa (New Guinea / Australia). The New Guinean taxa share a number of features not possessed by the Australian taxa. Both are larger than the Australian species and the body spination is relatively longer and more robust. Both possess a dis- tinctive pale v-shaped marking on the mesonotum, and the pair of medial lamellate spines on the posteri- or third of the meso- and metanotum are large and well developed. The paired medial lamellae of the fifth and sixth terga each occupy about one-third of the length of the segment and their basal width is less than their height (as in fig. 5). The posterior margin of the tenth tergite has large posteriorly directed spines (as in fig. 5) and the apex of the cercus is bi- lobed (as in fig. 14). The ventral expansion of the fore tibia has a denticulate margin (as in fig. 8) and the in- ner ventral margin of the median and hind femur is expanded ventrally towards its distal end (figs. 10, 12- 13). The egg of E. popa (fig. 100 in Sellick, 1980) is virtually identical in appearance and dimensions to that of E. carlbergi (figs. 3-4). The Australian taxa share the following characters: the pale mesonotal v-shaped marking (of E. popa and E. carlbergi) is absent in the adults (although present in newly hatched nymphs of £. tiaratum) and the pair of medial spines on the posterior third of the meso- and metanotum are poorly developed and, if double, are only weakly confluent basally. The paired medial BECCALONI: A new species of Extatosoma lamellae of the fifth and sixth tergites each occupy about one half of the segment and their basal width is greater than their height (as in fig. 6). The posterior margin of the tenth tergite has small posteriorly di- rected spines (as in fig. 6), which may be subobsolete, and the cercus is tapered towards its apex (as in fig. 15). The margin of the ventral expansion of the fore tibia is entire (as in fig. 7) and the inner ventral mar- gin of the median and hind femur is only weakly (or not) expanded ventrally towards its distal end (figs. 9, 11). The eggs of E. tiaratum (fig. 8 in Clark, 1976a) and E. bufonium (pers. observation) share a number of characters not possessed by eggs of the New Guinean taxa. Eggs of both species lack micropylar arms and the opercula are only c. half the length (cor- rected for differences in egg size) of E. popa | E. carl- bergi eggs. In addition, the chorionic membrane on the opercula of eggs of the Australian species is thick- ened and modified into an opaque hollow dome or capitulum. The chorionic membrane on the opercula of eggs of the New Guinean species is not modified into a capitulum, and instead is transparent and lies in contact with the surface of the operculum. (Note that the capitulum of £. tiaratum is relatively much larger than that of £. bufonium. Also note that the egg of E. bufoniumis c. half the size of that of £. tiaratum, the capitulum is dark brown in colour and the egg capsule has a pitted and irregular surface. The capitu- lum of £. tiaratum is cream in colour and the egg cap- sule is smooth and glossy like the capsules of the New Guinean taxa). It has not been possible to determine the polarity of the above characters, as appropriate outgroups to Extatosoma have not been identified. The other gene- ra (e.g. Podacanthus Gray, Tropidoderus Gray, Didymuria Kirby) placed with Extatosoma in the tribe Tropidoderini, differ greatly from Extatosoma in both adult and egg morphology preventing useful compar- ison. However, the non-polar characters described above imply that either the two New Guinean species are sister taxa and/or the Australian species are sister taxa. This ambiguity can only be resolved through polarity determination. Key to the adult females of Extatosoma 1. Mesonotum with conspicuous pale v-shaped marking; ventral expansion of fore tibia with den- tieulatermazsın (asin fig 6). D — Mesonotum lacking a pale v-shaped marking; ventral expansion of fore tibia with entire margin (ASUS) ot ann, 3 2. Lateral expansions of tergites 5-7 small, non- overlapping (fig. 2); dorsal expansions of median and hind femur arcuate (figs. 12-13); body color- ation pale green, with white and black mottling 121 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 pied seen che) tas MOD ba A aed ia carlbergi Beccaloni — Lateral expansions of tergites 5-7 large and over- lapping; dorsal expansions of median and hind femur broad, not arcuate (fig. 10); body colora- tion light to dark brown ....................... popa Stal 3. Tergites 2-4 and 8-10 bearing lateral expansions; dorsal expansions of median and hind femur ar- cuate (fig. 11); body coloration green, with dark- er green, white and black mottling...................... ART MA RIST EEN Ci bufonium Westwood — Tergites 2-4 and 8-10 without lateral expansions; dorsal expansions of median and hind femur broad, not arcuate (fig. 9); body coloration brommtongmeen MINO Ein otter ACKNOWLEDGEMENTS This paper is based on the author’s final year BSc (Hons) project, written while an undergraduate in Zoology at Imperial College (University of London). The author would like to thank Mrs J. Marshall and Mr J. Reynolds for assi- stance, advice and for enabling him to conduct much of his work in the BMNH. He is grateful to the following individu- als for either sending him specimens or enabling him to exa- mine material included in this paper: Mr S. Fellenberg (Sydney, Australia); Dr M. Hopkins (UPNG); Prof. R. Kumar (Department of Agriculture and Livestock, Port Moresby, PNG); Dr G. McGavin (Hope Dept. of Entomology, Oxford, UK); Dr G. B. Monteith (Queensland Museum, Brisbane, Australia); Dr M. S. Moulds (Australian Museum, Sydney, Australia); Dr H. Roberts (Forest Research Institute, Lae, PNG); Dr H. A. Rose (University of Sydney, Australia); Mr J. Somp (WEIC); and Mr J. S. Strazanac (BPBM). The author’s thanks also go to Mr P. D. Brock and Mr A. J. E. Harman (UK) for providing useful information, to Dr U. Carlberg (Sweden) for his enthusiasm and help, and to Jerry and Willie of Wau (PNG) for collecting the holotype of E. carlbergi. The author is also very grateful to Mr P. E. Bragg (UK), Dr V. K. Brown (Imperial College, UK), Mr R. I. Vane- Wright (BMNH) and Dr M. Wilkinson (BMNH) for reading and commenting on the manuscript and he is indebted to all the organisations and individuals who approved and spon- sored the ‘Imperial College Expedition to Papua New Guinea 1990’. REFERENCES Bragg, P., 1991. Species census returns 1991. — Phasmid Study Group Newsletter 46: 6. Brock, P. D., 1992. Rearing and studying stick and leaf in- sects. — The Amateur Entomologists’ Society, Handbook 22: i-v, 1-73. Carlberg, U., 1981. Defensive behaviour in adult female Extatosoma tiaratum (Macleay) (Phasmida). — Entomologist's Monthly Magazine 116: 133-138. 122 Carlberg, U., 1983. Bibliography of Phasmida (Insecta) I. 1970-1979. — Spixiana 6: 27-43. Carlberg, U., 1984. Oviposition behaviour in the Australian stick insect Extatosoma tiaratum. —Experientia 40: 888- 889. Carlberg, U., 1987a. Bibliography of Phasmida (Insecta) VI. 1980-1984. — Spixiana 10: 147-156. Carlberg, U., 1987b. Culturing stick- and leaf-insects (Phasmida)-A review. — Zeitschrift Versuchstierkunde 29: 39-63. Clark, J. T., 1976a. The eggs of stick insects (Phasmida): a review with descriptions of the eggs of eleven species. — Systematic Entomology 1: 95-105. Clark, J. T., 1976b. The capitulum of phasmid eggs (Insecta: Phasmida). — Zoological Journal of the Linnean Society 59: 365-375. Cogan, B. H. & K. G. V. Smith, 1974. Insects. — Instructions for Collectors 4a, 5th Edition, British Museum (Natural History), London, 169 pp. Compton, S. G. & A. B. Ware, 1991. Ants disperse the elaiosome-bearing eggs of an African stick insect. — Psyche 98: 207-213. Froggatt, W. W., 1922. Description of a new phasma be- longing to the genus Extatosoma. — Proceedings of the Linnean Society of New South Wales 47: 344-345. Giinther, K., 1929. Die Phasmoiden der deutschen Kaiserin Augusta-Fluss-Expedition 1912-13. — Mitteilung aus dem Zoologisches Museum in Berlin 14: 597-746. Günther, K., 1931. Beiträge zur Systematik und Geschichte der Phasmoidenfauna Ozeaniens. — Mitteilung aus dem Zoologisches Museum in Berlin 17: 753-835. Gurney, A. B., 1947. Notes on some remarkable Australasian walkingsticks, including a synopsis of the ge- nus Extatosoma (Orthoptera: Phasmatidae). — Annals of the Entomological Society of America 40: 373-396. Harman, A., 1985. The genus Extatosoma Gray. — Phasmid Study Group Newsletter 25: 5. Hughes, L. & M. Westoby, 1992. Capitula on stick insect eggs and elaiosomes on seeds: convergent adaptations for burial by ants. — Functional Ecology 6: 642-648. Johns, R. J., 1982. Plant zonation. p. 309-330. — In: J. L. Gressitt (Ed.), Biogeography and Ecology of New Guinea. Junk, The Hague. Redtenbacher, J., 1908. p. 380-381. — Die Insektenfamilie der Phasmiden 3. Verlag Von Wilhelm Engelmann, Leipzig. Sellick, J. T. C., 1980. A study of the eggs of the insect or- der Phasmida with particular reference to the taxonomic value of egg structure in this group. — Unpublished Ph. D. thesis, University of London, England. Verdcourt, B., 1979. — A manual of New Guinea Legumes. Kristen Press Inc., Madang, PNG, 645 pp. Vickery, V. R., 1983. Catalogue of Australian stick insects (Phasmida, Phasmatodea, Phasmatoptera, or Cheleutoptera). — CSIRO Australia, Division of Entomology Technical Paper 20: 1-19. Wilson, S. K., 1991. Throwing your babies at the enemy. — GEO Australasia’s Geographical Magazine 12: 106-113. Received: 25 May 1993 Accepted: 30 August 1993 APPENDIX The following records of Extatosoma popa were used to plot fig. 16. The data have been taken from material exam- ined by the author, unless stated otherwise. IRIAN JAYA: 16, Hollandia, 3°10’S 140°E, 300-600 m, i.1937-8, W. Stüber (BMNH). PAPUA NEW GUINEA: Madang Prov.: 19, Finisterre Mts., Damanti, 1065 m, 2-11.x.1964, M. E. Bacchus (BMNH); 19, Finisterre Mts., Budemu, c. 1200 m, 15-24.x.1964, M. E. Bacchus (BMNH); Morobe Prov.: 22, Wantoat, 1200 m, Casuarina, x.1957, R. W. Paine (BMNH); BECCALONI: A new species of Extatosoma 29, Wau, golf course, 1150 m, ex Calliandra surinamensis Benth., 13.vii.1990, G. W. Beccaloni (BMNH); 19 (nymph), Wau, c. 1200 m, ex Hibiscus, 1.vii.1990, G. W. Beccaloni (BMNH); 1 ® (nymph), Wau, 1150 m, 24.vi.1974, Gewise Otaweto (WEIC); 16, Wau, 1180 m, 18.x.1974, A. D. Hart (WEIC); 16, Wau, 1150 m, coll’d on Diospyros fer- rea, 6.x11.1975 (WEIC); 12, Bulolo, v.1990 (BMNH); Northern Prov.: 12, Mt. Obree, 1.x.1921, R. Neill (BMNH). The following records were taken from Gurney (1947): Papua NEW GUINEA: Morobe Prov.: Sattelberg, vic. Finschafen; Northern Prov.: Buna. 125 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 (FIORENZA. XX INTERNATIONAL CONGRESS OF ENTOMOLOGY Florence (Italy) August 25 - 31, 1996 Palazzo dei Congressi First Circular Scientific Program GENERAL STRUCTURE — Opening Plenary Lecture - Plenary Symposia — Sections Organized by Selected Convenors - Special Symposia Selected by Section Convenors - Closing Plenary Lecture. — Audio-visual Sessions — Poster Sessions - Special Interest Groups - Workshops — Exhibitions THE SCIENTIFIC PROGRAM WILL BE ARRANGED IN THE FOLLOWING SECTIONS 1) Systematics and Phylogeny 2) Zoogeography 3) Morphology 4) Reproduction and Development 5) Cell Biology, Physiology and Biochemistry 6) Insect Neurosciences 7) Insect Immunity 8) Genetics and Evolutionary Entomology 9) Insect Molecular Biology and Genetic Engineering 10) Ecology and Population Dynamics 11) Special Environments Entomology 12) Ethology 13) Social Insects 14) Apidology and Sericulture 15) Agricultural Entomology 16) Forest Entomology 17) Tropical Entomology 18) Urban and Stored Products Entomology 19) Ecology of Pesticides, Resistance and Toxicology 20) Entomophagous Insects and Biological Control 21) General and Applied Insect Pathology 22) Integrated Pest Management 23) Medical and Veterinary Entomology 24) Biodiversity and Conservation 25) History of Entomology 26) Entomology for the Third Millenium - Critical Issues BE UE Gt GE BE de GE GE LE GE AE LE GE EE LE AE Gt WE GE e at dè 124 DK) D CONGRESS PRESIDENT B. Baccetti SECRETARY GENERAL A. Vigna Taglianti NATIONAL ORGANIZING COMMITTEE B. Baccetti E. Tremblay M. Cobolli A. Vigna Taglianti C. Conci R. Zocchi F. Frilli SCIENTIFIC PROGRAM COMMITTEE E. Tremblay (Pres.) F. Le Moli A. Arzone C. Malva F. Bin L. Masutti G. Briolini A. Minelli L. Bullini S. Ruffo R. Cavalloro V. Sbordoni M. Coluzzi S. Turillazzi R. Dallai G. Viggiani G. Domenichini Organizing Secretariat O.I.C. Via A. La Marmora, 24 50121 Florence (Italy) Tel. ++39-55.5000631 Fax ++39-55-5001912 Entomologists wishing to propose sectional symposia, workshops or special interest group meetings should return the enclosed intent form or write to the Congress Organizing Secretariat O.I.C. with details. The proposals are expected to reach the Congress Secretariat by the end of 1993. see page 288 THOMAS W. DONNELLY Binghamton, N. Y., U. S. A. TWO NEW GENERA OF ISOSTICTID DAMSELFLIES FROM NEW BRITAIN, BOUGAINVILLE, AND THE SOLOMON ISLANDS (ODONATA: ZYGOPTERA) Donnelly, T. W., 1993. Two new genera of isostictid damselflies from New Britain, Bougainville, and the Solomon Islands (Odonata: Zygoptera). — Tijdschrift voor Entomologie 136: 125-132, figs. 1-23, tab. 1. [ISSN 0040-7496]. Published 10 December 1993. Titanosticta and Cnemisticta are new genera of the Isostictidae described from the eastern Papuan - Melanesian region. New species Titanosticta macrogaster (New Britain), Cnemisticta angustilobata (New Britain), and C. latilobata (Bougainville, Gizo, Malaita, and Guadalcanal) are also described. The two genera bear no particularly close relationship with any of the Papuan (Selysioneura, Tanymecosticta), New Caledonia (Jsosticta), or Australian (seven genera) taxa. Dr. Thomas W. Donnelly, 2091 Partridge Lane, Binghamton NY 13903 USA. Key words. — Odonata, New Guinea, Solomon Islands, new genera, new species. The damselfly family Isostictidae is reported from the Australian subcontinent, the Papuan region (ex- tending westward to Tanimbar and Halmahera), and the island of New Caledonia, where the type genus occurs. In Australia, Watson et al. (1991) recognize seven genera. Tanymecosticta is widespread on New Guinea, has a single species in New Britain and an- other on Woodlark on the east, and occurs on Misool and Tanimbar on the west. Selysioneura is widespread on New Guinea, and occurs on Misool, Halmahera, and Morotai on the west, and Woodlark, D’Entrecasteaux, and the Louisiade Archipelago on the east. I herein describe two new Papuan genera: Titanosticta and Cnemisticta, which occur in New Britain and the Solomon Islands. The new genera overlap only slightly in range with either of the two Papuan genera: thus, they fill a geographic gap in the range of the family. The Isostictidae had long been considered a sub- family of the Protoneuridae, but has recently been separated into a separate family, largely on the basis of larval characters (Lieftinck 1975). Adults resemble the protoneurids, and differ from most otherwise similar Oriental Coenagrionoid families by their nar- row wings, with the anal portion highly reduced. They differ from the Protoneuridae by their absence of bright colours (protoneurid adults are almost in- variably marked with bright colours, especially in the males), their generally longer CuP, and larvae (as far as known) with saccoid, rather than flat, caudal la- mellae with subapical constriction. Some genera of Oriental protoneurids have a very short anal vein; this vein is always totally lacking in isostictids. Another character of most genera of the isostictids is a promi- nently and elaborately developed hind lobe of the prothorax; Selysioneura and one species of Cnemisticta are Conspicuous exceptions. Fraser (1955) discussed this group (which he con- sidered a subfamily of the Protoneuridae) and sug- gested a relationship with the platycnemidids. In common with the Protoneuridae, but conspicuously different from the Platycnemididae, the Isostictidae have short legs with short leg spines. However Cnemisticta has a broader wing which is abruptly nar- rowed distal to the pterostigma and a relatively long CuP, both of which are reminiscent of the Platycnemididae. The larvae of the Platycnemididae have highly variable caudal lamellae and do not seem to form a single coherent group. In the absence of larval forms for most genera it is not profitable at this time to suggest relationships within the family. The two new genera do not seem to have an especially close affinity with any other Papuan or Australian genus. Cremisticta is notably distinct within the family because of its broader wing with abrupt terminal narrowing, and long CuP. Abbreviations for collections in which type materi- al is located: UZMC — Universitetets Zoologiske Museum, Copenhagen; RMNH — Rijksmuseum van Natuurlijke Historie, Leiden; TD — Collection of T. Donnelly, Binghamton. 125 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-2. Venation of hind wings. — 1, Cnemisticta an- gustilobata, gen. n., sp. n. 2, Titanosticta macrogaster, gen. n., sp. n. Titanosticta gen. n. (figs. 2, 3-7) This is the largest damselfly in the family Isostictidae and one of the longest Zygoptera in the Oriental region. It is distinguished by the short CuP, which is two cells long (rarely one). Also, Ac is just proximal to Ax1 in the fore wing, and slightly distal in the hind wing; the arculus is distal to the Ax2 of both wings; IR2 arises at the 7-8th Px of the fore wing and at 6 or 7th (rarely 5th) in the hind wing; R4+5 arises distal to the subnodus. The pterostigma is rhombic and longer than broad. The wing is nar- rowed gradually distal to the pterostigma. The inferi- or appendage of the sole species is much larger than the superior. In common with other genera of the family, the legs are short with short spines, and the body colours are dull. Type species: T. macrogaster sp. n.; the only known locality is New Britain. Etymology: ‘Titan’, from mythology, signifying very large size, and ‘-sticta’, the predominant suffix for genera in this family. In addition to its very large size, Titanosticta is dis- tinguished from all other isostictid genera by the fol- lowing combination of characters: inferior appendage much larger than the superior; CuP two cells long, vein R4+5 arising distal to subnodus, very small tarsal claws, and Ac close to Axl in the fore wing and slight- ly distal to Axl in the hind wing. Titanosticta macrogaster sp.n. (figs. 2, 3-7) Type material. — Holotype d : Papua New Guinea. New Britain. Yalom, 1000 m, 20 May 1962. Noona Dan Exped. (UZMC). Allotype 9: Papua New Guinea. New Britain. Metelen River, 20 mi. S.E. of Ruango, July 1970, V. Jindrich (RMNH). Paratypes: 2 d same locality as holotype, 19-20 May 1970 (RMNH, TD); 18 Papua New Guinea. New Britain. Metelen River, 20 mi. S.E. of Ruango, July, 1970, 126 V. Jindrich (rp). Additional material. — An additional 2 in the Donnelly collection, with the same data as Metelen River paratype, is not designated a paratype because it lacks both a head and the terminal segments of the abdomen. Male holotype. — This is an obscurely marked spe- cies varying from pale gray brown to dark, shining brown. Head: Labium pale with rounded triangular central excavation about 2/5 the length of the segment. Face mainly pale; labrum shining black with thin apical yellow rim and pale central, basal spot; postclypeus pale with prominent black anterior and lateral mar- gins; frons pale with small black spots anterior and posterior to antennae and irregular black central stripe enclosing front of lateral ocelli and median ocellus; pedicel about 1.5 times the length of the scape, both pale. Prothorax: Pale, small paired central black spots on hind lobe and very small black marks on remainder pronotum; hind lobe sharply recumbent, broadly T- shaped, with a basal constriction. Pterothorax: Mesostigmal laminae small, slightly tapered apically, with rounded tips; pterothorax gray- ish green; mesepisternum with stripe on dorsal carina occupying 1/6 of sclerite; very thin dark line on ante- humeral suture; 2nd lateral suture with small dark spot on caudal sixth. Legs: Yellow with contrasting dark spines: 4 on outer row of hind femur, 5 on hind tibia. Wings: Venation described in generic diagnosis; venation brown, rhombic, pale red brown. Abdomen: Pale, darker at tip; 1 and 2 greenish gray; 3 to 8 yellow, darkening apically; 9 and 10 shin- ing dark brown. Superior appendage dark, shorter than 10, constricted apically to terminate in rounded, cylindrical tip, with low, rounded basal-ventral prom- inence. Inferior appendage 1.5 times length of super- ior, forcipate, laterally flattened, expanded abruptly in caudal half, excavate on dorsal-apical corner to DonneLLy: /sostictid damselflies Table 1. Characters of genera of Isostictidae. — Explanation: CuP: Number of cells length; both wings considered. Inferior appendage: of male, whether large or smaller than superior appendage. Hind lobe of prothorax, male: whether prominent (flared, with horns, or with other processes) or low. Ac or Axl: which crossvein is in more distal position in hind wing. Tarsal claws: size compared with other coenagrionoids. R4+5: origin of vein with respect to nodus. Genus Range length of CuP, male hind lobe tarsal R4+5 hw: Ac or Axl cells inferior prothorax claws to nodus distal appendages male Isosticta New Caledonia 1-6 equal prominent normal distal, Axl; some spp. proximal, nearly aligned or aligned Selysioneura New Guinea, 0-1 smaller low very small distal Ax (nearly Misool, aligned in Halmahera, some spp) Morotai, Woodlark, D’Entrecasteau, Louisiade Arch. Tanymecosticta New Guinea, 1 slightly prominent small distal Ax New Britain, smaller Woodlark, Misool, Tanimbar Titanosticta New Britain 2 (1) larger prominent very small distal Ax Cnemisticta New Britain, (7)8-10(11) larger prominent normal proximal Ac Bougainville, or low Solomon I. Rhadinosticta Australia 1 larger prominent normal proximal Axl Oristicta Australia 1 smaller prominent very small distal Ac Neosticta Australia 5-6 smaller prominent normal proximal Ac (close) Labidiosticta Australia 0-2 smaller prominent very small distal Axl Eurysticta Australia 122 smaller prominent normal proximal Axl Lithosticta Australia 5-7 equal prominent normal proximal Axl Austrosticta Australia 4-6 larger prominent normal proximal Ac (close) form two rounded tips. Penis with terminal segment Y-shaped. Female allotype. — Similar to male, but face more pale: labrum yellow with obscure paired lateral spots; anteclypeus mottled brown; postclypeus pale with brown anterior and lateral margins, this dark colour extended posteriorly in center and at lateral extremes; top of head gray-green, black as follows: short trans- verse dashes anterior to median ocellus; jagged trans- verse stripe enveloping median ocellus extending pos- terior to lateral ocelli. Thorax: Hind lobe of prothorax quadrate, scarcely narrowed at base, the central portion of hind rim ex- cavate in the center with a medial, rounded knob. Colour of side of thorax greenish yellow. Basal seg- ments of abdomen as in male; 5 terminal abdominal segments lacking. Dimensions. — Holotype male: abdomen 60.5 mm, hind wing 32 mm. Allotype female, hind wings dissimilar, 29.5 and 31 mm. The dimensional range of the paratype series is as follows: two males have abdomens 46 and 63 mm, and hind wings 32-32.5 mm. The 2nd female has the hind wing 30 mm. Variation within paratype series. — The three males have notably different abdomen lengths. The Metelen River male has the abdomen (46 mm) more than a centimeter shorter than the Yalom males (60.5 and 63 mm), but the hind wing of this specimen (29 mm) is only slightly shorter than the other two (32 mm). There are only trivial differences among the three males in colour pattern. One male has a broad- er black stripe on mesepisternum (4 rather than % the width of the sclerite). The females, which are both re- grettably incomplete, appear essentially identical. Etymology. — The name, a noun in apposition, re- fers to the very long abdomen. Remarks. — The difference in abdomen length of the males (which have very similar hind wing lengths) violates the normally firm linear allometric relation- ship (in this case the ratio between abdomen and hind wing) among Zygoptera. The constancy of this ratio can be attributed to the necessity of maintaining a uniform dimensional ratio to enable the insect to fly vigorously. There are similar disparities in this di- 127 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 3-7. Titanosticta macrogaster, gen. n., sp. n. — 3, male hind lobe of prothorax, dorsal view. 4, female hind lobe of pro- thorax, dorsal view. 5, male abdominal appendages, dorsal - lateral - apical view. 6, male abdominal appendages, lateral view. 7, penis, ventral and lateral view. mensional ratio among New World Pseudostigmat- idae, which have extremely long abdomens and which fly much less vigorously than other Zygoptera. The violation of a constant ratio in the few specimens of Titanosticta raises the question as to whether the flight of this genus (which has not been recorded) might resemble the weak flight of the pseudostigmat- ids. Cnemisticta gen. n. (figs. 3, 8-23) This genus contains two species of large isostictids. It is immediately recognized by the length of CuP, which in the type species C. angustilobata is 8 to 10 cells long in the fore wing) and 8 (rarely 7) to 10 (rarely 11) in the hind wing. IR2 arises at the 7-8th Px in the fore wing (rarely 9th) and 5 to 6th in the hind wing. In C. latilobata IR2 arises at the 5-6th (rarely 7th) Px in the fore wing and 4-5th Px in the hind wing; CuP is 8-10 cells long in the fore wing and 7-9 in the hind wing. In both species vein Ac arises just distal to the Ax1 of the fore wing and %o the dis- tance between the Ist and Ax2 in the hind wing; the arculus is distal to the Ax2 in both wings; R4+5 arises proximal to the subnodus in both wings. The pteros- tigma is rhombic and as wide as long. The tarsal claws are of normal length and the inferior abdominal ap- 128 pendage of the male is much larger than the superior. The two species range from New Britain to Bougain- ville, Gizo, Malaita, and Guadalcanal. Type species. — C. angustilobata sp. n., from New Britain. Etymology. — ‘Cnem-’ from the slight resemblance to some platycnemidids. This genus is distinguished from all other isostic- tids by its much longer CuP and abrupt narrowing of the wing beyond the pterostigma. The two species vary from each other in the extent of development of the hind lobe of the prothorax. The species C. latilob- ata has a very prominent hind lobe, similar to that of T. macrogaster, and to most of the other members of the family. The species C. angustilobata has a highly reduced hind lobe, in common with the most wide- spread Papuan isostictid, Selysioneura. Cnemisticta angustilobata sp. n. (figs. 1, 8-15) Type material. — Holotype d : Papua New Guinea. New Britain: Yalom, 1000 m, 18 May 1962, Noona Dan Exped. _(UZMC). Allotype 9: same data as holotype, except coll. 19 May 1962 (UZMC). Paratypes: 3d 19 same data as holotype except coll. 18-19 May 1962 (RMNH); Papua New Guinea. New Britain. Metelen River, 20 mi. S.E. of Ruango, July 1970, V. Jindrich (rp); 1 & Papua New Guinea. New Britain: Gazelle Pen., upper Warango, 250-600 m, 28-30 Nov 1962, J. Sedlacek (RMNH). Additional material. — 4 additional d collected by Sedlacek (RMNH) along with last named paratype are not designated as paratypes because they lack the terminal seg- ments of the abdomen. Description Male holotype. — This is a dark, obscurely marked damselfly. Head: labium pale with rounded excavation in ap- ical 4; frons rounded; face and top of head black; la- brum black with pale, obscurely defined central stripe; genae pale, anteclypeus obscurely pale; pedicel slightly longer than scape, both dark; rear of head pale. Prothorax: Pale laterally, mottled dark on dorsum, with large tringular paired dark spots on median lobes; apodyne extended posteriorly as commissure nearly to hind lobe; hind lobe small and low, round- ed triangular with central part slightly thickened. Pterothorax: Mesostigmal laminae simple, parallel sided, rounded, slightly elevated lateral tip; mesepi- sternum black, mesepimeron dark with pale on mesal % (adjacent to antehumeral suture) and cephalic %; mesinfraepisternum pale on outside, central half dark; the dark colour of the mesepisternum essential- ly extensive to metepisternum with only thin pale lines on the two lateral sutures; metepisternum pale with black spot on caudal %; metepipimeron and metastermum pale. Legs: Pale with thin dark dorsal lines on femora; spines brown, not contrasting with legs, 5 on hind fe- mur and hind tibia. Wings: Venation as for genus; IR2 arising at 7 (fore wing) and 5 (hind wing); CuP 9 cells long in fore wing and hind wing; pterostigma rounded, rhombic, as long as wide, pale reddish brown. Abdomen: Pale, dark on dorsum of 1 and 2, ob- scurely dark on dorsum of 3 to 7, with apical % dis- tinctly darker on 3 to 6; 7 uniform; 8 with obscurely dark dorsal mark in caudal #; 10 with obscure paired dark spots on caudal 4. Superior appendage shorter than 10, blunt, rounded, slightly expanded and branched in dorsal view, with mesal branch very short; with ventrally directed spine. Inferior append- age more than twice as long than superior, forcipate, laterally flattened, with caudal half expanded into prominent rounded keel on dorsal and apical por- tions. Penis with terminal segment Y-shaped. Allotype female. — Similar to male. Face pale; post- clypeus pale with dark lateral margins; top of head pale, about half dark, with an irregular dark mark centered on the ocelli, small dark spots anterior to an- tennae, and short transverse dark stripes posterior to lateral ocelli. Thorax: Hind lobe of prothorax similar DonneELLY: Isostictid damselflies to male but slightly larger, low triangular, with a stepped appearance. Colour of thorax: mesepister- num with pale stripe above antehumeral suture % the width and in the cephalic % of the sclerite; mesepime- ron with short central elongate dark mark; Metepisternum with small caudal dark mark. Abdomen: Dorsum of 8 to 10 pale; ovipositor dark, extending distinctly beyond 10 (cerci). Dimensions. — Holotype male: abdomen 44 mm, hind wing 29 mm. Allotype female: abdomen 45.5 mm. The dimensional range of the paratype series is as follows: 5 paratype males have the abdomens 43.5 to 47.5 mm; 8 males have the hind wing 26 to 29 mm. A paratype female has the abdomen 45.5 mm and the hind wing 32. Variation within paratype series. — The Gazelle Peninsula and Metelen River males are paler than the Yalom series; the dark colour of the side of the thorax ranges to a short central dark mark on mesepimeron and metepisternum almost totally pale. Etymology. — The name refers to the vary narrow hind lobe of the prothorax. Cnemisticta latilobata sp. n. (figs. 16-23) Type material. — Holotype d : Solomon Islands. Malaita I., Kware’a River and tributaries, 8 km E of Dala, 100m, 19 Feb 1987, T. Donnelly (RMNH); Allotype 2 Solomon Islands. Guadalcanal I., Honiara, Rove Creek at Reservoir, 13-14 Feb 1987, T. Donnelly (RMNH). — Paratypes: 1d same data as holotype (TD), 1d, Papua New Guinea, Bougainville I., South Nasiooi, Agriculture Field Station, 19 May 1975, Howard R. Wimmer (RMNH). Additional material. — 16 Solomon Islands. Gizo I., low bush area in valley, 7,8 May 1975, H. Wimmer (specimen lacks terminal segments of abdomen) (RMNH), 1d Papua New Guinea. Bougainville I., Boku Bong, 5 June 1956, J. L. Gressitt (RMNH) (highly teneral specimen). Description Holotype male. — A pale, obscurely mottled, dam- selfly. Head: Labium white, with a quadrate apical exca- vation % of its length; frons rounded; face black, la- brum with thin yellow apical rim; genae brown with yellow mesal margin; pale pedicel slightly longer than dark scape; top of head black bordered at rear by yel- low; obscure pale spots lateral to lateral ocelli; rear of head yellow. Prothorax: pale, fore lobe with dark rim; medial lobes low, apodyne continued into a commissure nearly to hind lobe; hind lobe large, T-shaped, sharp- ly recumbent, basally constricted, and sharply notch in center of hind rim. 129 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 8 - 15. Cnemisticta angustilobata, gen. n., sp. n. — 8, male hind lobe of prothorax, dorsal view. 9, male hind lobe of pro- thorax and mesostigmal lamina, dorsal- lateral view. 10, female hind lobe of prothorax, dorsal view. 11, female hind lobe of prothorax and mesostigmal lamina, dorsal- lateral view. 12, male abdominal appendages, dorsal - lateral - apical view. 13, male abdominal appendages, lateral view. 14, ovipositor. 15, penis, ventral and lateral views. Pterothorax: Mesostigmal laminae parallel sided, flat, with rounded tips; colour of pterothorax pale; mesepisternum with dark stripe on dorsal carina oc- cupying % of sclerite; antehumeral suture with thin black line; 2nd lateral suture with tiny black spot on caudal %. Legs: Yellow, with 4 contrasting black spines (outer row) on hind femur, 5 on hind tibia. Wings: Venation as in generic diagnosis; IR2 arising at 6 (fore wing) and 5 (hind wing); CiP 8 (fore wing), 9 (hind wing) cells long. Abdomen: Pale laterally and darker dorsally; 3 and 4 dominantly pale dorsally, slightly darker in basal half and on apical 4; 5 obscure, darker on apical %; 6 slightly pale in basal %; 7-10 dark, with limited pale on sides, 8 and 9 with pale longitudinal dorsal lines on basal %, tapering caudally. Superior appendage brown, shorter than 10, rounded and expanded api- 130 cally in dorsal view, with rounded, upturned apices. Inferior apendage 3 times as long as superior, forci- pate, with blunt, laterally flattened, rounded tip and dorsal keel on expanded apical half, and with low, rounded, conical, dorsally directed spine in basal %. Penis with terminal segment Y-shaped. Allotype female. — Similar to male. Head: Face pale, paired dark spots anterior to lateral ocelli. Small rounded pale protuberances on rear of head. Thorax: Hind lobe of prothorax prominent, triangular, divid- ed into three nearly equally wide lobes by deep sulci; the central lobe emarginate, and the two lateral lobes low and tapering laterally to points. Colour of thorax: Mesepisternum pale, thin black line adjacent to pale dorsal carina; thin black line along caudal half of an- tehumeral suture; mesepimeron with an obscure cen- DonnELLy: /sostictid damselflies Figs. 16-23. Cnemisticta latilobata gen. n., sp. n. — 16, male hind lobe of prothorax, dorsal view. 17, male hind lobe of pro- thorax and mesostigmal lamina, dorsal-lateral view. 18, female hind lobe of prothorax, dorsal view. 19, female hind lobe of prothorax and mesostigmal lamina, dorsal- lateral view. 20, male abdominal appendages, dorsal-lateral-apical view. 21, male abdominal appendages, lateral view. 22, ovipositor. 23, penis, ventral and lateral views. tral dark mark deflected laterally in cephalic portion. Abdomen: 9 and 10 with prominent pale basal-later- al spots; ovipositor pale, extends to end of 10 (cerci). Dimensions. — Holotype male: abdomen 39 mm, hind wing 26 mm. Allotype female: abdomen 35.5 mm, hind wing 26.5 mm. The dimensional range of the paratype series is as follows: 2 paratype males have abdomens 37 and 38.5 mm; 3 paratype males have the hind wing 24 to 26 mm. The 2nd female speci- men is very teneral; the hind wing is 27 mm. Variation within paratype series. — The male from Bougainville has the pale colour of the top of the head more extensive than the holotype, extending anterior- ly nearly to the antennal bases. A very teneral female specimen has the hind lobe of the prothorax longer and apparently more Y-shaped than the allotype. However the apparent difference in shape could be due to twisting upon drying. Etymology. — The name refers to the wide and dec- orative hind lobe of the prothorax. Remarks. — The two species of Cnemisticta resem- ble each other in their peculiarly wide and apically abruptly narrowed wings, the long CuP, and the male abdominal appendages. The species angustilobata is slightly larger than /atilobata. The species differ most prominently in the hind lobe of the prothorax, which is prominently developed in /atilobata and reduced in angustilobata. The species latilobata is pale, but with a darker abdomen, and angustilobata is dark, but with a paler abdomen. The allotype female of latilobata has 131 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 a rounded prominence on the rear of the head; this is lacking in angustilobata. The female of angustilobata has a much longer ovipositor than /atilobata. The ve- nation of the two species (above) differs more than is appropriate for the slight dimensional differences between the two species. ACKNOWLEDGEMENTS I am indebted to the RMNH, Leiden, for the loan of specimens which M. A. Lieftinck had been studying at the time of his death. I am further very grateful to Vladimir Jindrich, at that time a graduate student in our department, for specimens of two of the new spe- cies collected during a geological trip to New Britain. Comments from Drs. Jan van Tol and Anthony Watson on a draft of the manuscript have been very helpful and are gratefully acknowledged; I am further 132 grateful to Dr. Watson for the loan of specimens of four Australian genera which enabled me to complete the table of generic characters. REFERENCES Fraser, F. C., 1955. A new isostictine dragonfly from Australia with some remarks on the subfamily Isostictinae nov. (Odon., Protoneuridae). — Entomologist’s Monthly Magazine 91: 227-230. Lieftinck, M. A., 1975. The dragonflies (Odonata) of New Caledonia and the Loyalty Islands. Part 1. Imagines. — Cahiers O. R. S. T. O. M., série Hydrobiologie 9(3): 127-166. Watson, J. A. L., G. Theischinger, and H. M. Abbey, 1991. The Australian dragonflies. — CSIRO, Canberra and Melbourne, 278 p. Received: 14 April 1993 Accepted: 14 June 1993 MATTI HAMALAINEN Department of Applied Zoology, University of Helsinki DESCRIPTION OF NEUROBASIS DAVIESI SP. N. FROM PALAWAN, WITH TAXONOMIC NOTES ON OTHER SRECIES OP DEE NCHINENSISGROUP (ODONATA, EAEOPTERYGIDAE) Hämäläinen, M. 1993. Description of Neurobasis daviesi sp.n. from Palawan, with taxonomic notes on other species of the N. chinensis group (Odonata, Calopterygidae). — Tijdschrift voor Entomologie 136: 133-136, figs 1-4. [ISSN 0040-7496]. Published 10 December 1993. Neurobasis daviesi sp. n. (holotype male, Brooke’s Point, S. Palawan) is described in both sexes. The new species is closest to the Bornean N. longipes Hagen, 1887, which is considered to be specifically distinct from N. chinensis (Linnaeus, 1758). A key for both sexes to the three recog- nized species of the N. chinensis group is presented. M. Hämäläinen, P. O. Box 53, FIN-02151 Espoo, Finland. Key words. — Neurobasis, Palawan; new species; key. As well pointed out by Lieftinck (1949, p. 13-15), the definition of the limits of each species in the ge- nus Neurobasis Selys, 1853 has always been difficult and subject to considerable personal interpretation. This applies well also to the present assessment of the taxonomic status of a new Neurobasis taxon from Palawan, work on which necessitated a reconsidera- tion of Lieftinck’s (1940, 1954) view of recognizing three subspecies in Neurobasis chinensis (Linnaeus, 1758), viz. chinensis, florida Hagen and longipes Hagen. In my opinion it is difficult to regard /ongipes and florida to have an equal status as ‘subspecies of chinensis. Whereas the Javanese florida comes quite near to the continental forms, the Bornean longipes differs markedly from them and represents a distinct good species. Neurobasis chinensis (Linnaeus, 1758) (figs. 1-2). N. chinensis is a widely distributed insect in South and South-East Asia. It is known to extend from NE Pakistan in the west to South China Sea in the east. In the western part of its range it spreads from NE Pakistan, Himachal Pradesh, Uttar Pradesh and Nepal in the north through whole India to Ceylon. In the eastern part its area extends from Fujien and Yunnan provinces in China southwards to the Malaya Peninsula, Sumatra (& Simalur and Nias) and Java. In males the breadth of the wings and the relative size of the metallic area in the hind wing and in fe- males the presence or absence as well as the size of pseudopterostigma and the size of the whitish nodal spots are subject to considerable variation within the vast range of the species. Documentation of this vari- ability and revealing its nature is worth of a thorough study. Results of this study would also be useful in decid- ing on the taxonomic status of the Javanese popula- tions, which indeed differ most from the topotypical Chinese populations. Lieftinck (1940, 1954) consid- ered the Javanese populations to represent a distinct subspecies - florida Hagen, 1854. He also pointed out the very confusing usage of the name florida in the early literature and concluded that the original de- scription of florida applies to the Javanese specimens only. However, the correct citation to the author of florida should be Hagen in Walker, 1853. Neurobasis longipes Hagen, 1887 stat. n. (fig. 3). This species is universally distributed in Borneo and it occurs in lowland streams in forested areas. The following brief comparison with chinensis ex- plains my decision to consider longipes as a distinct species. In longipes — the size is smaller and the whole appearance is ‘- slimmer’ — legs are proportionally longer in both sexes; espe- 133 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-4. Wings of males (not in scale). - 1, N. chinensis, North Thailand; 2, N. chinensis (ssp. florida), West Java; 3, N. lon- gipes, Borneo, E. Sabah; 4, N. daviesi sp. n., Palawan (paratype). cially the anterior femora are very long, in male as long and in female longer than the length of the 3rd abdominal segment (in chinensis much shorter) — in the male hind wing the opaque colouring is much more reduced at base (Fig. 3); in subcostal field starting at the level of ca. 10th antenodal and below at the level of apical end of quadrangle — in male the shape of hind wings is different (Fig. 3), the broadest point is situated usually beyond the mid- dle of its length; fore wing base with only 2 complete rows of cells between the lowest sector of IA and the wing border — female is differently coloured; metallic green colour on thorax has a strong coppery tint instead of emerald metallic green in chinensis. In abdomen the shining metallic green colour on dorsum is restricted to seg- ments 1 and 2, the rest being dark brownish; in chi- 134 nensis the dorsum of segments 3 and 4 is similarly shining metallic green as on segment 2; the metallic colour getting darker green more apicad — whitish nodal spots and pseudopterostigma are lacking in all wings of the female. The species was originally described by Hagen (1887) as ‘Abart’ of chinensis on the basis of a single male specimen from Mindai. Hagen’s contribution remained unnoticed by other authors, like Kirby (1890), de Selys Longchamps (1897) and Förster (1897), who treated the genus in detail. Lieftinck (1940), who had studied a series of specimens from different parts of Borneo, reintroduced the name /on- gipes to apply to all Bornean specimens and presented a complete list of early references on this taxon. Neurobasis daviesi sp. n. (fig. 4). Type material. — Holotype d: Philippines, Palawan, Brooke’s Point, (325 m), l.viii. 1980, Leg. R. B. Rodriquez; in Museum Leiden (RMNH). — Paratypes: 2d, locality and date as in the holotype (RMNH); 3d, Philippines, S. Palawan, Brooke’s Point, 1982, Leg. R. B. Rodriquez (RMNH); 198, 49, Philippines, Palawan, Matalangao River [between Roxas and Port Barton], 21.xii.1975, [collector data lost, probably R. B. Rodriquez] (coll. D. A. L. Davies, Cambridge, U.K. ; 6d, 29 in coll. M. Hämäläinen). Description Instead of a detailed description, I prefer to high- light the characters by comparison of the new species with N. longipes and N. chinensis. N. daviesi is a some- what larger insect than chinensis and very distinctly larger than longipes. Male. — Colouring of head resembles that of chi- nensis. However, the yellow colour on labrum is re- stricted to form two widely separated spots. Thorax brilliant metallic green as in the related species. Sides of thorax showing, however, a slight coppery hue. Legs long; anterior femora proportionally longer than in chinensis, but shorter than in longipes; shorter than the length of the 3rd abdominal segment. Abdomen metallic green with pale markings on sides of seg- ments 1-2 as in the other species. Wings. The shape of hind wing is closer to that in longipes than in chinensis (Fig. 4). Opaque metallic area in hind wing extends more basad than in /on- gipes, but base of hind wing distinctly less coloured than in chinensis. Hind wing border at base as broad- ly hyaline as in the Javanese specimens of chinensis (ssp. florida), but the line with the opaque patch much less distict. As in the other species the hind wing re- flects brilliant metallic green or blue colour according to angle of view; the apical third with violaceous re- flections. The intensity of the metallic reflections is less pronounced than in chinensis, thus the outer line of the metallic patch looks less sharp. Measurements. Fore wing 35.5-37 mm, hind wing 33.5-35 mm; abdomen 48-49 mm. Female. — Rather robust insect. Thorax less shining metallic green than in chinensis. The colouring of ab- domen resembles that of longipes, metallic green colouring being restricted to dorsum of segment 1 and the basal 4/5th of segment 2. Rest of the abdo- men brown, darker brown on dorsum of segments 6- 10. Dorsal carina narrowly yellow broadening to a typical band on segments 8-10. Legs very long, proportionally longer than in chi- nensis. Anterior femora as long as the 3rd abdominal segment. Hind femora extending (in enveloped speci- mens) beyond half of the 3rd abdominal segment as in longipes. HAMALAINEN: Neurobasis daviesi sp. n. Wings hyaline with yellowish tint; as in longipes, the hind wings only slightly more heavily tinted than the fore wings. As in longipes, no trace of pseudopte- rostigma or whitish nodal spots in fore and hind wings. Measurements. Fore wing 38.5-39 mm, hind wing 40-41 mm, abdomen 47.5-48.5 mm. Etymology. — The new species is named after Dr D. A. L. Davies (Cambridge, U.K.) to appreciate his diligent efforts to search for dragonfly rarities in dif- ferent corners of the globe. Discussion. — In spite of the striking size difference, daviesi seems to be nearer to longipes than to chinensis. This is apparent especially from the female characters: long legs, absence of nodal spots and pseudopterostig- ma in all wings and brownish colouring of abdomen. However, the colouring of thorax is slightly different. Males show more ‘intermediate’ characters between chinensis and longipes. I think that it is better to to treat N. daviesi as a dis- tinct species, rather than a subspecies of /ongipes. Both taxa have well defined separate ranges in different is- lands, although separated only by the narrow Balabac Strait. It is remarkable that M. daviesi is missing from the large dragonfly collection amassed by Roland A. Miiller and his coworkers from many localities (even from Matalangao River) in different parts of Palawan in 1985-1992. N. daviesi is either a very rare species or (most likely) its flying period is confined to the ad- vanced rainy season. The bulk of Miiller’s specimens have been collected at the beginning of the rainy sea- son in May-June or during the dry season in March. Key to the species of Neurobasis chinensis group Males 1. Hind wing broadly ‘hyaline at the base; opaque patch starts at the level of the apical end of quadrangle at the central part of the wing. In fore wing only two complete rows of cells between the lowest sector of IA and the wing border. Anterior femora as long as the length of the 3rd abdominal SESCNENRL: Has ee tone ear senses wale longipes — Hyaline area in hind wing base less extensive. In fore wing at least 3 rows of cells between the low- est sector of IA and the wing border. Anterior fe- mora shorter than the length of the 3rd abdomi- DAL SEP IMEN by us Lira ? 2. Athind wing base the whole subcostal and cubi- tal fields opaque and reflecting strong metallic colour (only 1-2 basal cells of cubital field hya- mes RUE PP LEE LP M eae chinensis — Several hyaline or subhyaline cells in subcostal 135 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Females 1. All wings with (more or less distinct) whitish no- dal spots. Abdominal segments 3-4 predominant- ly shining metallic green on dorsum .....chinensis — No whitish nodal spots present on wings. Ab- dominal segments 3-4 predominantly brownish OMdOrsumae A Lee A Aes ne 2 2. Anterior femora clearly longer than the length of the 3rd abdominal segment. Small species, hind fm lees nn en longipes — Anterior femora as long as the length of the 3rd abdominal segment. Large species, hind wing OVER SA sum: daviesi ACKNOWLEDGEMENTS I am intebted to Drs Jan van Tol (RMNH, Leiden) and Dr D. A. L. Davies (Cambridge, U.K.) for the loan of the necessary specimens and for valuable com- ments on an earlier version of the manuscript. My thanks are also due to Mr Matti Viitasaari and Mr Reino Tyynelä for the wing photographs. 136 REFERENCES Förster, F., 1897. Contributions a la faune odonatologique Indo-Australe. — Annales de la Societe Entomologique de Belgique 41: 204-211. Hagen, H., 1887.Ueber Neurobasis und Vestalis. — Abhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 37: 647-648. Kirby, W. F., 1890. A synonymic catalogue of Neuroptera Odonata or dragonflies. With an appendix of fossil spe- cies. — Gurney & Jackson, London. Lieftinck, M. A., 1940. Descriptions and records of South- east asiatic Odonata (II). — Treubia 17: 337-390. Lieftinck, M. A., 1949. The dragonflies (Odonata) of New Guinea and neighbouring islands. Part VII. Results of the Third Archbold expedition 1938-1939 and of the Le Roux Expedition 1939 to Netherlands New Guinea (II. Zygoptera). — Nova Guinea (N.S.) 5: 1-271. Lieftinck, M. A., 1954. Handlist of Malaysian Odonata. A catalogue of the dragonflies of the Malay peninsula, Sumatra, Java and Borneo, including the adjacent small islands. — Treubia (Suppl.) 22: i-xiii + 1-202. Selys Longchamps, E. de, 1897. Causeries Odonatologiques No 10. — Annales de la Societe Entomologique de Belgique 41: 427-432. Received 1 September 1993 Revised version accepted 14 October 1993 CHRISTOPH L. HAUSER Museum Alexander Koenig, Bonn AeA NO TATED GHECKLIST OF THESPECIES OF THE PARNASSIINAE (LEPIDOPTERA: PAPILIONIDAE) Hauser, C. L., 1993. An annotated checklist of the species of the Parnassiinae (Lepidoptera: Papilionidae). — Tijdschrift voor Entomologie 136: 137-146. [ISSN 0040-7496]. Published 10 December 1993. A total of 76 binominal taxa including 22 cases of questionable species status are listed for the Parnassiinae, a subfamily comprising eight genera of predominately palaeartic swallowtail but- terflies. The taxonomic history of all doubtful species is briefly reviewed and possible affinites of these taxa are indicated. The majority of taxa contributing to a considerable increase in spe- cies number compared to previous accounts represent changes in status rather than genuine dis- coveries of new taxa. This is seen in relation to a change of attitude in taxonomic practise. C. L. Hauser, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. Key words. - Papilionidae, Parnasiinnae, checklist, species number. The Parnassiinae represent one of the three cur- rently recognized subfamilies of the Papilionidae, the ‘Swallowtail Butterflies’. Among the swallowtails, the ‘Apollo Butterflies’ in particular have for long been highly sought after by butterfly collectors because of their great aesthetic appeal and considerable intraspe- cific variation in wing pattern. This specialized inter- est has led in the past to voluminous works devoted to catalog and illustrate all the many named species- group taxa and infraspecific forms of the Parnassiinae (e.g., Austaut 1889, Verity 1905-1911, Stichel 1907a, 1907b, Rothschild 1918a, 1918b, Bryk 1934, 1935, Eisner 1974, 1976). In contrast, comparatively little attention has hitherto been paid to species-level taxonomy, and to morphological or ecological pecu- larities of the group. The subfamily is largely restricted to the palaearctic region with three species occuring in western North America, and completely absent from tropical zones (Stichel 1907a, 1907b, Reinig 1937, Ackery 1975). In an evolutionary view, the Parnassiinae have always been regarded as a basic stock of the Papilionidae from which all other higher taxa of the family arose (e.g., Spuler 1892, Reuter 1897, Ford 1944, Munroe 1961). In phylogenetic terms, the Parnassiinae are considered accordingly as the sistergroup of the Papilioninae, which comprise all remaining swallow- tails, with the exception of one monotypic genus (Hancock 1983, Miller 1987, Hauser 1993). Since the last comprehensive review of the Parnassiinae by Ackery (1975) several species have been described as new, mostly in the genus Parnassius (e.g., Koiwaya 1987, Korshunov 1988, Weiss & Michel 1989, Watanabe 1990). Furthermore, a num- ber of formerly recognized species has been shown to consist of two or more closely related, but reproduc- tively well isolated species, often on the basis of accu- mulated knowledge on their early stages and biology (e.g., Larsen 1973, de Freina 1985, Kreuzberg 1985). It seems therefore appropriate to provide an updated species checklist for this group. This list is also meant as a preliminary basis for a much needed phylogenet- ic analysis of the Parnassiinae at the species level, with the intention to point out all cases of presently prob- lematic species delimitations. CHECKLIST The present list can only be regarded as a first step towards a complete revision of the Parnassiinae even at species level. For this list, a ‘splitter’ approach was followed in many cases, in that geographically well se- parated and morphologically distinct taxa are rather given species status than treated as conspecific with their geographic vicariants. None of the many availa- ble infraspecific names appear in this list, but a num- ber of former subspecies have been elevated here to species rank following suggestions elsewhere in the li- terature. Similarly, all recently described species in the genus Parnassius are retained here in binominal combination. However, all closely related, but strictly allopatric species-group taxa are indicated in the pre- sent list by indentation and omitting the generic na- me. Under a more rigorous application of the biolo- gical species concept, most of these cases might well turn out as conspecific, and could be treated as such 157 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 for purely classificatory purposes. The Parnassiinae are seen here to comprise the eight genera listed below following recent classifica- tions of the Papilionidae (Munroe 1961, Ackery 1975, Hancock 1983, Miller 1987), despite the fact that the subfamily as currently defined might not rep- resent a monophyletic group in phylogenetic terms (Hiura 1980, Hauser 1990, 1993). All taxa are listed fully with author and year, and their sequence follows current ideas about taxonomic and phylogenetic rela- tionships. Comments on individual taxa are present- ed as a series of notes, which are numbered consecu- tively and appear at the end of the checklist. Subfamily Parnassiinae Swainson, 1840 Genus: Hypermnestra Ménétriés, 1848 O1 Hypermnestra helios (Nickerl, 1846) Genus: Parnassius Latreille, 1804 02 Subgenus: Parnassius Latreille, 1804 03 Parnassius apollo (Linnaeus, 1758) P. nomion Fischer De Waldheim, 1823 P. phoebus (Fabricius, 1793) sacerdos Stichel, 1906 04 ruckbeili Deckert, 1909 05 P. actius (Eversmann, 1843) P. jacquemontii Boisduval, 1836 P. tianschanicus Oberthür, 1879 P. epaphus Oberthür, 1879 P. bremeri Bremer, 1864 P. honrathi Staudinger, 1882 P. apollonius (Eversmann, 1847) Subgenus: Driopa Korshunov, 1988 06 P. mnemosyne (Linnaeus, 1759) P. stubbendorfii Ménétriés, 1849 hoenei Schweitzer, 1912 07 P. glacialis Butler, 1866 P. eversmanni Menetries, 1855 felderi Bremer, 1861 08 P. ariadne (Eversmann, 1843) 09 nordmanni [Ménétriés] in Siemaschko, 1850 09, 10 clodius Ménétriés, 1855 09 P. orleans Oberthür, 1890 Subgenus: Tadumia Moore, 1902 11 P. hardwickii Gray, 1831 P. hunnyngtoni Avinoff, 1916 12 P. acco Gray, 1853 przewalskii Alpheraky, 1887 13 138 baileyi South, 1913 P. szechenyii Frivaldszky, 1886 P. cephalus Grum-Grshimailo, 1891 pythia Roth, 1932 P. maharaja Avinoft, 1916 labeyriei Weiss & Michel, 1989 nosei Watanabe, 1990 P. schultei Weiss & Michel, 1989 Subgenus: Koramius Moore, 1902 P. delphius (Eversmann, 1843) maximinus Staudinger, 1891 P. cardinal Grum-Grshimailo, 1887 P. staudingeri Bang-Haas, 1882 stenosemus Honrath, 1890 P. stoliczkanus Felder & Felder, 1865 nandadevinensis Weiss, 1990 P. patricius Niepelt, 1911 priamus Bryk, 1913 hide Koiwaya, 1987 P. acdestis Grum-Grshimailo, 1891 Subgenus: Sachaia Korshunov, 1988 P. tenedius Eversmann, 1851 P. arcticus (Eisner, 1968) ammosovi (Korshunov, 1988) P. simo Gray, 1853 andrei Eisner, 1930 simonius Staudinger, 1889 P. boedromius Püngeler, 1901 Subgenus: Kailasius Moore, 1902 P. charltonius Gray, 1853 P. loxias Piingeler, 1901 autocrator Avinov, 1913 P. imperator Oberthür, 1883 P. inopinatus Kotzsch, 1940 Genus: Archon Hiibner, [1822] Archon apollinus (Herbst, 1798) A. apollinaris (Staudinger, 1892) Genus: Zerynthia Ochsenheimer, 1816 Zerynthia rumina (Linnaeus, 1758) Z. polyxena ([Denis & Schiffermüller], 1775) Genus: Allancastria Bryk, 1934 Allancastria cerisy (Godart, [1824]) cretica (Rebel, 1904) louristana (Le Cerf, 1908) 37 38 39 40 41 42 43 44 A. deyrollei (Oberthiir, 1869) 45 A. caucasica (Lederer, 1864) 46 Genus: Sericinus Westwood, 1851 Sericinus montela Gray, 1853 Genus: Bhutanitis Atkinson, 1873 47 Bhutanitis mansfieldi (Riley, 1939) 48 B. thaidina (Blanchard, 1871) B. lidderdalii Atkinson, 1873 B. ludlowi Gabriel, 1942 Genus: Luehdorfia Criiger, 1878 Luehdorfia puziloi (Erschoff, 1872) L. japonica Leech, 1889 L. chinensis Leech, 1893 49 L. longicaudata Lee, 1982 50 Notes 01.—The genus Hypermnestra has long been inclu- ded in the subfamily (Spuler 1892, Stichel 1907a, Bryk 1935, Ackery 1975), but recent phylogenetic in- vestigations suggest that it might not form part of the Parnassiinae as a monophyletic group (Hiura 1980, Hauser 1993). 02. — The splitting of the genus Parnassius into sev- eral genera had started by Moore (1901-1903), and was further elaborated by Bryk (1935), and, most re- cently, by Korshunov (1988, 1990). This is not re- garded here as useful taxonomic practice (Sokolov 1929, Munroe 1961, Ackery 1975), particularly as long as the genus is assumed to be monophyletic. Some of the generic names proposed, however, are re- tained here to designate presumably monophyletic species-groups, which for taxonomic purposes can be treated as subgenera (Munroe 1961; see notes 03, 06, 205056) 03. — The subgenus Parnassius or the so-called ‘apollo-group’ comprises a morphologically and eco- logically homogeneous group of species with larval hostplants restricted to Crassulaceae and Saxifra- gaceae (Bryk 1935, Ackery 1975, Kreuzberg 1987a). The proper allocation of many taxa from Central Asia, where several species of the group coexist, is still problematic and awaits further detailed studies. For the present list, with the exceptions noted below, the arrangement of species by Ackery (1975) is followed. 04. — The separation of the European taxa as a spe- cies distinct from P. phoebus is largely based on pecu- liar ecological requirements and morphological dif- ferences of the larvae, in addition to considerable geographic separation. P. sacerdos is restricted to hab- Hauser: Parnasstinae checklist itats along and near small streams of running water at elevations between 1500 and 2600 m in the European Alps, and its larvae exclusively feed on Saxifraga aizoides (Weidemann 1986, Descimon et al. 1989, Schweizerischer Bund fiir Naturschutz 1991). In contrast, habitat preferences are quite variable in P. phoebus in Northern Asia and Western North America where the species can occur from near sea level in Eastern Siberia and steppe zone habitats in the Altai Mountains (Kurentzov 1970, Mracek 1989) up to the nival zone above the tree line in the Rocky Mountains (Ferris 1976). In North America, larval hostplants of P. phoebus are various species of Sedum, whereas local species of Saxifraga, as well as S. aizoides are not accepted by larvae in captivity (Scott 1986: 189, Nardelli 1991). 05. — P. ruckbeili had been originally described as a subspecies of P. phoebus, and was treated subsequent- ly as such by most authors (Rothschild 1918b, Bryk 1935, Eisner 1976). The taxon, however, shows also affinities both in external appearence and male geni- talia to P. actius (Bryk & Eisner 1935, Hering 1935), and Eisner (1961: 180) finally placed it as a separate species. The known geographic range of P. ruckbeili is restricted to mountains North-East of the Tarim ba- sin completely separated from both P. phoebus, and P. actius, which leaves the species status doubtful. The name is often spelt as ‘riickbeil? or ‘rueckbeilî (e.g., Bryk 1935, Hering 1935, Eisner 1976), both of which represent unjustified emendations. 06. — The ‘mnemosyne-group’ was in the older liter- ature often referred to by the (sub-)generic name Doritis Fabricius, 1807 (Moore 1901-1903, Sokolov 1929, Bryk 1935, Munroe 1961), which, however, has P. apollo as its type-species, and thereby is invalid as an objective synonym of Parnassius Latreille, 1804 (Hemming 1967). The name Adoritis Koçak, 1989 is invalid as an objective synonym of Driopa Korshunov, 1988. 07. — P. hoenei from the Japanese Island of Hokkaido had originally been described as a subspe- cies of P. stubbendorfii and was subsequently treated as such by most authors (Bryk 1935, Eisner 1974, Ackery 1975). Recently, the taxon has been elevated to species rank by several Japanese authors (Fukuda et al. 1982, Kitahara 1990), but the conclusive evidence in support of such a view is not clear. 08. — P. felderi had been described as species and was accepted as such by most early authors (Stichel 1907a, Verity 1905-1911, Rothschild 1918b; but see Elwes 1886). Later on, the taxon was generally placed as a subspecies of P. eversmanni (Bryk 1922, Bang- Haas 1927, Eisner 1961, 1974, Kurentzov 1970, Ackery 1975), but some doubts about its proper stat- us remained (e.g., Bang-Haas 1927: 8, Eisner 1974: 90). The allopatric distribution of P. felderi and typi- 139 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 cal forms of P. eversmanni (see Iwamoto & Inomata 1988), and the occurrence of apparent intermediate taxa (Bryk 1935, Eisner 1961), give little support to maintain it as a distinct species. 09. — P. ariadne, P. nordmanni, and P. clodius have always been treated as distinct species despite their close morphological and ecological similarity, and their strictly allopatric distribution (Elwes 1886, Stichel 1907a, Bryk 1935, Eisner 1974, Ackery 1975). This practice is followed here, but it should be pointed out that the degree of morphological differ- ence between these taxa, for example, in male and fe- male genitalia (Hering 1932, Orr 1988), especially, between P. nordmanniand P. clodius is not very large. For consistency, then, comparable cases of geograph- ically vicariant taxa should also be classified as differ- ent species (e.g. 04). 10. — The author of P. nordmanni is sometimes been given as ‘Nordmann 1851’ (Ackery 1975, Hancock 1983), based on conclusions by Hemming (1934) that a description of the taxon by Ménétriés was never published. However, according to biblio- graphic studies by Nekrutenko & Kerzhner (1986) the original description of P. nordmanni by Ménétriés was indeed published in 1850 and thus takes prece- dence. 11.—In the subgenus 7adumia, the so-called ‘acco- group, Stichel (1907a), Bryk (1935), and Eisner (1976), had included P. acco, P. simo, P. tenedius, and related forms, but not P. szechenyi, P. cephalus, P. maharaja, and P. hardwickii, which were placed in different (sub-)genera. The separation of the szmo- and tenedius- groups from Tadumia on morphologi- cal grounds (Munroe 1961, Ackery 1975) is further supported by biological characters (Kreuzberg 1985, 1987a). A close relationship between P. acco, P. sze- chenyii, P. hardwickii, and P. cephalus with all asso- ciated taxa is supported by similarity in male genital- ia, sphragis structure, and larval hostplants (Hering 1932, Kreuzberg 1987a, Weiss & Michel 1989). 12. — P. hunnyngtoni had originally been described as a separate species on the basis of its extremely small size and different wing pattern (Avinoff 1916), but it was subsequently treated as conspecific with P. acco by Bang-Haas (1927), Eisner (1976), and Ackery (1975). In contrast to all other Parnassius, this species flies at high altitudes in early spring (Riley 1927, W.Eckweiler, pers. com.). Furthermore, its sympatric occurrence with P. acco (Riley 1927, Weiss & Michel 1989) leaves little doubt as to place it as a separate species. The species name is often found spelt as ‘han- nyngtont or ‘hanningtonî (Bryk 1935, Eisner 1976, Weiss 1992), both of which are unjustified emenda- tions. 13. — P. przewalskii had been described as a separ- ate species and was treated as such by Stichel (1907a), 140 Verity (1905-1911), Bryk (1935), and Munroe (1961), whereas it was considered as conspecific with P. acco by Hering (1937), Ackery (1975), Eisner (1976), Hancock (1983), and Weiss (1992). Recent discoveries have extended the known range of P. prze- walskii considerably (Huang & Murayama 1991, Schulte 1992), which now covers most of North- Western Tibet and Qinghai. Some of the new taxa described appear intermediate in wing pattern with P. acco, and leave further doubts about the species status of P. przewalskii (Weiss 1992). 14. — P. baileyi is another questionable species geo- graphically separated but close to P. acco, which had originally been described as a subspecies of P. acco from South-Eastern Tibet. Subsequently Bryk (1932a) described P. rothschildianus as a species separ- ate from P. acco from Western Sichuan, which he lat- er united with baileyi and another new taxon (Bryk 1932b). All these taxa were later on placed by Bryk (1935) as subspecies of P. przewalskii, whereas Bang- Haas (1927), Eisner (1976), and Ackery (1975) con- sidered them all conspecific with P. acco. New reports of ‘typical’ P. acco occurring in South-East Tibet (Nosé 1990), in addition to reported constant differ- ences in genitalia (Weiss 1992) suggest that possibly two species might occur in that region, and give rea- son to tentatively maintain P. baileyi as a species. However, a thorough taxonomic revision of the entire acco-group is certainly needed and should yield a more definite conclusion. 15. — The description of P. pythia is based on a sin- gle small female specimen from South-Western Tibet. The taxon was placed as a subspecies of P. cephalus by Bryk (1935), but it was maintained as a separate species in a list by Munroe (1961). The type locality of P. pythia was at the time of its discovery well separated from the then known range of P. ceph- alus, but geographically intermediate populations have since been found, which all appear conspecific with P. cephalus (Weiss 1992). 16. — Originally, P. maharaja had been described as a separate species (Avinoff 1916), but it was placed as a subspecies of P. cephalus by Bryk (1922), Eisner (1958, 1976), and Ackery (1975) despite constant differences in wing venation and genitalia between the two taxa (Hering 1932, Bryk 1935). The sympat- ric occurrence of newly discovered taxa closely related to P. maharaja with P. cephalus in Southern and South-Eastern Tibet (see below, 17 + 18) give further support to maintain P. maharaja as a separate species from P. cephalus (Bryk 1935, Munroe 1961, Weiss 1992): 17. — The recently discovered ?. labeyriei from Southern Tibet is morphologically very similar to P. maharaja from Ladakh (see Michel & Weiss 1989, Watanabe 1990), and probably just represents a geo- graphic vicariant form of the latter. However, pos- sibly intermediate records are currently missing from Western Tibet, which certainly is one of the areas that have never been adequately surveyed. Additional new discoveries indicate that this species is more widely distributed in Western China (Weiss 1991); see be- low, 18. 18. — P. nosei from E. Tibet is also very similar to P. maharaja, and has already been regarded as conspecif- ic with P. labeyriei (Weiss 1991). However, both P. labeyriei and P. nosei reportedly occur next to each other in South-Eastern Tibet (Nosé 1990), which is the reason to retain the latter provisionally here as a distinct species. Probably at least one of the two taxa will turn out to be conspecific with P. maharaja. 19. — P. schultei has been described from only three worn specimens from Southern Tibet (Weiss & Michel 1989). The genitalia of the male holotype ap- pear to differ considerably from P. cephalus, and also from P. maharaja and P. labeyriei, which gives reason to place it here as a distinct species. 20. — The status of most of the following species listed under the subgenus Koramius is still problemat- ic, and a comprehensive revision of the entire ‘delphi- us-group’ is much needed. Previous authors have gen- erally recognized only a single (Stichel 1907a, Verity 1905-1911, Hering 1932), two (Elwes 1886, Bang- Haas 1915, Rothschild 1918b), three (Bang-Haas 1927, Ackery 1975, Hancock 1983), four (Bryk 1935, Collins & Morris 1985), or at most five differ- ent species (Munroe 1961). However, detailed com- parative investigations on the ecology and genital morphology (Kreuzberg 1985) indicate that in sever- al mountain ranges of Central Asia (Tian-Shan, Alai, Pamirs) two to four distinct species of the ‘delphius- group’ coexist. The species recognized by Kreuzberg (1985, 1987a) are accepted for the present list (Weiss 1992), but it should be noted that an exact delimita- tion of these species and a proper allocation of the more than 50 taxa named formerly lumped under P. delphius still awaits to be accomplished. 21. — P. maximinus was recognized as a distinct species from P. delphius by Kreuzberg (1985) on the basis of different ecological requirements and differ- ences in larval colouration. The species seems restrict- ed to the Western Tian-Shan range and shows no sig- nificant overlap in its range with P. delphius (Weiss 1992), which leaves some doubt about its status as a separate species. 22. — P. cardinal had originally been described as a subspecies of P. delphius, and was first recognized as a separate species by Stshetkin (1979) based on its sym- patric occurrence with another de/phius-group taxon. Subsequent studies by Kreuzberg (1985) confirmed the coexistence of P. cardinal and P. staudingeri in Tadzhikistan, where P. cardinal occurs mainly at ele- Hauser: Parnassiinae checklist vations between 3000 and 3200 m. 23. — P. staudingeri was reinstated as a different species from P. delphius by Kreuzberg (1985) on the basis of differences in male and female genitalia as well as wing pattern, in addition to ecological differ- ences regarding habitat and larval hostplants. The species appears to co-exist in parts of its range with P. cardinal, but is strictly allopatric with P. delphius and P. stenosemus (Weiss 1992). 24. — P. stenosemus had been described as a subspe- cies of P. delphius and it was treated as such or as a subspecies of P. stoliczkanus by Stichel (1907a), Verity (1905-1911), Rothschild (1918b), Bang-Haas (1927), Eisner (1959a, 1976), and Ackery (1975). The taxon was first given species status by Moore (1901-1903), followed by Bryk (1935), Munroe (1961), and Weiss (1992), on the basis of constant differences in wing pattern and male genitalia with the sympatrically occurring P. stoliczkanus. However, the known range of P. stenosemus shows no overlap with P. staudingeri, and further studies are needed to confirm its species status. 25. — P. stoliczkanus had been described as a separ- ate species, but was later considered a subspecies of P. delphius by Stichel (1907a), Verity (1905-1911), Avinov (1913), Rothschild (1918b), Eisner (1959a, 1976) and Ackery (1975). It occurs sympatrically with P. stenosemus, and had been already given species status by Elwes (1886), Bang-Haas (1927), Bryk (1935), Munroe (1961), and Collins & Morris (1985). 26. — The original description of P. nandadevinen- sis is based on a single worn male specimen from Northern Uttar Pradesh, India. According to illustra- tions of the holotype and parts of its genitalia, it ap- pears intermediate between P. acdestis and P. stolicz- kanus, and could possibly represent an aberrant specimen of the latter species. 27. — Since its original description P. patricius had generally been accepted as a species separate from P. delphius (Bryk 1935, Munroe 1961, Ackery 1975, Hancock 1983, Weiss 1992; but see: Bang-Haas 1915), and recent morphological and ecological stud- ies further support this view (Kreuzberg 1985). 28. — P. priamus had been originally described as a subspecies of P. acdestis, and subsequently the taxon was considered as either belonging to this species or to P. delphius (Bang-Haas 1915, Bang-Haas 1927). Marked differences in the shape of the sphragis between P. priamus and P. acdestis led Bryk (1932b, 1935) to separate it from the latter species and to place it as a subspecies of P. patricius. Apparent differ- ences in male genitalia between P. priamus and P. pa- tricius could support the view of two different species (Weiss & Michel 1989), but both taxa occur allopat- rically (Weiss 1992). 141 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 29. — P. hide has been described recently from the Kunlun range, and was subsequently also found in Western Sichuan and South-Eastern Tibet (Koiwaya 1987, Nosé 1990, Shinkai 1990). The taxon appears not to differ much in male genitalia from P. priamus from Southern Tibet, and probably is conspecific with the latter or with P. patricius (see Weiss & Michel 1989: 12). 30. — The type species of Sachaia is P. tenedius by original designation (Korshunov 1988), and Kreiz- bergius Korshunoy, 1990 is available as a separate ge- neric name for P. simo and related taxa in this group. 31. — Originally, P. arcticus had been described as a northern subspecies of P. simo, a species which does not occur in Siberia, but Eisner (1969) later himself corrected this misjudgement, and placed the taxon as a subspecies of P. tenedius (Eisner 1976). In 1984, however, another Parnassius species was discovered in Central Siberia that on ecological and morphological grounds is clearly separate from P. tenedius, without much consideration, this species has been repeatedly referred to as P. arcticus (Mracek 1989, Weiss 1991), but proper identification must still await critical ex- amination of the holotype (see below, 32). 32. — As judged from the original description (Korshunov 1988), P. ammosovi appears not to be conspecific with P. tenedius, and probably represents the second Siberian species of that group (see above, 31); this taxon, however, would then be a junior synonym of P. arcticus depending on the identity of the holotype of the latter. 33. — P. andreji has been separated recently togeth- er with another six subordinate taxa from P. simo by Weiss (1991), based on differences in wing pattern and male genitalia; both character sets, however, have not yet been adequately documented. The fact that P. andrei is not found sympatrically with P. simo, which occurs over most of Tibet into South-West China, makes this separation furthermore questionable. 34. — P. simoniushad been described as a subspecies of P. simo and was treated as such by most authors (Bryk 1935, Munroe 1961, Ackery 1975). However, Avinov (1913) already had remarked on constant dif- ferences in male genitalia between P. simonius and P. simo (see also, Hering 1932), but did not separate both taxa at the species level. In a comparative study on the group, Kreuzberg (1985) elevated the taxon to species rank supported by constant differences in postabdominal morphology. As far as known, how- ever, P. simonius and P. simo occur allopatrically and the ranges of the two taxa show no overlap (Weiss 199): 35. — P. boedromius had originally been described as a separate species, but was since generally regarded as a subspecies of P. simo (Stichel 1907a, Bryk 1935, Ackery 1975), except for Avinov (1913) followed by 142 Rothschild (1918b), who had recognized that it oc- curs sympatrically with typical forms of P. simo. Recent studies by Kreuzberg (1985) supported Avinov s early conclusions through the confirmation of sympatric occurrences of the two species, and eco- logical and morphological differences. 36. — The type species of Kailasius is P. charltonius by original designation, and a separate generic name, Eukoramius Bryk, 1935, is available for P. imperator (Hemming 1967). 37. — P. loxias had been described as a species and has since been treated as such by most authors (Verity 1905-1911, Bryk 1935, Eisner 1959b, Munroe 1961, Ackery 1975, Kreuzberg 1985), except for Bang-Haas (1927) who regarded it as a subspecies of P. charlton- ius. The recent discovery of its biology confirms its status as a separate species from P. charltonius (Kreuz- berg & Pljushch 1989). 38. — P. autocrator was originally described as a subspecies of P. charltonius (Avinov 1913, Bryk 1935), but because of its unique wing pattern was lat- er on universally accepted as a separate species (Bang- Haas 1927, Eisner 1959b, Ackery 1975). In terms of genitalia and ecology, however, it shows a close affin- ity to P. loxias (Bryk 1937, Kreuzberg 1987b), from which it is geographically well separated (Weiss 1991). 39. — A. apollinaris had always been regarded as a subspecies of A. apollinus (Stichel 1907a, Bryk 1934, Igarashi 1979), until de Freina (1985) proposed the existence of two distinct but allopatric Archon species based on differences in genitalia and larval coloura- tion. The recent discovery of areas of sympatry of two distinct taxa in the province of Mardin in Southern Turkey further supports this view (Carbonell 1991). 40. — The generic name frequently used for Zerynthia in the older literature is Thais Fabricius, 1807, which is invalid as a junior primary homonym (Hemming 1967). The older replacement name Parnalius Rafinesque, 1815 has been suppressed for the principle of priority by the Plenary Powers of the International Commission on Zoological Nomen- clature (Opinion 1134). 41.—The genus Allancastria has been synonymized by some authors with Zerynthia (Ackery 1975, Igarashi 1979, Olivier 1991), but it is retained here as a separate genus on the basis of considerable differ- ences in genitalic structures (see: Bryk 1934, Nekrutenko 1990). 42. — The species name has often been incorrectly cited as ‘cerisyi Godart, 1822’ (e.g., Rothschild 1918a, Bryk 1934, Bernardi 1970, Eisner 1974) based on a reference that was never published (see Cowan 1970: 17). A. cerisy was long regarded as the only species in the genus (Stichel 1907b, Bryk 1934, Munroe 1961, Eisner 1974, Ackery 1975, Igarashi 1979), and the discovery of two sympatric Allan- castria species in Lebanon (Larsen 1973, 1975) has fi- nally led to the separation of several other species (see below, 43-46). A complete revision of all Allancastria taxa appears, therefore, much needed (Olivier 1991). 43. — The elevation of A. cretica endemic to the isle of Crete as a different species from A. cerisy was first proposed by Koçak (1981), and is presumably based on differences in wing shape and colouration only. As a number of phenotypically different populations of A. cerisy from other Eastern Mediterranean islands have also been named as subspecies (see Bryk 1934, Olivier 1991), the single treatment of this taxon as a species (e.g., Kudrna 1986: 169) must appear ques- tionable, and cannot be justified by a few distinguish- ing features and geographic separation alone. 44. — À. louristana had been described as a subspe- cies of A. cerisy, and it was first treated as a separate species by Kuhna (1977), followed by Blom & Eisner (1979), de Freina (1979), and Collins & Morris (1985). The distinguishing characters supporting such a view, however, have not been clearly stated. Areas of sympatry between A. louristana and either A. cerisy or A. deyrollei are presently not known, and the status of this taxon therefore remains questionable. 45. — Although A. deyrollei had originally been giv- en species rank when first described, it was recognized only recently as a separate species from A. cerisy based on differences in wing pattern, genital morphology and early stages (Bernardi 1970, Larsen 1973, 1975). The species occurs partly sympatrically with A. cerisy in Southern Turkey and Lebanon (Kuhna 1977, de Freina 1979, 1986). 46. — A. caucasica had been described as a subspe- cies of A. cerisy, and it was first regarded as a different species by Kuhna (1977) based on characters of wing pattern, colouration and genitalia. This view has since been largely accepted (de Freina 1979, Collins & Morris 1985, Nekrutenko 1990), but the exact de- limitation of the species and the allocation of various subspecific taxa to A. cerisy and A. caucasica, respec- tively, is not clear. 47. — At present, the genus Bhutanitis is held to comprise the four species listed here (Ackery 1975, Collins & Morris 1985). Recent field studies in South-West China, however, seem to indicate that, at least, three different Bhutanitis species exist in that area (Hou 1992). As so far only two species were known to occur in the region (Bryk 1934, Ackery 1975), the existence of an additional species in this genus is quite likely. Although several new taxa were named in Hou’s abstract (1992), these names cannot be accepted as available. In the older literature, the ge- nus is often referred to by the name Armandia Blanchard, 1871, which is invalid as junior homo- nym (Hemming 1967). Hauser: Parnassiinae checklist 48. — A separate genus, Yunnanopapilio Hiura, 1980, has been proposed for B. mansfieldi, but a ge- neric separation of this species from Bhutanitis seems not justified (Saigusa & Lee 1982), particularly when considering its biology and early stages (Lee 1986a, 1986b). 49.— L. chinensis had been described as a subspecies of L. japonica, and was treated as such by most au- thors (Verity 1905, Rothschild 1918a, Ackery 1975), except for Bryk (1934), and later on Eisner (1974) who placed it as a subspecies of L. puziloi. The taxon was first recognized as a species by Lee (1978) based on marked differences in the early stages, a view which has now become widely accepted (Collins & Morris 1985, Nosé 1990). 50. — L. longicaudata was recently discovered in Western China, and occurs partly sympatrically with L. chinensis. The differences seen in adult morpholo- gy and early stages seem to furthermore justify its stat- us as a separate species (Nosé 1990). CONCLUSIONS In the present list, 76 species taxa are included in the subfamily Parnassiinae. Although 22 of these bi- nominal taxa are indicated as questionable species some of which will eventually have to be synonymi- zed, the remaining number of recognized species will still exceed the fourty to fifty species listed in most previous taxonomic reviews of the subfamily. Generally, species numbers for the eight genera inclu- ded here in the Parnassiinae range from 39 species re- cognized by Stichel (1907a, 1907b) at the turn of the century, to 42 (Eisner 1974, 1976), 44 (Bryk 1934, 1935, Ackery 1975), 48 (Hancock, 1983), 49 (Munroe 1961), or, at most, 53 species listed by Collins & Morris (1985). This new increase in spe- cies number should come somewhat unexpected, par- ticularly, in a much collected and taxonomically well studied group of papilionid butterflies. Most of the taxa recently recognized.as distinct species, however, were already known at the beginning of this century, and only very few represent genuine discoveries (e.g., Parnassius schultei, P. arcticus, Luehdorfia longicauda- ta). Thus, the majority of recently discovered ‘new’ species in the Parnassiinae came about by a change of status rather than by the description of new taxa. This recent increase in species number can prob- ably be explained by a change of general concepts in systematics, which is well reflected by the different at- titudes of specialists dealing with the Parnassiinae at different times. In the first half of this century, most authors were primarily concerned with the study of (intraspecific) variation in wing pattern exclusively based on large collections of adult specimens (e.g., Austaut 1899, Verity 1905-1911, Bryk 1934, 1935). 143 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 This approach produced an immense number of in- fraspecific taxa named for the Parnassiinae (Bryk 1922, Eisner 1974, 1976), and it naturally obscured somewhat the question of delimitation of species. In recent times, more and more authors tried to rely for their taxonomic decisions on other morphological, biological and ecological characteristics, many of which usually exhibit larger differences at the species level (e.g., Larsen 1975, Lee 1978, Kreuzberg 1985). This shift away from a purely descriptive taxonomy relying on a single set of adult characters was partly driven by and coincided with the change from a typo- logical towards a biological species concept in system- atics (Mayr 1982). In the Parnassiinae, the long ap- plied taxonomic practise to record and name all forms of infraspecific variation in wing pattern still persists today (e.g., Schulte 1992, Weiss 1992), but is slowly giving way to a more biological aproach considering all possible characteristics and pecularities of the group. From this historical perspective it appears therefore understandable that, despite the consider- able taxonomic efforts already undertaken in the past, a comparatively high number of unresolved cases at species level in the Parnassiinae remains. The primary purpose of this paper is to direct attention to these cases and encourage further, more detailed studies. ACKNOWLEDGEMENTS Helpful comments and criticisms on a preliminary draft of this paper were received from Dr. W. Eckweiler, Frankfurt/Main, Prof. Dr. C. M. Naumann, Bonn, and Ju. Ju. Stshetkin, Taganrog, Russia. 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Neumann-Neudamm, Received: September 1993 Accepted: 16 September 1993 WILLEM HURKMANS Institute of Taxonomic Zoology (Zoological Museum), Amsterdam A MONOGRAPH OF MERODON (DIPTERA: SYRPHIDAE). PART 1 Hurkmans, W., 1993 A monograph of Merodon (Diptera: Syrphidae). Part 1. — Tijdschrift voor Entomologie 136: 147-234, figs. 1-106. [ISSN 0040-7496]. Published 10 December 1993. This first part of a revision of the genus Merodon deals with 61 species characterized by two syn- apomorphies, viz. arcuate abdominal pruinose bands and abdomina with tapering second ter- gite. Eleven monophyletic species groups are recognized, accounting for 51 species. Seven spe- cies are placed in an undefined set while three species are discussed separately. Descriptions of the species groups are provided. All species are described and their important characters figured. Of the 91 nominal taxa mentioned in the checklist, 61 are species, 2 subspecies, 6 varieties and 22 synonyms, 4 of which are new. Of these, the following 28 species and one subspecies are new to science: M. mariae, M. ottomanus, M. testaceoides, M. altinosus, M. hypochrysos, M. mar- ginicornis, M. sophron, M. ankylogaster, M. auronitens, M. oidipous, M. persicus, M. tangeren- sis, M. xanthipous, M. aberrans isperensis subsp. n., M. cupreus, M. lusitanicus, M. splendens., M. warnckei, M. kaloceros, M. vandergooti, M. bequaerti, M. elegans, M. lucasi, M. nitidi- frons, M. satdagensis, M. schachti, M. taniniensis, M. toscanus, M. aureotibia. Lectotypes are designated for 23 nominal taxa; three new combinations are introduced. An an- notated list of species excluded from Merodon is given. Azpeytia shirakii nom. n. is proposed for Merodon scutellaris Shiraki. Willem Hurkmans, c/o Institute of Taxonomic Zoology (Zoological Museum), Department of Entomology, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. Key words. — Diptera; Syrphidae; Merodon; new subdivision; new species; lectotypes; excluded species; nomen novum. CONTENTS Index suites lose CORAL) zend oa LOLA 235 Ie Cao teen ite 147 facere LEE alesh SRO EA 148 Nie tera cn de Sn 148 The genus Merodon is a moderately large group of The genus Merodon Meigen, 1803 ..................... 149 syrphidflies naturally occurring in the palaearctic and ROretheSpEcIEStreaTe di... 153 ethiopian faunal regions. As far as known the deve- SA Pnt Gye i nai ina 157 lopment of the larvae takes place in bulbs or rhizomes desuncusassemblape>— ron 157 of Liliaceae and Amaryllidaceae. The last comprehen- TEBE CROWD eee per > cdi ae lu 162 sive discussion of the genus, by Sack (1931), is outda- TAD § > li, Sa Sek nude a 169 ted, contains several errors, and deals with only about slzumpespionpiio- > rr 175 half the species currently known. The descriptions [TE OAD). —— — > 185 and discussions of Merodon species published so far OA CON SOUPE. ari a 186 largely disregard important male genital characters DLC TOO EC DIE io Lot (i. (i. 188 considered suitable for grouping purposes and species LIT NEO DR sen 189 recognition. ZOE RON ne hen re ne 191 This paper deals with eleven monophyletic species GEASS EUTOTIS) PCO) sn 193 groups defined by apomorphies and a remaining un- ÉGALE e or oS 194 defined set of species. All species discussed in this CROIENT 197 paper are grouped by two synapomorphies, viz. arcu- Remamino species Lin 203 ate pruinose bands on the abdominal tergites, and an Annotated list of species discussed ...................... 204 abdomen with tapering second tergite. Only the Annotated list of species excluded from males are used for grouping purposes, since their gen- broder boni levi lana 207 italia yield enough data for subdivision. The female Neknowledpementst-.:- orione 208 genitalia are rather uniform; moreover the ornamen- References: teste mens en 208 tation of the legs and specialized pubescence areas are 147 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 usually much less developed in the females than in the males. MATERIAL Material was studied from the following institutes and private collections (with acronyms used in this paper): BMNH, British Museum (Natural History), London, England; MNHN, Muséum National d’ Histoire Naturelle, Paris, France; LSF, Museo Zoologico della Specola, Florence, Italy; MCSN, Museo Civico della Storia Naturale, Milan, Italy; BSA, Benediktinerstift Admont, Admont, Austria; NHMW, Naturhistorisches Museum Wien, Vienna, Austria; ZMHB, Museum für Naturkunde, Von Humboldt Universität, Berlin, Germany; DEIC, Institut für Pflanzenschützforschung, Eberswalde, Germany; HLMD, Hessisches Landesmuseum, Darmstadt, Germany; MENS, Museum für Naturkunde, Stuttgart, Germany; LAU, Musée Zoologique, Lausanne, Switzerland; KBIN, Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussels, Belgium; MRAC, Museum voor Midden Afrika, Tervuren, Belgium; ZMUC, Zoological Museum, Copenhagen University, Copenhagen, Denmark; sAL, Zoological Institute of Salamanca University, Salamanca, Spain; IEE, Instituto Espanol de Entomologia, Madrid, Spain; ZMAS, Zoological Institute of the Russian Academy of Sciences, St. Petersburg, formerly Leningrad, Russia; TMA, Természettudomänyi Museum Allättara, Budapest, Hungary; BTM, Bakonyi Természettudo- mänyi Museum, Zirc, Hungary; LHW, Entomological Institute, Agriculture University, Wageningen, the Netherlands; RMNH, Rijksmuseum van Natuurlijke Historie, Leiden, the Netherlands; JLR, J. A. W. Lucas, Rotterdam, the Netherlands (the most impor- tant single source of material; material from several other, mainly private collections was made available by Mr Lucas as well); ZMAN, Institute of Taxonomic Zoology, Zoological Museum, University of Amster- dam, Amsterdam, the Netherlands; PC, S. J. Para- monov collection. Many types of species described by Paramonov were formerly preserved in his private collection. After Liepa (1969) this collection is consi- dered to have been lost. METHODS Preparation of specimens In order to study the male genitalia, specimens pri- or to examination were relaxed using dilute ethanol vapour at room temperature for 6 to 48 hours depen- ding on specimen size. After relaxation the genitalia were extracted with an insect pin with hooked tip. Isolation of the genitalia often proved necessary; in such cases the genitalia were stored in microvials con- 148 taining a 60%-40% mix of glycerol and ethanol. The genitalia were examined in the dry state since boiling or prolonged submersion in KOH led to maceration and damage to the pubescence. The pubescence found in many species could hardly be seen properly in wet state. We were confronted with the same pro- blem with genitalia already isolated and stored in Canada balsam. Genitalia already stored in microvials (in ethanol/glycerol mixture) were repeatedly washed with 70% ethanol and left to dry prior to examina- tion. In Merodon, the female genitalia have been of little use for taxonomic purposes since they are hardly scle- rotized. If dried they shrivel completely, and if stored in a liquid medium the various pubescences are diffi- cult to distinguish. Methods used in grouping The species have been grouped on the basis of apo- morphic characters. Characters were considered apo- morphic after a preliminary survey of the correspon- ding characters in other species groups and in the genus Eumerus, the presumed sister group of Merodon, on the ground of similarities in the genita- lia and the banding pattern on the abdomen of some species of Eumerus formerly placed in the genera Megatrigon Johnson and Amphoterus Bezzi (see also Hull 1949: 395). On the basis of this comparison a slightly oblique to parallel, straight banding pattern, comparatively simple male genitalia and a comparati- vely stout abdomen are therefore considered to be plesiomorphic. Terminology The terms used to describe the various structures are illustrated in figs. 1-4 (head), 5 (thorax), 6 (wing), 7 (leg) and 8-17 (male genitalia). The terms are main- ly derived from McAlpine (1981), but some terms from Speight (1987), Van der Goot (1981), Hull (1949), Sack (1931) and Metcalf (1921) have been used for elements occurring specifically in Syrphidae. A few terms are introduced here to describe structures in the male genitalia; they are illustrated in figs. 8-17. The vertex angle and ocellar angle, as used here in de- scribing the head, are illustrated in fig. 3. The ratio obtained by dividing the compound eye touchline length by the vertex height, measured from its angle to the border with the occiput, is cited as the tl-v ratio. This ratio is often a useful distinguishing character. The antennal ratio, illustrated in fig. 2, is also frequently used. Sternites and tergites are count- ed in Roman numerals and abbreviated S and T, re- spectively. Drawings and maps All drawings are original; they were made with the aid of various stereo microscopes with drawing tubes under 10-80 X magnification; all scale lines represent 1 mm. Distribution maps are provided for all species. On the distribution maps only those records are repre- sented which are based on examined material or on reliable references. References from literature for spe- cies easily confused with other species have not been included. Where many records for a species occur, single records are represented by small symbols. The large symbols represent five records in the same area. THE GENUS MERODON MEIGEN, 1803 Merodon Meigen, 1803: 274. — Type-species Syrphus clavipes Fabricius, 1781: 427 [designation of Westwood 1840: 137]. Lampetia Meigen, 1800: 34 [Suppressed by the I. C. Z. N., Opinion 678]. — Type-species Syrphus clavipes Fabricius, 1781: 427 [designation of Coquillett 1910: 557]. Exmerodon Becker, 1912: 604 (as subgenus of Merodon). — Type-species Merodon (Exmerodon) fulcratus Becker, 1912: 604 [original designation]. Taxonomic and nomenclatural history From 1781 to c. 1900 In 1800, Meigen established the genus Lampetia to accommodate Syrphus clavipes Fabricius, 1781 (Meigen 1800). In 1803 he introduced the new name Merodon for this genus, mentioning that Syrphus cur- vipes [error for clavipes| belongs in Merodon (Meigen 1803). The work of Meigen (1800) has been placed on the Official Index of Rejected and Invalid Works in 1963 (I. C. Z. N. Opinion 678); since then only the Meigen (1803) names should have been used. Several species described by Fabricius and Rossi prior to the description of Merodon were later incorporated in this genus by e.g. Fabricius (1805). At the start of the nineteenth century species now included in Merodon were described in several genera such as Eristalis Latreille, 1804, and Milesza Latreille, 1804. As, in the first half of the nineteenth century, the concept of the various syrphid genera crystallized, many new species were rightly described in Merodon. During this period Fabricius (1805), Meigen (1822- 1838), Macquart (1834, 1842) and Rondani (1843- 1868) described and redescribed many species. Several synonyms were created through interpreta- tion of colour forms as species. Walker (1849-1860) assigned many species to this genus, all but one incor- rectly. Loew (1862a-b, 1869) described several new species and also provided redescriptions for several known species. Strobl (1893-1909) described several new, and redescribed many known taxa. The authors mentioned so far almost never discussed the differ- ences between species, except when establishing a Hurkmans: Monograph of Merodon new taxon; they were not concerned with subdividing the genus or with discussing its position within the Syrphidae. The twentieth century The first author providing a comprehensive generic description for Merodon was Verrall (1901) who loca- ted this genus in the subfamily Eristalinae. Villeneuve (1909-1934) published several new species and recog- nized a number of synonyms within Merodon. Becker (1912) subdivided Merodon into two subgenera, the nominal subgenus Merodon and the new subgenus Exmerodon Becker, 1912. The latter subgenus at that time comprised only the type-species Merodon fulcra- tus Becker, 1912. Stackelberg (in Stackelberg & Richter 1968) included his new species dichopticus in the same subgenus, established to accommodate spe- cies with dichoptic eyes in the males. In the only mo- nograph on Merodon so far, by Sack (1913a), Merodon was not assigned to any subfamily although it was compared with several genera in the Eristalinae. Other publications by Sack (1913-1938) contain a wealth of information on Merodon, many descrip- tions of new species and a subdivision of the genus based on external similarities. Major contributions to the knowledge of Merodon were made by Paramonov (1924-1937). He divided the genus into species groups, not all of which were published. These groups, like those of Sack (1913a), were based on ex- ternal similarities. Moreover Paramonov critically re- viewed papers by Becker (1912) and Sack (1931), re- jecting some species and expressing his doubts about others. Other contributions were made by Gil Collado (1930), Curran (1939) and, more recently, Séguy (1961), who surveyed the species known from west- ern Europe. Van der Goot (1981) in his work on the north-west European Syrphidae provided a key for the species occurring in that area. The works mentioned so far did not make use of genital characters to identify the species. In Merodon many problems existed as to the separation of species, which easily could have been solved by superficial ex- amination of the genitalia. The first papers in which genitalia of Merodon are discussed and figured were by Glumac (1958) and Gaunitz (1969). Regrettably the drawings provided are insufficient, but they af- ford an insight into the great variety within the genus. There is a considerable amount of literature on one single species, the narcissus bulb fly Merodon equestris (Fabricius, 1794), which is a pest in flower bulb cul- tivation. This species probably received more atten- tion in terms of literature produced, than all other species together. This literature is mainly concerned with economical aspects, genetics and life cycle. Some of this literature is referred to under that species. No 149 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 other species of Merodon are of known economical importance. Diagnostic characters The genus Merodon is characterized by the follo- wing characters: (1) Radius 4+5 curves deeply into cell r5 (e.g. fig. 19 d); (2) the apical part of Media is strongly recurrent; its subapical portion is slightly si- nuate (e.g. figs. 6, 19 d); (3) dm-cu is weakly, smoothly curved, not sinuate (e.g. figs. 6, 19 d); (4) femora 3 have a concave lower face and apically bear a single projection at the distal end (e.g. figs. 20 a through e). These features are found in both sexes; the size and shape of the projection on femora 3 is rather variable, but this projection is always present although it is re- duced in some species. | Species of Merodon have sometimes been assigned to other genera. More often, species of Mallota Meigen, 1822, and Eumerus Meigen, 1822, are wrongly assigned to Merodon. The genus Azpeytia Walker, 1865, includes species that can be mistaken for Merodon. In Azpeytia the femora 3, which are less stout than in Merodon, bear two projections or series of spinules distally; the scutellum is much wider than in Merodon; R4+5 curves only weakly into cell r5; dm- cu is distinctly sinuate to kinked; moreover the oral margin is less projecting than in the majority of Merodon species (figs. 18 a, 19 b, 20 k, ]). In Eumerus the femora 3 have a convex lower face and bear two projections or spinule series as in Azpeytia, the 3rd antennal article is relatively larger and shorter in Eumerus than in most Merodon species; R4+5 does not or hardly curve into cell r5 while dm- cu is strongly kinked in Eumerus, in several Eumerus small appending veinlets of the dm-cu are found; these are not found in Merodon (figs. 18 c, 19 a, 20 f, g, h, j). In Mallota the apical part of vein M is not recurrent as in Merodon; moreover Mallota species have a dis- tinct facial knob, not found in Merodon (figs. 18 d, 19 c, 20 m, n). Platynochaetus Wiedemann, 1830, is rather similar in appearance to Merodon; the males may be distin- guished by their curious antennae, while both sexes show a facial knob; the position of the bands on the abdomen is different (anteriorly in Platynochaetus, in mid-region in Merodon). However, the wing venation and the shape of femora 3 are quite similar to that in Merodon (figs. 18 e, 19 e, 20 0, p). The larvae of Merodon may be separated from those of other genera by using the key of Heiss (1938), who figured the larva of Merodon equestris. As the larvae of most species are still unknown, identifi- 150 cation may be problematic. According to Maldonado Capriles & Berrios (1977) the puparium of Merodon can be confused with that of Copestylum species. Description of the imagines (fig. 23 d) In all respects species of Merodon conform to the criteria established for the Syrphidae by many au- thors, e.g. Verrall (1901), Lundbeck (1916), Sack (1931), Hull (1949) and Van der Goot (1981). Habitus: Stoutly built to moderately slender, rarely very slender syrphid flies. Body length 5-25 mm, wing span 6-35 mm. Head, thorax and abdomen about equally wide. Legs rather stout, the trochanters 3 and femora 3 in many species swollen and orna- mented in the males. In many species pruinose bands of microtrichiae are present on the thorax and abdo- men. The thoracic surface is black or dark brown, in several species metallic lustrous; the abdomen is black to brown, often with yellow to red lateral spots which may extend over all the abdomen. Wings mostly clear although infuscate wings rarely occur. A typical repre- sentative of the genus, M. distinctus Palma d is shown in fig. 23 d. Many species mentioned in this paper are superficially similar, especially those of the alagoezicus, avidus, crassifemoris, nigritarsis and van- dergooti groups. Head: In the majority of species the males are hol- optic while intermediate and dichoptic conditions oc- cur in males of several species; females always dichop- tic. Compound eyes dark brown to reddish, usually bearing moderately dense whitish pubescence. Antennae shorter than head except in a few species with lengthened third article; third article usually longest and widest, often conspicuous; arista inserted rather basally on dorsal margin. Coloration of anten- nae most variable. Face lacking knob; oral margin clear, often conspicuous, seldom weak; genae absent; facial surface little sculptured, dark, often metallic lustrous, often yellow to white pruinose and pubes- cent. Frons concolorous, often denser pubescent than face; in females lateral strips on frons often pruinose, leaving free a lustrous midstripe; species with an all lustrous or all pruinose frons occur. The pruinose frontal strips do not occur in dichoptic males. In males with incompletely touching eyes the narrow frontal strip is bald. Holoptic males show consider- able variety in length of the touchline of the eyes. The vertex is dark, in some species metallic lustrous, and often conspicuously dark pubescent in the ocellar re- gion. Occiput dark, pubescence concolorous, often white to yellow pruinose down sides (figs. 3-4, 18 b). Thorax: Slightly elongate, humeral calli well marked, dorsal suture clear laterally only; colour dark, often metallic lustrous on dorsum, scutellum, kate- pisternum, anepisternum and anepimeron. Pube- scence often conspicuous, sometimes strongly coloured; length and density most variable; species with very even, and with uneven pubescence occur. Pruinose bands, if present, are longitudinal, often co- alescent anteriorly; in some cases weak crossbands oc- cur along the dorsal suture. The scutellum is without specialized structures or specialized pubescence (fig. 5). Wings: The wing surface is clear to partially or strongly infuscate, in most species slightly, evenly troubled. All species show the vena spuria characteris- tic of the Syrphidae. Stigmal crossvein oblique, stig- ma absent; cell rl open anteriorly; R4+5 strongly curving into r5, dm-cu proximally sinuous, distally rather strongly recurrent; r-m reaches M1+2 at ap- proximately the middle of the anterior border of cell dm; no appending veinlets occur on R4+5 or dm-cu; the curves in CuAl and dm-cu are smooth. The colour of the veins ranges from dark yellow to black and is rather constant within species. The alula is in- conspicuous; the squama and antisquama (upper and lower calypters) are pale and bear a fringe of pale, yel- low, even pubescence. The halteres usually are well clear of the thoracic pubescence; the stalk is yellow, the knob usually pale, sometimes dark (fig. 6, 23 d). Legs: Legs 1 and 2 moderately robust. Leg 3 very robust, with specialized structures: a single, usually triangular projection on the femora 3 at the distal end in all Merodon. Various ridges and spines occur on the trochanters 3, femora 3, tibiae 3 and metatarsi 3 in the males, seldom so in the females, of many species. In several species the femora, the metatarsi, or both, may be swollen or modified strongly in the males, weakly in females. The colour is most variable, species with all orange and with all black legs occur. In many species the dark legs bear short, recumbent, golden pubescence giving a false impression of yellow leg colour. Specialized pubescences occur on the tro- chanters 3 where a tuft of stout hairs is found in some species, often inserted on a projection; on the tibiae 3 where a tibial floccus (a bunch of hairs) occurs in some species, in most cases inserted on a spur; and on the metatarsi 3 where stout short hairs occupy part of the lower face in some species (figs. 40 c, 42 d). Abdomen: Outline variable, usually comparatively stout in the females, with some notable exceptions, more slender in the males. The abdominal apex in the males is posteriorly rounded with the genitalia often clearly visible, in females it is posteriorly rounded to more or less acute. In males of some species T IV is conspicuously swollen to hold the large genitalia. Colour in most species dark throughout with yellow lateral spots on T II; in other species all dark, or mainly to all red, yellow or orange. Pubescence often concolorous with surface, most variable in density, Hurkmans: Monograph of Merodon evenness, length and colour; pruinose bands often present, in many cases combined with yellow to red background. These bands are quite variable in width and curvature throughout Merodon and are lacking altogether in several species. In the males S IV is usu- ally slightly modified posteriorly; in some species large paired appendages or a strong keel occur (figs. 36 b, 39 c, 40 b, 41 d, 42 e, 43 e, 44 b). Male genitalia: The male genitalia are often intri- cate and show enormous variation. They consist of two main parts, the aedeagus and the surstyle. Both parts are basally connected, by epandrium and basale respectively, to the inside of the genital cap. The ae- deagus is relatively the least variable, consisting of a usually tapering shaft, which apically bears paired chi- tinous plates, encasing the sperm duct. These plates vary in shape, size and degree of sclerotization and can be recumbent, erect or intermediate. The outer face of the shaft may bear paired spines, lobes or strongly sclerotized plates, or may have a subbasal ex- cavation. The inner face may show pubescence. The curvature of the aedeagus varies between species; both slender and stout aedeagi occur, while the apical shaft part (defined in fig. 10 a) may be weakly to strongly lengthened. The surstyle which envelops the cercus is much more variable. In its most simple form it con- sists of a paired, more or less S-shaped sclerotized plate, the apical part of which is identified as the pos- terior surstyle lobe which is always present. In most species additional lobes of the surstyle occur; often there is an anterior surstyle lobe which may either be free and projecting, or coalescent to various degrees with the posterior surstyle lobe; in some cases a mid- dle surstyle lobe is present as well. The anterior and posterior surstyle lobes may bear accessory lobes and / or specialized pubescences. The cercus is variable in shape and in the length and density of the pubes- cence. The genital cap (S X of Metcalf 1921, S VIII of McAlpine 1981, T VIII of Speight 1987) encasing the genitalia proper does not show much variability, although in one species, Merodon caudatus Sack, 1913, this cap is strongly elongate and bears an apical floccus conspicuous to the naked eye (figs. 8-17). Female genitalia: Simple, consisting of a telescope tube terminally fringed with pubescence (see figs. 52- 55 in Speight 1987). The structure of female genital- ia is quite uniform throughout Merodon and of little significance in establishing species groups or in spe- cies recognition. Sexual dimorphism: Rather pronounced in most species. In species with dichoptic males, eyes of fe- males wider apart. Metallic lustre in females usually less strong. Wings in females often less infuscate. In females pubescence on head, thorax and abdomen shorter and duller, but in many species the pruinose bands on thorax and abdomen clearer. In species with 191 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 males showing interalar pubescence bands these are sometimes lacking in females. Biology Merodon is widely distributed over the Palaearctic and Ethiopian regions and contains about 150 spe- cies. In general, very little is known of the biology, in spite of several species being quite common. Merodon species may account for a considerable part of the Syrphidae in the fauna of a region or country. There are over 50 species of Merodon in Turkey, where ma- ny species are very common. In Israel about 20% of the Syrphid species belong to Merodon (Kaplan & Thompson 1981: 198). The known larvae all develop in underground bulbs or rhizomes of monocotyledons. It is assumed that most if not all species share this way of life. Larvae were reared of Merodon amaryllidis Villeneuve de Janti, 1934 (Villeneuve de Janti 1934), Merodon bombiformis Hull, 1944 (Stuckenberg 1956) and Merodon equestris (Fabricius, 1794) (e.g. Lindner 1949). In the temperate zone the larvae survive the winter underground. Various species occur in regions having severe winters, e.g. the eastern Turkish mountains at 3000 m altitude, where species of Merodon are com- mon. The larvae are sheltered by the snow cover. On the other hand, Merodon occurs in tropical and sub- tropical regions as well. In these regions the larvae survive unfavourable climatic conditions, mainly drought, in their sheltered underground habitat. In relatively dry surroundings the various species seem to prefer relatively wet places (Hurkmans 1988). Some are mainly found at higher altitudes (e.g. Hurkmans 1988, Marcos Garcia 1989) where lower temperatures cause less evaporation, and therefore less extreme drought. In view of these adaptations Merodon should be considered xero-tolerant rather than xerophilic. In the northern and moister parts of the range, Merodon occurs in relatively drier habitats. This probably reflects that dry areas warm up quicker in spring and therefore allow a longer season. This is important given the restricted flight period. In view of the overall distribution, Merodon can be considered thermophilous, but this holds true only for the flight period. Cold season or drought conditions seem less important in view of the sheltered conditions during larval development. The various species of Merodon appear to have a preference for flowers of Umbelliferae (e.g. Suster 1959, Hurkmans 1985, 1988). Territorial behaviour was recorded in several Merodon species, e.g. by Fitzpatrick & Wellington (1983) and Hurkmans (1985, 1988). Defence of ter- ritories is more or less aggressive according to territo- 152 rial space available, presence of females, population density and weather. In some species the males emit piping sounds presumed to play a role in territorial defence and mating (Conn 1978; Treiber 1987; Hurkmans 1988). Copulation is either airborne or sedentary; both modes may occur in the same species (Conn 1978; Hurkmans 1988) and may be linked with morpho- logical differences (Conn 1978). Merodon equestris is capable of prolonged flight in copula. Duration of copulation is variable, from few seconds (Kabos 1939) to several minutes. In Merodon rufus copula- tion is accompanied by a loud piping sound (Treiber 1987). During oviposition several modes appear to occur: Kabos (1939) recorded that in Merodon equestris eggs were not attached to plants but pressed into the soil, while Hurkmans (1988) found that in Merodon loewi (Van der Goot, 1964) the females pressed their ab- dominal tip to the stems of Ornithogalum plants as if ovipositing; regrettably no larvae could be reared from the plants dug out. Apparently the number of generations per year var- ies throughout the range. Northern records are main- ly from July, while Mediterranean records of a single species in one region may extend from May through September. Marcos Garcia (1985) recorded that Merodon avidus (Rossi, 1790), and Merodon aeneus Meigen, 1822 are bivoltine in western-central Spain. Split occurrences of many species reflect only avail- able collection data and do not indicate that these species are bivoltine. Several species of Merodon have a dense and colourful pubescence and mimic bumblebees by their size and laborious flight, close to the soil. Brown (1951) noted that in Merodon equestris the colour forms most numerous in his samples mimicked the most common bumblebee. A cluster of small, strong- ly lustrous species resemble halictid bees in size, ap- pearance and behaviour. Division into species groups The divisions of Merodon by Becker (1912), Sack (1913a) and Paramonov (1926b) are based on extern- al similarities. The genitalia were not considered while some problematical species were not included in the species groups presented. The subgenus Exmerodon Becker, 1912, estab- lished to accommodate the type-species [by original designation], Merodon (Exmerodon) fulcratus Becker, 1912, is based on the dichoptic condition in males of fulcratus. Sack (1913a, 1931) and Paramonov (1924- 1937) did not follow Becker’s subdivision. The sub- division by Becker is not followed here for two rea- sons: first, dichoptic males are also found in other species not closely related to fulcratus and second, the fulcratus group does not occupy an isolated position. A first attempt to present a subdivision based on synapomorphous characters, including those of the male genitalia, is given here for the species having ar- cuate pruinose bands and abdomina with tapering second tergite. These two characters are judged to be apomorphic. The corresponding plesiomorphic con- ditions are: straight, oblique pruinose bands and non- tapering abdomina. Key to the species treated Note: Several species had to be left out of the key. Their genitalia could not be checked, since type-ma- terial could not be studied. These are: Merodon bes- sarabicus Paramonov, 1924 (the types are presumably lost); Merodon dobrogensis Bradescu, 1982; Merodon kiritshenkoi Stackelberg, 1960; Merodon tener Sack, 1913 (the only types left are 2, the d types are pre- sumably lost). Males: Key to species groups and species A. No anterior surstyle lobe present on genitalia, or: the surstyle consists of one mass in which the an- terior part may be differentiated (e.g. figs.); or: the outline of the surstyle ridge is fluent, semicir- cular or broadly rounded; or. anterior surstyle lo- be weak, posterior lobe very long (African spe- cies); or: anterior surstyle lobe medially recurved, totally coalescent with surstyle mass; or: three sur- style lobes of equal size, separated by two very deep sulci; or : abdomen relatively stout, T II not tapering, pruinose bands parallel to posterior ter- gite margins ... key in part II of this monograph — Anterior surstyle lobe well-defined, moderately large to large, the surstyle lobes are well differen- tiated. The surstyle lobes are not all separated (by two deep sulci), if a middle lobe occurs it is small and coalescent with either anterior or posterior lobe, and the lobes are not of equal size. Anterior lobe not medially recurved and totally coalescent with stylus mass, but free (although the apex may be recurved). Abdomen relatively slender, T II (more or less strongly) posteriorly tapering, abdo- minal pruinose bands arcuate. Species fulfilling all these requirements can be identified using the following key. 1. Anterior surstyle lobe short, rounded, pubescent, coalescent with surstyle mass (distinctus assem- beretta 12 — Anterior surstyle lobe more elongate, often well separated from surstyle mass, often with accesso- malobes or. specialized. parts"... 2 2. Anterior surstyle lobe narrow, posterior lobe me- 10. 11. 122 Hurkmans: Monograph of Merodon diolaterally aligned, often overlapping (alexeji Te) ioni rari 17 Posterior lobe dorsoventrally aligned ................ 3 Basitarsi 3 swollen, with specialized (dense, short, golden) pubescence on medial face; tibiae 3 often with apical floccus; sternite IV often keeled or strongly modified (tarsatus group) … 25 Sternite IV never strongly modified, if keeled the basifarsy3imormal ae a 4 Legs all dark, many species with conspicuous pu- bescence; anterior surstyle lobe wide, elongate, Er (A OD) EE 32 Wepsiparthy yellow. "2... 7 a 5 Flanges present on outer face of aedeagus (elegans PROUD) jets one eher aen seems’ 42 No such flanges present Secondary, lateral lobe present on posterior sur- style lobe; femora 3 much swollen, tibiae and tar- si all yellow; anterior surstyle lobe large, rather el- ongate, moderately wide; aedeagus with apical shaft part normal (vandergooti group) ................. RE Re DEL vandergooti sp. n. No secondary lateral lobe, or if present, femora 3 less swollen, anterior surstyle lobe narrow, elon- gate, ventrally recurved, anterior surstyle lobe narrow, ventrally recurved, apical shaft part of ae- deagus)lenethenede. 2. u... 2 0 002 7 Anterior surstyle lobe separated from surstyle mass/by.a sulcuss-. dda 8 Genitalia not with sulcate surstylus ................. 9 Large species, averaging 17 - 21 mm body length; abdomen predominantly yellow, antennae nor- mali(przzeronp) een pruni Rossi Smaller species, 7 - 15 mm, abdomen dark with crimson or dark red, antennae notably to ex- tremely lengthened (longicornis group) ........... 45 Posterior surstyle lobe with large, ventral accesso- walobeilerasssfemzonseroup) 22 ee crassifemoris Paramonov This lobe without large ventral accessory lobe .... Apical shaft part of aedeagus lengthened; anterior surstyle lobe usually not apically widened but if so, then bifid (alagoezicus group) .................... 46 Apical shaft part normal, anterior surstyle lobe apically widened, mot bifid 2 11 Aedeagus with 2 pairs of spines on outer face, an- terior surstyle lobe rectangular to diamond- shaped (nigritarsis group) Aedeagus smooth, anterior surstyle lobe ending in spathulate-semicircular apex (avidus group) ... EE EE er avidus (Rossi) Femora 3 very much swollen .......................... 13 Femora 3 only moderately swollen ................. 14 153 15 14. 15: 16. 17. 20. 21. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Larger species, body length 13 - 15 mm; genital- ia: aedeagus stouter, apical shaft of aedeagus shorter Smaller species, body length 10 - 13 mm; genital- ia: aedeagus slenderer, apical shaft of aedeagus JON LEE nn clunipes Sack Thorax and abdomen densely yellow pubescent; abdomen stout, T II strongly tapering; genitalia: both surstyle lobes elongate; body length 15 mm SIREN ALTRO Jee ull Aa. EEA ROIO mariae sp. n. Pubescence less dense, abdomen slenderer; if body length approx. 15 mm then abdomen very slender Genitalia: Posterior lobe very short; third anten- nal article subacute - acute; pubescence overall ver long ane) ASU Sono ottomanus sp. n. Genitalia: Posterior lobe longer; third antennal ATE CIETODTUSE e O PIE Eee .16 Body length 13 - 15 mm; both surstyle lobes co- alescent; abdomen very slender, dark with large yellow lateral spots ..................... distinctus Palma Body length 9-12 mm; surstyle lobes separated by a sulcus; abdomen yellow throughout ................ RR A]: NEE ENE AEE testaceoides sp. n. Genitalia: Anterior surstyle lobe showing elon- gate bald area; S IV deeply emarginate posterior- lyrics dee marginicornis sp. n. Anterior surstyle lobe without bald area ......... 18 . Genitalia: Posterior surstyle lobe with rounded bald knob in antero-lateral part ...................... 19 No such knob present, or knob very weak ......21 . Both surstyle lobes short; cercus relatively large, short,wide; T II with orange lateral spots bearing submetallic lustre; apical part of aedeagus covered by overlapping posterior surstyle lobes ............... star «ple ERLE ARENA RE ch Po ER sophron sp. n. Both surstyle lobes longer, cercus relatively small- er, more elongate; no submetallic lustre on lateral spots; aedeagus entirely visible ........................ 20 Pubescence on thorax and abdomen recumbent to oblique, golden; triangular processus with clear apical spur and short stout bristles ..................... BAALDE VUS. RER SALE RR altinosus sp. n. Pubescence on thorax and abdomen less conspic- uous, erect; no clear apical spur on triangular pro- cessus which bears longer, less stout bristles … … SARE SOIA IV) SACS TIGRI e alexeji Paramonov Anterior surstyle lobe with length/width ratio 1.2 - 1.3 in lateral view; posterior surstyle lobe short, rather spathulate in ventral view, coalescent … … HAT ALARM LU ZM kawamurae Matsumura Anterior surstyle lobe with length/width ratio of 2.0 or over; if posterior surstyle lobe short: acces- sory lobe present where anterior and posterior lobes meet, and anterior lobe strongly spathulate DIA 23: 24. DS) 26. 29) 30. Antennal ratio 2.1; genitalia: cercus placed far ventrad; accessory lobe present on posterior sur- style lobe; anterior lobe very elongate (an incon- spicuous bald knob present antero-medially on posterior surstyle lobe) ........... alexeji Paramonov Antennal ratio under 2.0; genitalia: cercus in nor- mal position; no accessory lobe on posterior sur- style loef e ee 23 Abdomen with strong golden lustre; genitalia: an- terior surstyle lobe widest in its mid-region; ae- deagus with two separate subapical cavities, apical shaft part slightly lengthened ...4ypochrysos sp. n. Abdomen without strong golden lustre; genitalia: anterior surstyle lobe equally wide throughout, or spathulate; aedeagus with one subapical cavity ... D NE A I ah ak I Re Pre cd Ai MEE nti oa 24 Touchline of compound eyes incomplete, very short; all legs with tarsi orange, tibiae orange with wide dark distal band; genitalia: anterior surstyle lobe equally wide throughout .....rufitarsis (Sack) Touchline complete, tl-v ratio 0.9; all legs dark except joints; genitalia: anterior lobe somewhat Spatial ateo en an hirsutus Sack Genital cap much elongate, with floccus ............ ARI ESTATE WERE eRe Clero le en caudatus Sack S IV normal, slightly arched, emarginate posteri- orly; basitarsi 3 with length/width ratio 2.5 and dense, short, golden pubescence on lower face; clear apical floccus present on tibiae 3 ................ PrO Lo A ri ri diga risi ia auronitens Sp. n. S IV with small (single or paired) median projec- ODA UE i SE ILE iis ilo Di S IV with large paired appendages … 29 . Antennal ratio over 2.8 ............ tangerensis sp. n. Antennal ratio under 2.3 ............................... 28 . Antennal ratio approx. 2.1; S IV with small pro- jections; legs pale brown ........... xanthipous sp. n. Antennal ratio approx. 1.3; S IV with single me- dian projection ending in acute spine ................. deste Re REES ankylogaster sp. n. S IV with long slender appendages; posterior proximal face of basitarsus 3 showing ovoid area of widely spaced setaceous pubescence bordered by long setae; apical floccus of tibiae 3 inserted on ENON PUS ee tarsatus Sack S IV with less long, robust appendages; if special- ized area present on basitarsi 3, pubescence in centre and on margin similar... 30 Genitalia: anterior surstyle lobe widening apical- ly, angular, its apical part projecting ventrad, with a fringe of short yellow pubescence PR Se nan tried enne persicus Sp. n. Genitalia: anterior lobe not widened apically, not Projecting MENACE een ee een 31 3. SPA 34. >: 36: STA 38. 32. Head: antennal ratio over 2.0; leg 3: length/width ratio of basitarsus 2.0; genitalia: anterior surstyle lobe tapering apically; aedeagus rather stout ....... MORN ce turkestanicus Paramonov Head: antennal ratio approx. 1.6; leg 3: length/ width ratio of basitarsus 1.4; genitalia: anterior surstyle lobe slightly spathulate; aedeagus slender- CO? iy ee As. oidipous sp. n. Wings strongly infuscate, deep brown except api- cally; large species (body length 15 - 18 mm) e antennen due crie velox Loew Wings clear or slightly infuscate (yellowish at most) . Thorax and abdomen densely pubescent, obliter- ating background colour; pubescence often red- an Or VELO nue 38 Thorax and abdomen moderately dense pubes- CENT surface well visiblen ev overs ten 34 Pubescence on thorax uniform; relatively stout species, with T II only moderately tapering ....35 Pubescence on thorax with dark interalar band; T II strongly tapering, slenderer species Genitalia: anterior surstyle lobe with long stalk; posterior lobe anteriorly with a low well-defined projection bearing dense pubescence; aedeagus rather stout; body length 14-15 mm SIR RC E warnckei sp. n. Genitalia: anterior surstyle lobe with short stalk; dense hair-tuft anteriorly on posterior lobe not inserted on a projection; aedeagus slender; body lede mmie brevis Paramonov Triangular processus modified, ending in large hooked spur; 3rd antennal article orange; Body ENEN na hamifer Sack Triangular processus on femora 3 normally shaped; abdomen slender, with bronze lustre; body length 10-15 mm Body length 11 - 15 mm; abdomen slender, with bronze lustre; pubescence yellow, rather dense; Éosdanithroughoutt een. na a ne BEER u... aberrans ssp. aberrans Egger Body length 12 - 14 mm; abdomen slender, with bronze lustre; legs with yellow markings ............ Pate mere aberrans ssp. flavitibius Paramonov Body length 10 - 11 mm; abdomen extremely slender; with slate grey lustre and often purple re- flections; pubescence grey to greyish yellow, rather sparse; legs dark throughout ..................... RE aankaart ledee aberrans ssp. isperensis ssp. n. Large species (body length 17 - 22 mm) ............. PPP RAMA Se eo sise nota clavipes Fabricius Smaller species (body length up to 16 mm) ....39 Femora 3 normally swollen; triangular processus with a strong, erect, apical spur; body length 12 - Bani ei vad ch Pate tte 40 Femora 3 very much swollen; triangular proces- 40. 41. 42. 43. 44. . Anterior surstyle lobe recurved mediad Hurkmans: Monograph of Merodon sus without apical spur, or this spur low, stout; body length 14 - 16 mm Tarsus 3: stout, with first article twice as long as second; apical spur of triangular processus on fe- mora 3 shorter; pruinose band on T IV chevron- shapeda ss karadaghensis Zimina Tarsus 3: slender, first article 1.4 x as long as sec- ond; apical spur of triangular processus on femo- ra 3 erect, longer; pruinose band on T IV round- [N es Cee dzhalitae Paramonov Antennal ratio 1.4; body length 14 - 16 mm; tri- angular processus on femora 3 finely serrate, nor- MARS. armadi. orione rn cupreus sp. n. Antennal ratio 1.8; body length 13 mm; triangu- lar processus strong, with slight apical spur ........ tee splendens sp. n. Anterior surstyle lobe with rounded profile; sul- cus between both surstyle lobes marked, but not very deep or wide; aedeagus rather stout Anterior surstyle lobe with angular profile; sulcus between both surstyle lobes very wide; aedeagus slenderer Head: oa 35°; abdomen very slender; anterior surstyle lobe with lateral ridge; flanges on outer ace orales one 0 testaceus Sack Head: oa 50°; abdomen moderately slender; an- terior surstyle lobe without lateral ridge; flanges onjaedeapusiighene See manicatus Sack Pruinose bands on abdomen conspicuous; S IV strongly arched; triangular processus on femora 3 bearing 6 - 9 bristles; posterior surstyle lobe with free anterior part separated by an oblique sulcus; cercus long, well projecting posteriorly ............... Resana elegans sp. n. Pruinose bands inconspicuous but visible; S IV strongly arched; triangular processus bearing 10 bristles; posterior surstyle lobe without free ante- rior part; sulcus rather quadrate; cercus shorter, not much projecting ................... bequaerti sp. n. . Antennal ratio 4.0 - 4.5 .............. longicornis Sack Antennal ratio 2.5 - 3.0 kaloceros sp. n. . Anterior surstyle lobe stalked, widening apically Anterior surstyle lobe narrow throughout . Both surstyle lobes separated by a deep sulcus... EE TR A taniniensis sp. n. Surstyle lobes coalescent; anterior lobe bifid ....... SR SR En eee PESTE toscanus sp. n. stia i n nitidifrons sp. n. Anterior lobe recurved ventrad to ventro-caudad . Anterior lobe recurved ventro-caudad, apical part more or less parallel with basal part ................ 50 Anterior lobe recurved ventrad, apical part pro- HAINE entri elet sente doge Sil 50: il 52: TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Tibiae 3 without large distal spur; posterior sur- style lobe with semicircular ventral projection … Secs ntl FEN PR LEHE lucasi sp. n. Tibiae 3 with large, triangular spur; posterior lobe without semicircular projection; lateral spots on T II with postero-lateral digitiform dark in- CENONE ccc ies alagoezicus Paramonov Anterior surstyle lobe simple; surstyle guarded by large lateral sclerotized plate, projecting far cau- dad; posterior surstyle lobe straight; aedeagus ba- SAIL eee eres en nn schachti sp. n. Anterior lobe with accessory lobe; no guarding lateral plate; posterior lobe twisted; aedeagus slender throughout ................... satdagensis sp. n. Anterior surstyle lobe with apical part rhomboid- al; posterior lobe higher; aedeagus with outer face all convex; trochanter 3 often with clear knob; rather robust species … … … … nigritarsis Rondani Anterior surstyle lobe suddenly widened apically, profile of apical part quadrate; posterior lobe low- er; aedeagus concave on basal outer face; tro- chanter 3 smooth; slenderer species Key to females of species 1. Abdomen dark throughout on surface (the pu- bescence may be brightly coloured!), often metal- lic lustrous, without red or yellow lateral spots . Abdomen showing yellow, orange or red lateral spots on T II, or with even more extensive yellow or red on abdomen Legs all dark, at most femoral apices and tibial bases very narrowly paler (but check 7: M. aber- rans ssp. flavitibius) Legs with yellow markings Pruinose bands present on abdomen, pubescence on posterior abdomen concolorous .................. 4 No pruinose bands present on abdomen ......... 10 Abdomen slenderer, in some cases with clear BISI eee 5 Abdomen stouter, dark, not clearly bluish lus- trous (but if clear grey lustre is present, see 7: AUCTIGIESSS PENISDENETISL) SORTE eee 8 Femoral ore LEALYASWOILENE ese eee ee 6 Femora 3 normal, not much swollen ................ 7 Abdominal surface colour brownish, with nar- rower pruinose bands ..................... clunipes Sack Abdominal surface colour blackish, with wider Abdomen with bluish metallic lustre, thorax of- ten with inconspicuous pruinose bands; wings apically slightly infuscate; femora straight; trian- gular processus normally shaped; body length 13 - 15 mm (in the ssp. isperensis the metallic lustre 156 10. 16. We . Thorax with pruinose bands is more greyish, while the abdomen is slenderer than in the nominal subspecies; the ssp. flavitibi- us, of which no records exist postdating the pub- lication of the original description, shows yellow zend mes). EE aberrans Egger Habitus similar, triangular processus ending in larger hacken en hamifer Sack Thoracic pubescence unicolorous, species of some 12 mm, bands on abdomen rather clear, nearly continuous, paired pruinose spots on T II BS LR oA a vain Pelee ti es hirsutus Sack Thoracic pubescence bicolorous Thoracic pubescence grey, dark interalar pubes- CEN CED AND LES speoacodacnaconcecanesse cupreus sp. n. Thoracic pubescence golden, but clearly paler posteriorly; relatively stout species with marked apical spur on triangular processus .… … de nis och haben ee karadaghensis Zimina Larger species, over 12 mm, densely pubescent at least on thorax, abdomen dark, blackish pubes- cent, with 2 pairs of outstanding, snow-white pu- DESCESDOL n quadrinotatus (Sack) Smaller species, sparsely pubescent throughout, slight bluish lustre on thorax and abdomen, basi- tarsus 3 swollen; body length 7.5 - 9 mm ........... Berane I tert aie turkestanicus Paramonov Thorax without bands, tibiae and tarsi partially dark, basitarsi 3 rather swollen . Species strikingly black-and-white with strong pruinose bands on abdomen Duller coloured species, bands inconspicuous .... ee hal warnckei sp. n. . Bluish lustre on thorax and abdomen, pruinose abdominal bands nearly continuous el le Air me nal ankylogaster sp. n. No bluish lustre, bands interrupted .................... . Well marked whitish pruinose bands present on HOTA rosie tee cee eet ee ee nee 15 Pruinose bands on thorax very weak, or longitu- dinal metallic bands pe or bands of different OO ee eee 32 Thorax unicolorous, without traces of bands...36 . Tergite II posteriorly strongly tapering, thorax with strong pruinose bands and extremely short, even pubescence; habitus very slender, body IRR n 16 Tergite II less tapering or habitus stouter, or smaller or larger species, thoracic pubescence longer sn eee ere er er 107 Femora 3 reddish to orange; midstripe on frons ExXtremielyznanrow ee distinctus Palma Femora 3 dark, midstripe normal ....................... "atea biarcuatus Curran 19: Dijk 28. 29. Antennal ratio 2.0 or less nn. 21 . Abdomen relatively stout, tapering posteriorly ... Abdomen slenderer, more cylindrical, lateral spots extended over much of 2nd tergite at least, in many cases over 3rd as well, crimson coloured Femora slender, abdomen with grey hue pro- duced by overall whitish pubescence; no blue lus- tre on abdomen … kaloceros sp. n. Femora swollen, abdomen dark with bluish lustre MM ee eee lusitanicus sp. n. . Antennal ratio 3.4 to 5.0 ............ longicornis Sack Antennal ratio 2.2 to 2.8 ..erivanicus Paramonov MSec ansthrouglioltss. tnt tn... 22 Legs yellow marked, or all yellow/orange ....... 25 . Large, robust species (17 mm), wings often infus- cate, thorax with dense pubescence ...velox Loew Smaller species (8 - 11 mm), wings clear ............ e ian alexeji Paramonov . Large species (17 - 21 mm), wings slightly infus- cate, abdomen clearly conoid in outline ............. MEI et. pruni Rossi Smaller species up to 15 mm, abdomen more cy- Fina bce En no. via 24 . Abdomen mainly pale orange, slender, pruinose bands on abdomen arcuate, posterior end of ab- CPU ER testaceus Sack Nbdomcnimamiyidark in 25 . Antennal ratio 2.0, triangular processus strongly SOM nt affinis Gil Collado Arena ton NME 26 . Tarsi 3 with dark upper face and contrasting, golden pubescence on sides; pruinose bands on thorax inconspicuous ............ nigritarsis Rondani Tarsi without this contrast or if so, thorax with RISAIE ANAS init 27 Legs nearly all orange ............... aureotibia sp. n. Legs with extensive dark surface .................... 28 Abdomen relatively lustrous, thorax with strong pruinose bands, often a fifth anteromedial band BESSER een alagoezicus group Comprising the following species: alagoezicus Paramonov, /ucasi sp. n., nitidifrons sp. n., satda- gensis sp. n., schachti SP. n., taniniensis sp. n. and toscanus sp. n. These species cannot at present be well separated in the female sex, as the pairing with the males is not certain. M. toscanus is known only from Italy, the other species are mainly known from Turkey. Abdomen duller, usually no 5th band on thorax DO a oe ae dee ARRE La A TR (RE RA CRT COREE 29 Small species, about 10 mm body length, thorax clearly but weakly banded .................. tener Sack Larger species, 12 mm or over, bands more men Si EN Sh DE a 30 Hurkmans: Monograph of Merodon 30. Frontal depression present; 2nd tergite bright CNEL Tobey, ERRO Mae EOS elegans sp. n. = Naldepressio MON {[0Nst ete ee ek ne, 31 Sle Femorasiswolleny curved nse eee PRE SS OLE ITER BaD femoratoides (Paramonov) — Femora 3 less swollen, straighter … avidus Rossi S2igltepsallidariso ne ce, pies 33 — Legs at least with yellow markings, thorax rather dulledark aero, A RE SIA 34 33. Femora 3 strongly swollen, abdomen bluish lus- trous, body length 10 - 13 mm ./usitanicus sp. n. — Femora3 slenderer, if in doubt: no blue abdomi- nal lustre; approx. 17 mm, wings infuscate ........ BERS | Anche MARI Pee Ne SERRE A VAE. VE velox Loew 34. Large species, 17 - 22 mm, wings often infuscate A TT SIRIA SIRO DS NLL PES pruni Rossi VAI E 35 35. Abdomen mainly orange, legs dark ..................... PO LEA DEM SIRIA manicatus Sack — Abdomen mainly dark, third antennal article slender, acute to subacute, size 7.5 - 10 mm, prui- nose abdominal bands conspicuous .................... aE eee cate he kawamurae Matsumura 36. Lateral spots on abdomen fiery orange or red, basitarsi 3 widened ........................ caudatus Sack — Lateral spots less conspicuous, abdominal pubes- cence bands clear, large to very large species ...37 37. Large species, 12 - 15 mm, abdomen stout with parallel, often yellow pruinose, conspicuous Dandie esis ie MN Ro mariae sp. n. — Very large species, 17 - 22 mm, bands whitish, less conspicuous, or dense pubescence obliterat- ID ED EUROS EE e eno clavipes (Fabricius) SYSTEMATIC PART The distinctus assemblage This set is a paraphyletic group sharing no other than the ‘general’ synapomorphies of this section of the genus, viz. the tapering T II and arcuate pruinose bands. Merodon biarcuatus Curran (figs. 21 a-c, 103) Merodon biarcuatus Curran, 1939: 7. Holotype 9 : ‘Forest of Namora, Robot, Morocco’ [near Rabat, approx. 33° 55’ N, 7° W] (AMNH) [examined]. Material examined. — France: 22 (BMNH, RMNH); Greece: 19 (ZMAN); Italy: 32 (NHMW, ZMAN, WH); Morocco: 22 (RMNH); Turkey: 122 (JLR, ZMAN, BMNH, WH). Description Female. — Head: Antennae orange, antenna 3 with upper margin convex, apex rounded, antennal ratio 157 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 2.2, ocellar angle 55°; pubescence pale yellow, rather sparse, erect, but moderately dense and darker on me- dian, lustrous 1/3-1/6 of frons and on vertex; face densely pruinose except on narrow median line. Thorax: Katepisternum, anepisternum and ane- pimeron, lateral dorsum and scutellum lustrous, bear- ing very short yellow even oblique pubescence; dark intaralar band present. Wings clear; halteres, squamae and antisquamae pale orange to yellow. Legs: Trochanters brown, femora dark, tibiae and tarsi orange to yellow; pubescence yellow, trochanter 3 smooth, femora 3 swollen with normal triangular processus bearing 7-9 bristles on serrate distal margin, apical bristle conspicuous. Abdomen: Extremely slender; T II orange, some- times with small dark medio-anterior lunule, strongly tapering; T III with conspicuous wide arcuate whitish pruinose band, background orange anterior, dark posterior to band; T IV-V dark, with conspicuous ar- cuate pruinose band on T IV; pubescence on all ter- gites concolorous to pale throughout. Body length 13-17 mm. Male. — Unknown. Diagnosis. — The extremely slender habitus will distinguish this species from other species, but M. bi- arcuatus is apparently closely related to distinctus. The pubescence on the frons of distinctus female is denser, oblique and golden; the lustrous midstripe is narrow- er, occupying 1/8 of its width; the pubescence on the thoracic dorsum is darker and shorter; the femora 3 are orange (dark in biarcuatus); the pruinose bands on the abdomen are narrower. Period of flight and distribution (fig. 103). — This species seems to occur throughout the mediterranean where it flies from May to August. Merodon clunipes Sack (figs. 22 a-c, 104) Merodon clunipes Sack, 1913a: 444. d Holotype: ‘Smyrna 49417’ ‘Billar (on lower face of same label) [= Izmir, Turkey] (ZMHB) [examined]. Merodon clunipes, Peck 1988: 169. Lampetia clunipes, Sack 1931: 311. Material examined. — Austria: 1d, 19 (RMNH); France: 154,29 (RMNH); Greece: 156,59 (NHMW, KBIN, RMNH, JLR, WH, MNHN); Hungary: 12 (RMNH); Italy: 146, 79 (NHMW, RMNH); Lebanon: 2d, 12 (NHMW, KBIN); Spain: 13 (RMNH); Switzerland: 1d (RMNH); Turkey: 148, 129 (BMNH, NHMW, ZMAN); Yugoslavia: 10d, 19 (RMNH). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apex rounded, antennal ratio 1.2, 158 pubescence dense, especially on face, erect, pale yel- low; vertex angle 35°, ocellar angle 45°, tl-v ratio 0.5. Thorax: Dark, without conspicuous lustre; pubes- cence dull yellow, pruinose bands clear. Wings very slightly infuscate; halteres, squamae and antisquamae yellow. Legs: Femora dark, tibiae 1-2 apically brownish, tibiae otherwise dark; tarsi brownish, darker apically; trochanter 3 with ridged projection; femora 3 very much swollen, curved, with normally shaped triangu- lar processus bearing 8-9 bristles on strongly serrate distal margin. Abdomen: Dark; slender to very slender; T II-IV bearing arcuate somewhat vague rather wide pruinose bands; pubescence dark, paler laterally and on T IV; densest on pruinose bands of T II; S IV narrowly emarginate posteriorly. Genitalia: Anterior surstyle lobe rather short, rounded, bearing dense, short, yellow, even pubes- cence throughout, showing a ridge bearing 5-6 strong bristles on medial face; posterior surstyle lobe coales- cent, elongate, posteriorly with dense, moderately long, yellow pubescence. Cercus rounded, bearing long, yellow pubescence. Aedeagus moderately long, apical shaft part slightly lengthened, fringed plates on thecal apex recumbent. Body length 8.5-13 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Ocellar angle 50°, frons with lustrous midstripe occupying 1/3 of width. Thorax: Pruinose bands quite conspicuous, pubes- cence less dense, rather recumbent on sides. Legs: Apices of femora paler, femora 3 less swollen, but if compared to females of other Merodon species still very much so. Abdomen: T II largely reddish to orange laterally, this colour may extend over T III as well; dark parts brown, not black. Body length 9-12 mm. Diagnosis. — The males of this species are rather similar to femoratus from which they differ in being smaller (range 8.5-13 vs. 12-16 mm) and having a more slender aedeagus with slightly lengthened apical shaft part. In the females the size ranges are 9-12 and 12-17 mm in clunipes and femoratus, respectively; the females of clunipes have a more slender abdomen with clear orange markings (all dark in femoratus females) with vague bands; the bands in the females of femor- atus are conspicuous and sharply bordered. Variation. — The ridge on trochanter 3 is rather variable in size, and seems correlated with specimen size: small specimens have small ridges, large speci- mens having large ones. Period of flight and distribution (fig. 104). — This species occurs through central Europe and the north- ern Mediterranean where it flies in June and July. Discussion. — One of the NHMW specimens identi- fied as clunipes has an identification label ‘M. striatus Sack det. Sack’. As far as could be traced, this name is unpublished. Merodon distinctus Palma (figs. 23 a-d, 103) Merodon distinctus Palma, 1863: 46. Holotype d : ‘from the field of Sanseverino’ [in the vicinity of Naples, Italy] [not located, not examined]. Lampetia dimorpha Szilady, 1940: 66. Holotype d: (and 9 Paratype) ‘Karacabey bei Brussa’ [West of Bursa, Turkey on coast of Marmara inland sea] [lost, formerly in TMA, not examined]. Syn. n. Merodon distinctus, Rondani 1868a: 22; Sack 1913a: 444; Peck 1988: 169. Lampetia distincta, Sack 1931: 313. Material examined. — France: 16 (BMNH); Greece: 236, 62 (KBIN, RMNH, BMNH, NHMW, JLR, WH); Italy: 16 (BMNH); Turkey: 52 (LR); U. S. S. R.: 12 (RMNH); Yugoslavia: 1d, 22 (LR, RMNH). Description Male. — Head: Antennae bright orange, antenna 3 with upper margin convex, apex obtuse, antennal ra- tio 1.5, pubescence golden with dark tuft in ocellar region; vertex angle 40°, ocellar angle 45°, tl-v ratio 0.6. Thorax: Dark, velvety lustrous on sides produced by recumbent pubescence, with clear pruinose bands on dorsum; pubescence golden, short, even. Wings clear; halteres, squamae and antisquamae yellow. Legs: All femora dark, not black, femora 1-2 apical- ly often brown to orangish; tibiae and tarsi yellow, legs with conspicuous golden pubescence through- out; trochanter 3 bearing distinct knob with apical obtuse ridge; femora 3 slightly swollen, rather straight, triangular processus with serrate distal mar- gin; in some specimens vague orange spot present ba- sally on femora 3; tibiae 3 rather abruptly widened distally (best seen in posterior view). Abdomen: Slender; T II-IV dark with rather wide sharply defined arcuate yellowish pruinose bands, narrowly to hardly interrupted, the band on T IV widened laterally and in some specimens slightly re- curved; T II with conspicuous yellow lateral spots; pubescence concolorous with surface; sternites I-III yellowish orange, IV darker, narrowly emarginate posteriorly, clearly though not strongly vaulted. Genitalia: The anterior surstyle lobe short, round- ed, with even, yellow, short, dense pubescence, co- alescent with posterior surstyle lobe which is low and Hurkmans: Monograph of Merodon elongate and bears long, erect, yellow pubescence. Cercus rounded to obtusely angular, with same pu- bescence. Aedeagus rather stout, slightly tapering, ap- ical shaft part somewhat lengthened, fringed plates on thecal apex rather wide and recumbent. Body length 12-16 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Midstripe on frons occupying 1/8 of width or even less; antenna 3 with dark upper margin in some specimens. Thorax: Pruinose bands more conspicuous, a thin praesutural crossband present in some specimens; pubescence ex- tremely short, as if shaven; pubescence in mid region of dorsum often dark. Legs: Deep orange to yellow all through, but triangular processus often darker. Abdomen: Even more slender than in d ; lateral spots larger, deeper coloured, pruinose bands less wide; T V bearing arcuate wide pubescence band; pubescence shorter than in d. Body length 13-15 mm. Diagnosis. — The combination of the very slender habitus, clear, sharply bordered pruinose bands, large lateral spots and genital structure will easily separate the males of distinctus from any other Merodon. The females are separated from the females of most other species by their relatively large size and extremely slender habitus. They can be separated from the fe- males of the alagoezicus group by the more cylindrical abdomen, and from the very similar females of biar- cuatus by the narrower frontal midstripe and the orange femora 3. Period of flight and distribution (fig. 103). — M. distinctus occurs from May through August in the mediterranean parts of Europe and Turkey. Discussion. — M. dimorphus Szilady, 1940 is tenta- tively considered a synonym of distinctus. In the orig- inal description dimorphus is stated to be allied to clu- nipes Sack, but the description points to very apparent differences with that species. From the description it can be inferred that dimorphus is very similar to, if not conspecific with, distinctus, the only difference appar- ent from the description is the colour of the pubes- cence on the head which is said to be pale in dimor- phus. Regrettably several features are not or very superficially described; the ‘spots’ on the abdomen may either be lateral spots or pruinose bands, or both. The figure of the thoracic dorsum as given by Szilady closely resembles this element in distinctus. No mate- rial of dimorphus was examined, as the type-material (formerly in TMA), the only material known, was lost in 1956 (Dr. A. Dély-Draskovits, in litt.). 159 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Merodon femoratus Sack (figs. 24 a-b, 104) Merodon femoratus Sack, 1913a: 446. Syntypes: ‘Korsika, Griechenland und Kleinasien’ [Corsica, Greece and Turkey]; depository unknown [not examined]. Merodon femoratus, Van der Goot 1981: 216, 218; Peck 1988: 169. Lampetia femorata, Sack 1931: 315; Séguy 1961: 179. Material examined. — Greece: 26, 52 (ZMUC, BMNH, KBIN, ZMAN, WH); Italy: 48 (ZMAN); Turkey: 156, 359 (JLR, BMNH, ZMAN, WH). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apex obtuse, antennal ratio 1.8; pubescence pale yellow, face and occiput densely white pruinose; vertex angle 35°, ocellar angle 45°, tl-v ratio 0.5. Thorax: Dark, katepisternum, anepisternum, ane- pimeron, lateral dorsum and scutellum metallic lus- trous, hardly pruinose except four longitudinal bands on dorsum; pubescence pale brown, moderately dense, erect, more even on dorsum but less so on sides. Wings very slightly infuscate; halteres, squamae and antisquamae pale brown. Legs: Dark, except for apices of femora and bases of tibiae of legs 1-2; trochanter 3 showing conspicuous hump, femora 3 very strongly swollen, strongly curved, with normally shaped triangular processus with strongly serrate, sometimes undulating margin bearing 5-10 bristles. Abdomen: Dark; rather slender; T II tapering, without lateral spots; T II-IV with interrupted mod- erately wide strongly arcuate whitish pruinose bands; pubescence concolorous, densest laterally on T II; S IV strongly vaulted, deeply narrowly emarginate pos- teriorly. Genitalia: The anterior surstyle lobe short, round- ed; bearing even, erect, short, dense, yellow pubes- cence apically; coalescent with posterior surstyle lobe, which shows long yellow pubescence posteriorly, is elongate and medio-laterally aligned apically. Cercus wide, short, rounded, not far protruding beyond pos- terior surstyle lobe, bearing dense, long, yellow pu- bescence. Aedeagus short, stout, smooth on outer face, fringed plates on thecal apex rather wide, recum- bent. Body length 12-16 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Ocellar angle = 50° to 60°, frons with lustrous midstripe occupying 1/4 of width. Thorax: Pruinose bands wider and clearer, often striking. Abdomen: Distinctly wider, typically black-and-white patterned, pruinose bands slightly less arcuate. 160 Body length 12-17 mm. Diagnosis. — M. femoratus is nearly similar to clu- nipes from which it is distinguished in both sexes by the greater size (12-17 mm vs. 9-12 in clunipes), in the males by the stouter aedeagus, and in the females by the black and white abdomen (brownish red in clunipes). From other species it differs in having a combination of very strongly swollen and curved fe- mora 3 and very simple anterior surstyle lobe in the males, and by a combination of strongly curved and swollen femora 3 and a striking black-and-white ab- domen in the females. Period of flight and distribution (fig. 104). — M. fe- moratus occurs in southern Europe and Turkey from May through August, both in lowland and mountain habitats (up to over 2000 m). It is absent from the western Mediterranean. Merodon mariae sp. n. (figs. 25 a-c, 105) Type-material. - d Holotype ‘France, Corse V. S. van der Goot / Asco 620 m [42. 15’ N, 9. 10° E] 11.vii.1967 (ZMAN). — Paratypes: 296, 102: same data as holotype 24, 12 (ZMAN); topotypic, 20.vii.1956, leg. v.d. Goot, 19 (ZMAN); topotypic, 5/6.vii.1961, leg. J. A. W. Lucas, 7d (JLR); topotypic, 9 /11.vii.1967, leg. Lucas, 3? (JLR); topo- typic, 1/11.viii.1959, leg. P. M. F. Verhoeff, 18 (ZMAN); topotypic, 25.vii.1977, leg. P. Goeldlin, 2d (coll. Goeldlin, Lausanne); topotypic, 27.vii.1977, leg. P. Goeldlin, 54 (coll. Goeldlin, Lausanne); “Corsica, La Foce de Vizzavona, 30.vi/ 31.vii.1893, Col. Yerbury 56 (BMNH); ‘Corte, Corsica, 2.vii.1893, Col. Yerbury 18, 12 (BMNH); ‘Evian, Corsica, 19.vii.1899 Col. Yerbury 16 (BMNH); Corsica, Vizzavona 900 1000 m, 2.x.1906, leg. F. Guglielmi’ 1d (BMNH); ‘Italia Sardegna Alghero 27/28.iv.1957, leg. C. A. W. Jeekel’ 16 (ZMAN); ‘Italia Sicilia Misterbianco 215 m, 27.v.1966 leg. H. J. P. Lambeck 16 (LR); “Environs de Salonique [Thessaloniki, Greece], Region du Mt. Profitis Ilias 768 m, Dr. A. Berton v.1918’ 12 (MNHN); Greece, Phokis, Delphi 19/22.v.1984, 700-850 m, leg. W. Hurkmans’ 2d, 12 (wn). Description Male. — Head: Antennae dark, antenna 3 with up- per margin straight, slightly sulcate longitudinally, apex acute, antennal ratio 1.6; vertex angle 25 to 30°, ocellar angle 40°; pubescence whitish yellow with dark tuft in ocellar region, very dense above antennae, causing silky lustre; tl-v ratio 0.35. Thorax: Dark, katepisternum, anepisternum, ane- pimeron, lateral dorsum and scutellum slightly lus- trous; pubescence rather dense, deep yellow, very even, short; vestigial longitudinal bands present in ‘most specimens. Wings slightly infuscate; halteres, squamae and antisquamae pale yellow. Legs: Dark, but tibial bases yellow marked; basitar- sus 2 brown, not black in many specimens; trochan- ter 3 smooth, femora 3 slightly swollen, rather straight, triangular processus with distal margin wavy, weakly serrate, but apically with two strong pedestals, holding 6-9 bristles; tibiae 3 with short rounded api- cal spur on inner face, basitarsi 3 somewhat swollen. Abdomen: Mainly yellow; moderately slender, T II conspicuously tapering; T II-IV bearing slightly arcu- ate, narrowly interrupted to continuous pale yellow pruinose bands and showing dark brown anterior strips and posterior lunules; pubescence erect to oblique, rather dense and long, pale orange to golden throughout; S IV strongly vaulted, posterior margin entire. Genitalia: Anterior surstyle lobe apically bearing short, even, yellow pubescence; subapical-medial ac- cessory lobe seen as acute angle in lateral view; coales- cent with posterior surstyle lobe; this lobe very elon- gate, low, posteriorly showing long, erect, rather dense, yellow pubescence and bearing a subapical, medial accessory lobe as well. Cercus large, elongate, with membraneous ventral plates basally, with dense, long, erect, yellow pubescence apically; Aedeagus rather stout, the outer face with elongate subapical cavity showing paired elongate projections; apical shaft part slightly lengthened; fringed plates near the- cal apex suberect. Body length 15 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Habitus: slightly stouter than males. Head: Midstripe on frons occupying 1/3 of width, noticeably tapering posteriorly. Thorax: Longitudinal bands slightly clearer; less lustrous on sides. Abdomen: T II hardly tapering; pruinose bands wider, less arcuate, strongly marked; apart from the quite different colour, not unlike those in femoratus. Body length 15-16 mm. Diagnosis. — The males of mariae can be distin- guished from those of other species by their possess- ing accessory lobes on both anterior and posterior surstyle lobe and robust habitus with typical dark yel- low to orange abdominal pubescence. The females are stout and show a very sharply defined pattern of prui- nose bands on the abdomen; they are unlikely to be confused with other species. Period of flight and distribution (fig. 105). — M. mariae is apparently common on Corsica and is known also from Sardinia, Sicily and Greece. This distribution pattern is not known from other Merodon. The flight period is from May through October, probably in more than one generation. Etymology. — It is a pleasure to dedicate this species HURKMANs: Monograph of Merodon to my wife, Marijke Wester, to commemorate her support and stimulation of my work. A noun in gen- itive case. Merodon ottomanus sp. n. (figs. 26 a-c, 106) Type-material. - d Holotype ‘Turkey, Hakkari, Tanin- Tanin pass 2200 m, 12.vi.1984 leg. J. A. W. Lucas’ (ZMAN). Paratypes: 7d, 39, all with same data as holotype (JLR). Description Male. — Head: Antennae orange, antenna 3 with upper margin straight, apex subacute, antennal ratio 1.7: vertex angle 50°, ocellar angle 65°; pubescence long, pale ochre yellow, moderately dense, with dark tuft in ocellar region; pubescence on compound eyes pale yellow to white, moderately dense, conspicuous- ly long; tl-v ratio 0.5. Thorax: Dark, faintly blue metallic lustrous to naked eye, under magnification showing very strong lustre partially dulled by disperse whitish pruinosity, the lustre strongest on katepisternum, anepisternum and anepimeron; pubescence as on face but longer still. Wings slightly yellow tinged, halteres, squamae and antisquamae yellow to pale yellow. Legs: All femora dark with yellow apices, tibiae yel- low with wide dark distal band, tarsi yellow; trochant- er 3 smooth, femora 3 slender, triangular processus low and bearing 4-6 bristles on serrate distal margin. Abdomen: Slender; dark; bluish metallic lustrous laterally on T II, there showing dense pubescence, otherwise pubescence moderately dense, pale yellow, and conspicuously long; T III with vestigial narrow arcuate pruinose band in some specimens; S IV strongly vaulted, posteriorly hardly emarginate, par- tially hidden by curved sides of T IV and its pubes- cence. Genitalia: Anterior surstyle lobe strongly yellow, short even pubescent, somewhat angular, coalescent with the short posterior surstyle lobe bearing long pu- bescence posteriorly. Cercus quadrate, with long dense yellow pubescence. Aedeagus with apical shaft part lengthened, fringed plates on thecal apex recum- bent, aedeagus subapically constricted in lateral view. Body length 9 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: ocellar angle 70°, frons lustrous, diffusely pruinose laterally. Thorax: Less lustrous. Abdomen: Pubescence slightly less dense and long, but still most conspicuous. Body length 9 mm. Diagnosis. — Apart from differences in the male genitalia, ottomanus can be distinguished from other 161 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 species by the long, pale erect pubescence of all ex- posed parts of the body; viewed with the naked eye this pubescence produces a hazy fringe around the in- sect. Period of flight and distribution (fig. 106). — The species occurs in south-eastern Turkey where it flies in June. Etymology. — The name ottomanus (an adjective) refers to the former Ottoman empire and was chosen to indicate the region of occurrence of the species, viz. Turkey. Merodon testaceoides sp. n. (figs. 27 a-c, 105) Type-material. — d Holotype ‘Yugoslavia, 30 km ZW Gradsko, 41. 26 N, 21. 50’ E, 13.viii.1963 exc. Leiden’ (RMNH). — Paratypes: 16 (ZMAN), 19 (RMNH), 22 (LR), same data as holotype. Description Male. — Head: Antennae dark, antenna 3 with up- per margin slightly convex, apex obtuse, antennal ra- tio 1.5; vertex angle 25°; ocellar angle 40°; pubescen- ce pale yellow; tl-v ratio 0.5. Thorax: Dark, rather lustrous on katepisternum, anepisternum, anepimeron, lateral scutellum and dorsum; pubescence moderately dense, pale yellow; dorsum showing inconspicuous pruinose bands. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: All femora dark with yellow apices, tibiae yel- low with broad dark distal band, tarsi yellow to red; trochanter 3 with low ridge; femora 3 swollen, curved, bearing normally shaped triangular processus with 8 bristles on slightly serrate distal margin, widest spaced basally. Abdomen: Slender; conspicuously orange, T II ta- pering considerably, with dark anterior lunule in some specimens; T II-IV with arcuate pruinose bands, interrupted on T II-III; pubescence pale throughout, some dark pubescence often present on dark anterior lunule of T II; S IV orange, strongly vaulted, incised up to 2/3 its length from posterior margin. Genitalia: Anterior surstyle lobe low, rounded-el- ongate, with short, dense, even, yellow pubescence throughout, separate from posterior surstyle lobe by moderately deep and wide sulcus; posterior surstyle lobe rounded, rather low, bearing moderately dense, erect, yellow pubescence throughout, slightly denser apically. Cercus rounded, rather small, bearing dense yellow pubescence. Aedeagus showing paired humps basally on outer face,and marked subapical cavity; ap- 162 ical shaft part slightly lengthened, fringed plates on thecal apex recumbent to somewhat suberect. Body length 10-12 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Lustrous midstripe occupying 1/3 of frons, ocellar angle 50°. Abdomen: pale reddish to yellow throughout, T II less tapering. Body length 10-12 mm. Diagnosis. — The relatively small size and striking orange abdomen in both sexes precludes confusion with all but a few species. M. testaceoides is similar ex- ternally to testaceus from which it is distinguished in the males by the very different genitalia; the females of testaceoides can best be distinguished by the paler colour of the abdomen and longer, denser abdominal pubescence. Both sexes differ from the species of the longicornis group by the all red to orange abdomen, which in that species group is partially crimson, par- tially black; moreover, the antennal ratio is less in tes- taceoides. Period of flight and distribution (fig. 105). — This species is known only from Macedonia, where it was collected in August. Etymology. — The epitheton testaceoides refers to Merodon testaceus to which this species is externally similar; the suffix -ides means ‘in likeness of. An ad- jective. The alexeji group Apomorphies: Anterior surstyle lobe elongate; pos- terior surstyle lobe high to very high, curving medial- ly. Description. — Head: Antennae dark, 3rd article red or orange in some species, antennal ratio 1.5 to 2.4; eyes with touchline incomplete to equalling 0.15-0.9 vertex height; pubescence golden to pale yel- low with more or less conspicuous dark tuft in ocellar region. vertex angle and ocellar angle variable, oral margin well defined. Thorax: Dark with some lustre and in some species with traces of longitudinal pruinose bands; wings mostly clear, veins brown, paler in rufitarsis and soph- ron; halteres, squamae and antisquamae yellow. Legs: All dark or with yellow markings on tibiae and tarsi; trochanters 3 smooth, with faint (Aypochry- sos) or strong (marginicornis, rufitarsis) edge or knob; femora 3 mostly rather swollen, straight; basitarsi 3 somewhat swollen in marginicornis. Abdomen: Rather slender; T II moderately taper- ing, with slightly to more strongly arcuate pruinose bands of variable clarity; lateral spots yellow to orange, or bluish, or lacking; S IV normal. Genitalia: The anterior surstyle lobe elongate to ex- tremely elongate, bearing short, dense pubescence at least apically, separated from the posterior surstyle lobe by a shallow sulcus, or coalescent; the posterior surstyle lobes much extended medially, often overlap- ping, hiding part of the aedeagus from view. Aedeagus smooth on outer face, mostly slender, api- cal shaft part slightly lengthened, subapical cavity well defined. Merodon alexeji Paramonov (figs. 28 a, b, 29 a, 81) Merodon alexeji Paramonov, 1925: 155. Holotype d (and ? paratype) ‘Kechanowka, Balta, Odessa, Ukraine, U. S. S. R., 1.vi.1924, leg. A. Paramonov’ [not examined, pre- sumably lost]. Merodon alexeji, Paramonov 1935: 164, 166; Liepa 1969: 20; Van der Goot 1981: 215, 218; Violovitsh 1983: 124; Peck 1988: 167. Lampetia alexeji, Sack 1931: 304. Material examined. — Algeria: 28 (ZMAN, WH); France: 56, 42 (MNHN, KBIN, ZMAN); Greece: 58, 42 (BMNH, KBIN, ZMUC, NHMW); Italy: 14 (NHMw); Mongolia: 19 (TMA); Morocco: 66, 19 (MNHN); Spain: 2d (ZMAN, WH); Tunisia: 28 (LR); U.S. S. R. : 54, 39 [Id identified by original author, in KBIN] (BMNH, KBIN, ZMHB, H. Hippa, Helsinki). Description Male. — Head: Antenna dark brown, antenna 3 ta- pering, upper margin rather straight, apex subacute, antennal ratio 2.1; pubescence whitish to pale yellow, rather sparse with dark tuft in ocellar region; vertex angle 40°, ocellar angle 50°, ocelli rather large; tl-v ra- tio 0.45; pubescence on eyes pale, inconspicuous. Thorax: Dark; katepisternum, anepisternum, ane- pimeron, dorsum and scutellum metallic lustrous, with short moderately dense, erect, uneven yellow pubescence, with variably clear interalar band; in some specimens vague pruinose bands present. Wings clear, brown veined; halteres, squamae and antisqua- mae yellow. Legs: All dark; trochanters 3 smooth, femora 3 rather swollen, with normal triangular processus bear- ing 6-12 bristles, basitarsi 3 and other tarsi 3 normal, tibiae 3 rather slender. Abdomen: Rather slender, with tapering T II, somewhat triangular, dark with slight metallic lustre overall (strongest laterally), bearing dark orange later- al spots on T II; T I-IV with rather narrow, arcuate, interrupted whitish pruinose bands, pubescence pale, inconspicuous medially. Genitalia: The anterior surstyle lobe much elon- gate, slightly spathulate, with short, even, dense, erect, yellow pubescence apically and along dorsal Hurkmans: Monograph of Merodon margin; separated from posterior surstyle lobe by a sulcus; posterior surstyle lobe divided into basal and apical parts by a suture, anteriorly with smooth knob; apical part mediolaterally aligned, rather high, these parts on both sides overlapping; posterior surstyle lobe with rather dense, yellowish, erect pubescence but sparser on apical margin. Cercus conspicuous, rectangular, with dense, long, uneven, yellow pubes- cence. Aedeagus rather slender, smooth on outer face, apical shaft part slightly lengthened, fringed plates on thecal apex oblique to recumbent. Body length 9-11 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Ocellar region with darker pubescence; ocellar angle = 60°; lustrous mid- stripe on frons occupies all but fringes. Thorax: Pruinose bands clearer, less lustrous. Legs: The femo- ra 3 rather variable in the degree of swelling. Abdomen: Triangular, pruinose bands wider, lateral spots variable, not always clear. T V acute, almost bald, mostly inconspicuous. Body length 9-11 mm. Variation. — Especially in the males M. alexeji is variable in the clearness of the abdominal lateral spots, the number of bristles on the triangular proces- sus, and slightly so in the elongation of the knob on the posterior surstyle lobe and the position (not the shape) of the cercus. Diagnosis. — M. alexeji can be separated from other species in the alexeji group by the presence of both the knob on the posterior surstyle lobe and the rectangu- lar well protruding cercus. The latter feature separates from altinosus (cercus inconspicuous) and sophron (cercus short, very wide) which also have the posteri- or surstyle lobe knob. alexeji is separated from altino- sus by the lack of dense abdominal pubescence as well. Period of flight and distribution (fig. 81). — The imagines occur from May in the Mediterranean to July in the northern parts of the range, which in- cludes the mediterranean countries and the temperate parts of western and central Eurasia. Discussion. — Although type-material of this spe- cies was not available there is no doubt about the identity of alexeji. The original description mentions a number of diagnostic characters; moreover material examined and identified by S. J. Paramonov has been examined. 163 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Merodon altinosus sp. n. (fig. 29 b, 102) Type-material. — d Holotype ‘Turkey, Hakkari, Suvarihalil pass 1250 m., W. side near Habul Deresi [37° 35° N, 43° 20° E], 13.vi.1984, leg. Lucas (ZMAN). Description Male. — Head: Antennae blackish, antenna 3 with upper margin slightly concave, apex subacute, anten- nal ratio 2.4, pubescence golden, a dark tuft present in ocellar region; vertex angle 30°, ocellar angle 45°, tl-v ratio 0.6. Thorax: Dark, slightly metallic lustrous on katepis- ternum, anepisternum, anepimeron, dorsum and scu- tellum, with moderately dense, longish, uneven, golden pubescence, without pruinose bands. Wings slightly troubled evenly, especially basally and on an- terior margin; halteres, squamae and antisquamae yel- low. Legs: All dark; trochanters 3 smooth, femora 3 swollen, rather straight, bearing normal triangular processus with strong apical spur and 8 extremely short stout bristles; tibiae 3 and tarsi 3 normal. Abdomen: Dark, tapering posteriorly; T II with yellow lateral spots; dense, golden, oblique pubes- cence present throughout; vestigial arcuate pruinose bands present on T II-IV, clearest on T III. Genitalia: The anterior surstyle lobe elongate, ante- riorly with dense, short, even, yellow pubescence, co- alescent with posterior surstyle lobe; that lobe medio- laterally aligned in its apical part, overlapping; an elongate smooth knob present anteriorly on the pos- terior surstyle lobe; posterior part with long, rather dense, erect, uneven, yellow pubescence. Cercus moderately large, not far protruding, bearing dense, erect, long, yellow pubescence. Aedeagus rather slen- der, smooth on outer face, fringed plates on thecal apex suberect. Body length 9.5 mm. Female. — Unknown. Diagnosis. — M. altinosus resembles alexeji from which it is separated by the conspicuous golden pu- bescence on the abdomen, the extremely short bristles on the triangular processus with its strong apical spur and the less protruding cercus; it is distinguished from sophron by the same features, and from other species in the alexeji group by the presence of a knob on the posterior surstyle lobe. Period of flight and distribution (fig. 102). — The species is known from the holotype only, collected in eastern Turkey in June. Etymology. — The name altinosus (an adjective) de- 164 notes the golden habitus of the species, and is derived from the Turkish altin, meaning gold, and the Latin suffix -osus, meaning full of. Merodon hirsutus Sack (figs. 30 a-c, 82) Merodon hirsutus Sack, 1913a: 435. Lectotype d (here des- ignated): ‘Djebel Akra [35° 35’ N, 36° 15° E] vi.85 [1885], N. Syria, Dr. E. Leuthner/ Lampetia hirsuta Sack det. Sack’ (NHMW) [examined]. Merodon hirsutus, Peck 1988: 171. Lampetia hirsuta, Sack 1931: 318; Séguy 1941: 14. Material examined. — Israel: 32 (BMNH, RMNH); Jordan: 29 (RMNH); Syria: 9d Paralectotypes, with same data as lec- totype (NHMW, ZMAN) [examined]; 1% Paralectotype, ‘Syria/ hirsutus Sack/ coll. J. Villeneuve: Lampetia hirsuta Sack RMHN Belg. 15.392’ (KBIN); Turkey: 22 (BMNH). Description Male. — Head: Antennae blackish, antenna 3 with straight upper margin, apex acute, antennal ratio 1.9; pubescence yellow, dark tuft in ocellar region weak; vertex angle 40°, ocellar angle 50°, tl-v ratio 0.9; pu- bescence on eyes inconspicuous. Thorax: Dark, slightly metallic lustrous on katepis- ternum, anepisternum, anepimeron, dorsum and scu- tellum; pubescence dense, erect, uneven, yellow, moderately long. Wings clear, brown veined; halteres, squamae and antisquamae yellow. Legs: Dark with yellow markings on both ends of tibiae, rather slender; bearing normal triangular pro- cessus with short stout bristles, tibiae 3 and tarsi 3 normally shaped. Abdomen: Dark, rather slender to somewhat stout, lacking strong lustre (not due to surface sculpturing); laterally slight micaceous lustre present; T II-IV with slightly arcuate, moderately wide whitish interrupted pruinose bands; pubescence rather long, yellowish, densest laterally, dark and sparser on posteromedian parts of T III-IV; sternites dark, with scattered whit- ish pubescence; S IV slightly vaulted. Genitalia: Anterior surstyle lobe extremely elon- gate, anteriorly rather rectangular, bearing dense, short, yellow pubescence, slightly spathulate, separat- ed from posterior surstyle lobe by shallow but clear sulcus; posterior surstyle lobe rounded, showing un- even, long, dense, yellow pubescence especially poste- riorly. Cercus conspicuously protruding, with dense, long, yellow pubescence. Aedeagus smooth on outer face, stout basally, slender apically, apical shaft part slightly lengthened, fringed plates on thecal apex sub- erect. Body length 10 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Ocellar angle 55°, shining midstripe on frons occupies 0.6 of width. Thorax: Pubescence comparatively shorter, more even. Legs: femoral apices very narrowly brownish. Abdomen: Slightly more slender; pruinose band on T II reduced; those on T III-IV clearer, wider, nearly continuous, more arcuate. Body length 9.5-10.5 mm. Variation. — There is little variation in the males (all specimens examined are from same locality); the females show slight size variation. Diagnosis. — Merodon hirsutus is externally similar to M. hirtus Sack from which it is easily distinguished by its very different genitalia; study of the genitalia is essential since both species fly together in the same re- gion. From the other species of the alexeji group, hir- sutus can be separated by the extremely long anterior surstyle lobe, the relatively low posterior surstyle lobe and lack of yellow lateral spots on T II. Period of flight and distribution (fig. 82). — M. hir- sutus occurs in the eastern Mediterranean region from March through June. Merodon hypochrysos sp. n. (figs. 31 a-c, 82) Type-material. . — 4 Holotype ‘Turkey, Adiyaman, 10 kms. North of Celikhan, Alu Haral Golii [38° 05’ N, 38° 22° E], 1450 m, 1.vii.1986, leg. Lucas’ (ZMAN). Paratypes: 206 ‘Turkey, Hakkari, Sat Daglari, Varegòs [37° 25’ N, 43° 55° E] 1600-1650 m, 15/18.vi.1984, leg. Lucas’ (JLR). Description Male. — Head: antennae dark, antenna 3 red, upper margin convex, apex rounded, antennal ratio 1.5, ver- tex angle 40°, ocellar angle 45°, tl-v ratio 0.2; pubes- cence pale yellow, with dark tuft in ocellar region. Thorax: Black, lustrous on katepisternum, anepis- ternum, anepimeron, dorsum and scutellum; pubes- cence rather sparse, golden, paler down sides and on scutellum, rather even. Wings slightly tinged, veins brown, halteres, squamae and antisquamae pale yel- low. Legs: Dark, apices of femora and bases of tibiae paler; trochanters 3 with inconspicuous edge; femora 3 much swollen, straight, bearing normal triangular processus with weakly serrate distal margin holding 5- 8 small bristles; tibiae 3 long, slender, bearing small apical spur distally on medial face; tarsi 3 normal, rel- atively small. Abdomen: Dark, golden metallic lustrous with bluish to bronze reflections on sides; T III-IV with moderately arcuate, whitish to pale yellow, interrupt- Hurkmans: Monograph of Merodon ed, moderately wide pruinose bands; pubescence pale golden, densest laterally. Genitalia: Anterior surstyle lobe elongate, slightly diamond-shaped, apically with short, even, dense, yellow pubescence, posterior surstyle lobe more or less coalescent but steeply rising, rather rounded, slightly mediolaterally aligned apically; posterior por- tion showing dense, erect, yellow pubescence. Cercus well protruding, moderately large, rounded, bearing dense yellow pubescence. Aedeagus basally stout, ta- pering, subapical cavity elongate and showing con- spicuous paired chitinous plates; an accessory triangu- lar cavity present just below the recumbent fringed plates on the thecal apex (which acutely protrudes well beyond the plates). Body length 7 mm. Female. — Unknown. Diagnosis. — M. hypochrysos can be distinguished from other species in the alexeji group by its golden surface and pubescence of the abdomen; the lack of a knob on the posterior surstyle lobe distinguishes from the larger but superficially similar altinosus. Period of flight and distribution (fig. 82). — This species is known from the mountains of eastern Turkey where it occurs in June and July. Etymology. — The epitheton is derived from the Greek words hypo, i.e. under, less than, and chrysos, meaning gold, on account of the moderate golden lustre of the male. An adjective. Merodon kawamurae Matsumura (figs. 32 a-e, 79) Merodon kawamurae Matsumura, 1916: 257. Described af- ter an unknown number of specimens from Kumamoto, Kyushu, Japan, leg. Kawamura. [not examined]. Lampetia micromegas Hervé Bazin, 1929: 111. Lectotype d (here designated): ‘Tchen-Kiang, 13.iv.1918/ Lampetia micromegas H. B. type’ [examined] (MNHN). Syn. n. Merodon kawamurae, Paramonov 1925: 160; Shiraki 1930: 206. Lampetia kawamurae, Hervé Bazin 1929: 111; Sack 1931: 319. Merodon kawamurai; Shiraki 1968: 198; Peck 1988: 171. Merodon micromegas, Peck 1988: 172. Material examined. — China: Paralectotypes of micromeg- as: 198, 399: Topotypic 9d, 182 (MNHN); ‘Shia-Shu’ 62 (MNHN, BMNH); ‘Che-Mo’ 96, 102 (MNHN, BMNH); ‘Nanking’ 39 (MNHN, BMNH); ‘Hoa-Chan 1d, 29 (MNHN), Collection dates from 1.iv.1918 to 16.v.1918. — Non types: China: 19 (NHMW); Japan: 2d, 19 (BMNH). 165 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Description Male. — Head: Antennae orange, antenna 3 with upper margin convex, apex acute to rounded, most variable, antennal ratio 1.6; vertex angle 35°, ocellar angle 50°, tl-v ratio 0.15 to 0.3; vertex sides straight to slightly concave; pubescence pale yellow, with dark tuft in ocellar region. Thorax: Metallic lustrous on katepisternum, ane- pisternum, anepimeron, dorsum and scutellum, bear- ing moderately dense and long, uneven, yellow to white pubescence. Wings slightly tinged, halteres, squamae and antisquamae yellow; very faint longitu- dinal striation present in some specimens. Legs: All femora dark brown, tibiae yellowish with broad dark band in distal half (in some specimens more extended), tarsi, especially tarsi 3 dark above, with golden pubescence underneath; trochanters 3 smooth to distinctly edged (seen as little spine in end- on view), femora 3 rather swollen, with normal trian- gular processus bearing 12-15 spinules of which the apical 1-3 are robust, the others rather fine; tibiae 3 and tarsi 3 normal. Abdomen: Dark; T II-IV bearing widely to hardly interrupted, slightly arcuate, white, laterally widening pruinose bands; T II with rather large orange lateral spots, dark-centred in some specimens; pubescence inconspicuous, pale yellow to white, densest laterally on T II-IV and abdominal tip; S IV weakly vaulted, posterior margin entire to shallowly emarginate. Genitalia: The anterior surstyle lobe rather large, slightly spathulate, elongate, rounded, apically with dense, short, even, yellow pubescence; posterior sur- style lobe coalescent, strongly mediolaterally aligned, overlapping, posteriorly with dense, erect, yellow pu- bescence. Cercus wide and short, bearing dense, erect, long, yellow pubescence. Aedeagus smooth on outer face, apical shaft part short, fringed plates on thecal apex large, suberect to erect. Body length 7-11.5 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Antenna 3 generally less acute than in males; ocellar angle 60°; frons strongly golden lustrous throughout, diffusely prui- nose on margins, showing white pubescence; vertex slightly but clearly edged anteriorly in some speci- mens. Thorax: Clearly more lustrous; pubescence less dense; traces of striation clearer. Legs: Often relative- ly more yellow, especially on tibiae. Abdomen: Slenderer, lateral spots larger. Body length 6-12 mm. Variation. — M. kawamurae is one of the most var- iable species in Merodon. The variation observed seems to be size-correlated: the largest specimens show the strongest lustre, the most acute antenna 3, 166 most edged trochanters 3 and most vaulted and emar- ginate S IV (males), or strongest edged vertex (fe- males). In view of this variability, including size vari- ation of 100%, more than one species would seem to be involved. There are however several strong argu- ments for considering all material conspecific: 1. there are gradual transitions in all variable features; 2. all males are completely similar in the genitalia; 3. many cases of such extreme variation are known in other insect species, some of which have been shown to be the direct effect of food availability. Diagnosis. — M. kawamurae can be distinguished by its clearly spathulate anterior surstyle lobe in the males and by the golden lustrous vertex, slender ab- domen and (some specimens) the edge on the vertex. Moreover it is the only small-sized Merodon species currently known from China and Japan, the only larger-sized species being equestris Fabricius. Of other Merodon species the only one possibly occurring in this region is alexeji which has a much longer anterior surstyle lobe and more protruding cercus (fig. 28 a). M. alexeji has been found in Mongolia but more east- ward localities are unknown. Period of flight and distribution (fig. 79). — M. kawamurae is known from eastern China and Japan, and occurs in April and May. Discussion. — The synonymy of M. micromegas with M. kawamurae might be questioned since no type-material of M. kawamurae has been examined. After Horn and Kahle (1936) the Matsumura types might be at the Agricultural Department of Hokkaido University, Sapporo, Japan, but inquiries there have not confirmed the presence of the kawam- urae types at this institute. The illustrations and de- scriptions of kawamurae and micromegas leave hardly any doubt as to their conspecificity; moreover, Hervé-Bazin himself already stated that micromegas might be conspecific with kawamurae and described micromegas since he ‘thought the description of kaw- amurae as given by Matsumura (1916) insufficient’ (Hervé-Bazin 1929). The Japanese material currently known is in the BMNH. Merodon marginicornis sp. n. (figs. 33 a-c, 82) Type-material. — d Holotype ‘Iran, Chiraz [26° 40° N, 52° 30’ E], 11.iv.1937, Coll. F. H. Brandt (RMNH). Description Male. — Head: antennae large, antenna 3 orange, apex acute, upper margin edged, concave, antennal ratio 2.4; vertex angle 35°, ocellar angle 65°, tl-v ratio 0.4, pubescence pale yellow. Thorax: Dark; katepisternum, anepisternum, ane- pimeron, dorsum and scutellum bronze lustrous; pu- bescence rather sparse, short, uneven, pale yellow; no traces of pruinose bands present. Wings slightly tinged, brown veined; halteres, squamae and anti- squamae pale yellow. Legs: Dark, but tarsi brownish orange; trochanters 3 with distinct knob; femora 3 swollen, rather straight, with low, strongly serrate triangular proces- sus bearing 8 spinules; tibiae normal, basitarsi 3 slightly swollen, other tarsi 3 normal. Abdomen: Dark, vaguely bluish lustrous, with rather vaguely bordered but well marked, slightly ar- cuate, interrupted whitish pruinose bands on T II-IV; T II bearing strongly lustrous bluish lateral spots; pu- bescence rather sparse, whitish, densest antero-lateral- ly; S IV deeply emarginate posteriorly, somewhat vaulted. Genitalia: Anterior surstyle lobe elongate, bearing short, even, yellow, erect pubescence dorsally and along margins, showing elongate bald area, bordered by a rim, on lateral face; the posterior surstyle lobe separated by a shallow sulcus, elongate, posteriorly with yellow, dense, erect pubescence. Cercus incon- spicuous, with long, dense, yellow pubescence. Aedeagus smooth on outer face, with long subapical cavity, showing paired elongate, heavily sclerotized structures basally; fringed plates suberect, apical shaft part short. Body length 9 mm. Female. — Unknown. Diagnosis. — M. marginicornis can be easily distin- guished by the bald, rimmed area on the lateral face of the anterior surstyle lobe. Period of flight and distribution (fig. 82). — M. marginicornis is known only from the holotype, col- lected in south-western Iran in April. Discussion. — M. marginicornis has been assigned to the alexeji group in view of the structure of its male genitalia; it has somewhat swollen basitarsi 3 [a syn- apomorphy for the tarsatus group], but lacks the modification of S IV [the other synapomorphy for the tarsatus group]. Therefore this species may in fact occupy a position in between these two groups. Etymology. — The epitheton marginicornis refers to the edged upper margin of the antenna 3; it is derived from the latin margo, meaning edge, and cornu, meaning horn, in insects often used in referring to an- tennae. A noun in apposition. Hurkmans: Monograph of Merodon Merodon rufitarsis Sack stat. n. (figs. 34 a-c, 83) Merodon fulcratus Sack, 1913, subsp. rufitarsis Sack, 1913a: 438. Holotype d: ‘Alai mont [the Alajsski Mountains, Tadzhikistan, approx. 39° 30’ N, 71° to 73° E] (ZMHB) [examined]. Merodon (Exmerodon) fulcratus ssp. rufitarsis Peck 1988: 165. Description Male. — Head: Antennae dark, antenna 3 red, apex rounded, upper margin convex, antennal ratio 1.6; vertex angle 30°, ocellar angle 50°, pubescence pale yellow; touchline incomplete, tl-v ratio 0.15; comp- ound eyes with inconspicuous white pubescence. Thorax: Dark; katepisternum, anepisternum, ane- pimeron, dorsum and scutellum slightly lustrous, bearing moderately long and dense, yellow, rather un- even pubescence, without traces of pruinose bands. Wings clear, veins pale brown, halteres, squamae and antisquamae pale yellow. Legs: All femora dark, tibiae orange with wide dark distal band, tarsi orange; trochanters 3 with low sharp edge, femora 3 moderately swollen, rather straight, bearing normal triangular processus with 10 spinules; tibiae 3 and tarsi 3 normal. Abdomen: Dark, without lateral spots or pruinose bands; pubescence rather dense, uneven, yellow, sub- erect; S IV strongly keeled; abdominal surface slight- ly lustrous throughout, strongest laterally. Genitalia: Anterior surstyle lobe much elongate, with short yellow even pubescence apically and down margins; posterior surstyle lobe separated by shallow sulcus, large, strongly mediolaterally aligned, showing rather long, yellow pubescence apically and posterior- ly. Cercus rather short, wide, with dense, long, yellow pubescence. Aedeagus smooth on outer face, with paired chitinous plates in middle region of shaft; api- cal shaft part rather short, subapical cavity well marked, fringed plates on thecal apex suberect. Body length 9 mm. Female. — Unknown. Diagnosis. — M. rufitarsis can be distinguished by the combination of lacking abdominal bands and lat- eral spots with the mediolaterally aligned posterior surstyle lobe. Period of flight and distribution (fig. 83). — M. ru- fitarsis is known from Tadzhihistan; the flight period is unknown. Discussion. — M. rufitarsis cannot be considered a subspecies of M. fulcratus Becker as originally estab- lished by Sack (1913a). From their respective descrip- 167 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 tions it is clear that Becker and Sack described two different species, although both were based on infor- mation provided by Loew. Type material belonging to the nominate form of M. fulcratus Sack has not been recovered. Merodon fulcratus Sack, 1913 (pub- lished August, 1913) is a junior homonym of Merodon fulcratus Becker, 1913 (published April, 1913), and should be renamed when its identity has become clear. The male genitalia are very different. For instance, in M. fulcratus Becker the aedeagus is abruptly tapering basally and the surstyle lobes are smaller. M. rufitarsis has completely touching eyes in the male, whereas in males of fulcratus Becker the eyes are widely separated. These species are assigned to dif- ferent species groups, and are not considered closely related. Merodon sophron sp. n. (figs. 35 a-c, 81) Type-material. — 4 Holotype ‘Morocco, Azrou [33° 25° N, 05° 20’ W], 29.v.1925, E. Hartert’ (BMNH). Description Male. — Head: Antennae dark, antenna 3 with slightly convex upper margin, apex acute, antennal ratio 2.0; vertex angle 30°, ocellar angle 60°, tl-v ratio 0.6; pubescence pale yellow, with dark tuft in ocellar region. Thorax: Dark, katepisternum, anepisternum and anepimeron lustrous; pubescence greyish yellow, slightly uneven; no pruinose bands present. Wings clear, orange veined; halteres, squamae and antisqua- mae pale orange. Legs: All dark; trochanters 3 smooth; femora 3 swollen, with normal triangular processus bearing 6 bristles; tibiae 3 and tarsi 3 normal. Abdomen: Dark, comparatively stout; T II-IV with slightly arcuate, narrow, widely interrupted whitish pruinose bands and slight metallic lustre overall; T II with clear metallic lustrous lateral spots bearing rath- er dense, golden pubescence; pubescence otherwise paler, less conspicuous. Genitalia: Anterior surstyle lobe elongate, rectan- gular, markedly curving dorso-anteriorly, spathulate, with dense, yellow, even pubescence apically; posteri- or surstyle lobe separated by small clear sulcus, rather large, very high, apical parts mediolaterally aligned and largely overlapping; the posterior surstyle lobe bears a bald, slightly elongate knob anteriorly; from here a suture runs toward the posterior margin, divid- ing the posterior surstyle lobe in basal and apical parts; the posterior margin bears dense, long, erect, yellow pubescence. Cercus very large with same pu- bescence. Aedeagus smooth on outer face, excavated on this face in hardly lengthened apical shaft part; 168 fringed plates on thecal apex rather large, erect; in natural position aedeagus entirely covered by posteri- or surstyle lobe. Body length 9.5 mm. Female. — Unknown. Diagnosis. — M. sophron is distinguished from oth- er species by its very large overlapping posterior sur- style lobe and its golden pubescence on the metallic lustrous lateral spots on T II. Period of flight and distribution (fig. 81). — M. sophron is known only from the holotype, collected in Morocco in May. Etymology. — The epitheton sophron is derived from the greek words soos, meaning chaste, and phronein, meaning to act, and refers to the ‘chaste’ covering of the aedeagus in this species. A noun in ap- position. Merodon tener Sack (fig. 81) Merodon tener Sack, 1913a: 443. Lectotype 2 (here desig- nated): ‘Sarepta [= Krasnoarmeysk near Volgograd, after Peck 1988]/ M. tener Sack det. Sack/ coll. Lichtwardt/ coll. D. E. I. Eberswalde’ [examined]. Merodon tener, Becker 1921: 54; Van der Goot 1981: 215. Lampetia tenera; Sack 1931: 329. Material examined.-® Paralectotype with same data as lectotype (DEIC). Description The lectotype and paralectotype are quite similar to alexeji females. The main differences are: ocellar ang- le 70° in the tener types, 60° in alexeji, coloration of abdomen reddish throughout in tener and slightly stouter than in alexeji. Lectotype and paralectotype differ slightly in the degree of swelling of the femora 3 but are otherwise similar among themselves and to alexeji. Period of flight and distribution (fig. 81). — M. ten- er is known from the southern and eastern parts of Russia, where it has been collected in June (Sack 19315529); Discussion. — Since no male type-material of tener has been traced it is impossible to ascertain the status of tener as a separate species. In view of the differences stated tener is here considered to be a separate species. In view of the similarity in the females the species is tentatively assigned to the alexeji group. Merodon trizonus (Szilady) Lampetia trizona Szilady, 1940: 67. Syntypes d and 9: La Calle [el Kala], Algeria and Ain Draham, Tunisia [not ex- amined]. Discussion. — The extremely brief description pro- vided by Szilady is insufficient to separate trizonus from other Merodon. The original description ex- pressly states that the species is related to hirsutus Sack. Therefore the species is tentatively assigned to the alexeji group which also includes hirsutus. The types of this species, formerly stored in TMA, have been lost in 1956 (Dr. A. Dely-Draskovits, in litt.). No other material assigned to this species is known. The tarsatus group Apomorphies: Basitarsus 3 strongly swollen, spe- cialized pubescence present on inner face; S IV at least keeled, mostly strongly modified. Description. — Head: Antennae orange to brown, antennal ratio 1.3 to 1.7, compound eyes in male usually touching with tl-v ratio 0.1 to 0.3, pubescence yellow to white, oral margin well marked. Thorax: Dark, metallic lustrous. Wings clear to slightly tinged, orange to dark veined, halteres, squamae and antisquamae yellow. Legs: Femora 3 with normally shaped triangular processus, tibiae 3 in male often bearing apical floccus inserted on distal spur, basitar- si 3 swollen strongly to enormously, less conspicuous- ly so in females; showing short thick-set pubescence on inner face. Abdomen: Tergites metallic lustrous, S IV strongly modified in males of all species except au- ronitens and caudatus where it is only keeled. Genitalia: Anterior surstyle lobe simple, often with short, dense pubescence distally, separated by small suture from higher posterior surstyle lobe bearing long posterior pubescence. Cercus well protruding. Aedeagus with apical part short. Merodon ankylogaster sp. n. (figs. 36 a-b, 84) Type-material. - d Holotype ‘Iran, Fars, Road Chiraz- Kazeroun, Ft. Sine-Sefid [Southwestern Iran, about 29° 35’ N, 52° E, in mountains], 29.iv.1937, coll. F. H. Brandt’ (RMNH). Paratype 9, same data as holotype (RMNH). Description Male. — Head: Antennae orange, antenna 3 with upper margin convex, apex obtuse, antennal ratio 1.3; pubescence pale yellow, dark hairs present in ocellar region; vertex angle 40°, ocellar angle 45°, vertex lar- ge, d-v ratio 0.15. Thorax: Coppery lustrous on katepisternum, ane- pisternum, anepimeron, dorsum and scutellum, dor- Hurkmans: Monograph of Merodon sal surface rather roughly sculptured, showing yellow, erect, uneven pubescence. Wings orange veined, clear. Legs: Dark, tibiae basally yellow; tarsi with dense golden pubescence; trochanters 3 smooth, femora 3 much swollen, slightly metallic lustrous, with normal triangular processus; tibiae 3 without apical floccus; basitarsi 3 strongly swollen, bearing dense, golden, short pubescence medially; other tarsi 3 normally shaped. Abdomen: T II much tapering posteriorly, without lateral spots; pruinose bands interrupted, slightly ar- cuate; pubescence moderately dense, golden, oblique; S IV with conspicuous median projection; abdominal surface roughly sculptured on medial region of all ter- gites. Genitalia: The anterior surstyle lobe elongate, slightly spathulate, apically bearing short, dense, erect, yellow pubescence, posterior surstyle lobe sep- arated from anterior surstyle lobe by shallow sulcus, steeply rising, posteriorly with long dense yellow pu- bescence. Cercus subangular, oblong, with similar pubescence. Aedeagus stout, with fringed plates on thecal apex recumbent, apical part screened by the ap- ical portion of the posterior surstyle lobe in natural position. Body length 7.5 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Ocellar angle 60°; frons lustrous throughout, laterally yellowish metallic lustrous. Thorax: Stouter than in male; sculpturing less strong, pubescence more whitish. Abdomen: Stouter, pruinose bands nearly continuous; pubes- cence whiter; S IV normal. Body length 9 mm. Diagnosis. — The male is unmistakable in view of the morphology of S IV; the female ankylogaster can be separated from turkestanicus by the partially orange legs, and from caudatus by the lack of lateral spots on the abdomen. : Period of flight and distribution (fig. 84). — The species is known only from the types, collected in Iran in April. Discussion. — The shape of the anterior surstyle lobe and inward curve of the posterior surstyle lobe in ankylogaster suggest a certain degree of relationship with alexeji and its allies [note the discussion of mar- ginicornis above]; in view of the modified S IV and the swollen basitarsi 3, ankylogaster has been assigned to the tarsatus group. Etymology. — The epitheton ankylogaster is derived from the Greek ankylos, meaning spine, and gaster, 169 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 meaning belly, hence abdomen, and refers to the pro- jection of S IV in the male of this species. The epithe- ton is to be treated as a noun in apposition. Merodon auronitens sp. n. (figs. 37 a-c, 84) Type-material. — Holotype d ‘Israel, Mt. Carmel, Bat Slomo 32° 35’ N, 35° 00’ E, 22.iii.1981, W. B. M. Brantjes’ (ZMAN). The antennae and the right leg 3 on cardboard tag, both anterior surstyle lobes attached to another cardboard tag. Description Male. — Head: Antennae black, 3rd article with up- per margin concave, apex subacute, antennal ratio 1.7, vertex angle 35°, ocellar angle 40°, tl-v ratio 0.35; pubescence very dense, long, glossy golden on frons, slightly shorter and sparser on face and vertex. Thorax: Dark; golden metallic lustrous laterally, bearing moderately dense, even, golden brown pubes- cence. Wings clear, halteres, squamae and antisqua- mae yellow. Legs: Dark throughout, tibial bases and tarsi slight- ly lighter; trochanters 3 smooth, femora 3 much swol- len, showing normally shaped triangular processus bearing 8 bristles; tibiae 3 show a clear, golden apical floccus; basitarsi 3 swollen, with very dense, short, golden pubescence on inner face. Abdomen: T II slightly tapering, with traces of lat- eral spots; T II-IV with arcuate greyish pruinose bands, that on T II inconspicuous; pubescence rather long, even, yellow, moderately dense; sternite IV keeled, posteriorly slightly emarginate, but not de- formed. Genitalia: Anterior surstyle lobe somewhat elon- gate, with dense orangish pubescence throughout; connection to posterior lobe severed; posterior lobe rather high, curving mediad apically, with scattered yellow pubescence on apical margin; posterior end with dense yellow pubescence. Cercus well protrud- ing, bearing dense, long, yellow pubescence. Aedeagus smooth on outer face, fringed plates on the- cal apex recumbent. Body length 11.5 mm Female. — Unknown. Diagnosis. — M. auronitens can be distinguished from other species in this group by the lack of a de- formed S IV, like caudatus, but differs from that spe- cies by not having a deformed genital cap. Period of flight and distribution (fig. 84). — M. au- ronitens is known only from the holotype, collected in Israel in March. 170 Etymology. — The epitheton auronitens is latin, and derived from the word aurum, meaning, gold, and the verb nitere, meaning, to shine, the construction meaning, shining with gold, as a reference to the glossy pubescence on the head. The epitheton is to be treated as an adjective. Merodon caudatus Sack (figs. 38 a-d, 84) Merodon caudatus Sack, 1913a: 446. Holotype: Asia Minor [Turkey] [not examined, considered lost]. Merodon caudatus, Paramonov 1926b: 322; Peck 1988: 168. Lampetia caudata; Sack 1931: 309. Material examined. — Israel: 26, 12 ‘Mt. Carmel, Beth Slomo, 22.3.1981 leg. Brantjes’ (LR); 12 ‘Palestine, Beth Hakerem [near Jerusalem], 17.3.1951 leg. O. Theodor’ (RMNH). Description Male. — Head: Antennae dark, antennal ratio 1.8, antenna 3 with apex obtuse, somewhat edged; vertex large, metallic lustrous, vertex angle 50°, ocellar ang- le 65°, pubescence yellow, whitish on face and occipi- tal sides; compound eyes showing grey pubescence, tl-v ratio 0.2-0.25. Thorax: Dark, brassy lustrous on katepisternum, anepisternum and anepimeron, dorsum and scutel- lum, showing dense, erect, yellowish, even pubes- cence, diffusely pruinose on lateral dorsum, scutellum and sides. Wings quite clear, halteres, squamae and antisquamae pale yellow. Legs: All femora dark, apically paler, tibiae yellow with dark distal band, tarsi orange to brown, apical articles darkest; pubescence dense on tibiae and tarsi, yellow underneath; trochanters 3 smooth, femora 3 rather straight, triangular processus somewhat re- duced, forming only slight knob; tibiae 3 curved and twisted, bearing apical floccus on drawn-out distal medial spur; basitarsi 3 short, swollen, with furrow giving twisted aspect, bearing short, even, dense, yel- low pubescence; other tarsi 3 normal. Abdomen: Tergites dark, metallic lustrous, T II with orange lateral spots and erect, orange pubescence (densest and longest on lateral spots); pruinose bands arcuate, interrupted on T III, continuous on T IV; S normal but S VIII (the genital cap) strongly modified, elongate, with apical floccus conspicuous to naked eye (hence the name caudatus). Genitalia: The anterior and posterior surstyle lobes separated by deep wide sulcus; anterior surstyle lobe rounded-triangular, with short, even, dense, yellow pubescence, showing ridge running from ventro-pos- terior angle to mid region; posterior surstyle lobe short and high, with longish, uneven, yellow pubes- cence apically. Cercus well protruding, angular, with very long, yellow pubescence. Aedeagus with short apical shaft part, the fringed plates on thecal apex re- cumbent. Body length 12 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Ocellar angle 70°, frons metallic lustrous, diffusely pruinose on sides; frons clearly tapering toward vertex. Thorax: Markedly stouter than in male. Legs: Slenderer than in males, basitarsi 3 quite similarly modified but pu- bescence duller. Abdomen: Stouter, rather triangular, with conspicuous lateral spots bearing fiery orange hair-tufts; pruinose band on T III nearly continuous. Body length 9.5 mm. Diagnosis. — The males are easily separated by their unique genital cap; the females are similar to those of ruficornis Meigen and allied species from which they are separated by the swollen basitarsi 3. Period of flight and distribution (fig. 84). — The species is known from Israel and Turkey where it oc- curs in March. Discussion. — In spite of the fact that no type-ma- terial was available, the identification of this species is certain in view of the extraordinary features present. The assignment of the female of caudatus as present- ed here seems justified in view of the striking similar- ities (basitarsi 3, lateral spots on T II) and the concur- rent flight in the same area. Merodon oidipous sp. n. (figs. 39 a-d, 84). Type-material. — dé Holotype ‘Turkey, Hakkari, Suvarihalil pass 2100 m, 4.vi.1984, leg. J. A. W. Lucas’ (ZMAN). Paratypes: 2d , same data as holotype (JLR). Description Male. — Head: Antennae orange to brown, anten- nae III paler than I-II, upper margin straight to slight- ly concave, apex rounded, antennal ratio 1.6; vertex angle = 40°, ocellar angle = 60°, tl-v ratio 0.25; face and occiput slightly, vertex strongly blue metallic lu- strous; pubescence whitish to pale yellow, face more- over diffusely pruinose; compound eyes with greyish pubescence. Thorax: Dark, with some steel blue lustre on later- al dorsum and scutellum, stronger on sides; pubes- cence erect, moderately dense, rather short, uneven. Wings clear; halteres with knob yellow, stalk darker, squamae and antisquamae yellow. Legs: All femora dark with narrow yellow apices, tibiae yellow with broad dark distal band, tarsi dark Hurkmans: Monograph of Merodon above, with dense pale pubescence underneath, upper faces orangeish to brown on legs 1-2, darker on 3; tro- chanter 3 bearing low projection with yellow hair- tuft; femur 3 swollen, bluish metallic lustrous; trian- gular processus rather large, with conspicuous apical spur, otherwise finely serrate, bearing some 9 bristles; tibiae 3 with long yellow apical floccus inserted on drawn-out distal inner angle; basitarsi 3 enormously swollen, almost globular, brownish, with dense gold- en short erect pubescence medially; other tarsi 3 nor- mal. Abdomen: Rather slender, dark; T II tapering pos- teriorly, T I-IV bearing slightly arcuate moderately wide interrupted pruinose bands; pubescence whit- ish, rather sparse, slightly denser laterally and on prui- nose bands, darker medially; at certain angles vague metallic blue lustrous lateral spots on T II may be seen; S IV with paired, conspicuous appendages. Genitalia: The anterior surstyle lobe somewhat spathulate, bearing dense, yellow, even, erect pubes- cence on distal margin; separated from the slightly el- ongate rounded posterior surstyle lobe by a shallow sulcus; posterior surstyle lobe incised dorsally, bearing dense, erect, yellow pubescence. Cercus conspicuous, elongate, with same pubescence. Aedeagus rather slender with short apical shaft part, fringed plates on thecal apex recumbent to suberect. Body length 8.5 mm. Female. — Unknown. Diagnosis. — M. oidipous can be separated by its enormously swollen basitarsi 3 which are differently shaped than in other species of the tarsatus group, by the clear blue lustre on the vertex, and the more pro- truding cercus. Period of flight and distribution (fig. 84). — The species, known only from the types, has been found in south-eastern Turkey in June. Etymology. — The epitheton oidipous derives from the Greek and means, swollen foot. It refers to the enormously swollen basitarsi 3, conspicuous to the naked eye. Merodon persicus sp. n. (figs. 40 a-d, 85) Type-material. — d Holotype ‘Iran, Fars, Ft. Sine-Sefid, Rd. Chiraz Kazeroun [29° 35’ N, 52° E], Coll. F. H. Brandt 25.v.1937 / Lampetia smirnovi det. P. H. van Doesburg’ (RMNH). Description Male. — Head: Antennae brown, antenna 3 orange, 1721 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 upper margin straight to slightly concave, apex acute, antennal ratio 2.3; pubescence pale yellow, with dark hair-tuft in ocellar region; oral margin more protru- ding than in the other species in this group; face slightly diffusely pruinose; vertex angle 40°, ocellar angle 60°; tl-v ratio 0.25; compound eyes with whi- tish pubescence. Thorax: Rather dull, surface dark brown, bearing uneven, short, erect yellow pubescence on dorsum, scutellum, katepisternum, anepisternum and anepim- eron, with weak bronze lustre on these elements. Wings clear, halteres, squamae and antisquamae yel- low. Legs: Dark but bases of tibiae and tarsi brown; tro- chanter 3 showing well-defined low hump bearing rather short black bristly pubescence; femur 3 swol- len, curved, with rather small triangular processus, strongly serrate on distal margin; tibiae 3 short, stout, with distinct apical floccus on distal medial angle; basitarsi 3 strongly swollen, showing dense, golden, short, erect pubescence throughout (densest on medi- al face). A slightly elevated, well defined area proxi- mally on lower face bearing erect, dense dark bristly short pubescence obliterating the surface of this area; other tarsi 3 normal. Abdomen: Moderately slender, all dark, with rath- er strong olivaceous metallic lustre; vaguely defined, arcuate pruinose bands present on T III-IV; pubes- cence golden throughout, some dark hairs present in posteromedian parts of T II-III. All S rectangular, S IV bearing conspicuous, paired appendages visible to naked eye. Genitalia: The anterior surstyle lobe elongate, api- cal part widening and projecting dorsally, showing short, yellow, erect pubescence along ventral and dis- tal margins; separated from posterior surstyle lobe by shallow suture, from which the rounded-rectangular posterior surstyle lobe steeply rises; posterior surstyle lobe with yellow, scattered, erect pubescence, longest on ventral and posterior margins. Cercus largish, bearing dense, long, yellow, erect pubescence. Aedeagus stout, fringed plates on thecal apex recum- bent, apical shaft part short; in natural position the apical portion of the aedeagus is covered by the poste- rior surstyle lobe tops. Body length 9.5 mm. Female. — Unknown. Diagnosis. — M. persicus seems to be most closely related to tarsatus, from which it differs in the smaller size, the entire basitarsi 3 with a small raised area (as opposed to excavated basitarsi 3 with large area in tar- satus), the elongate posterior surstyle lobe with dorsal- ly projecting apical part, and the stronger inward curve of the apical posterior surstyle lobe region. M. 172 persicus differs from the description of smirnovi in the smaller size, the smaller tl-v ratio and the colour of the antennae. Period of flight and distribution (fig. 85). — The species is known to occur in Iran; it flies in May. Etymology. — The epitheton persicus is derived from the classical name Persia, indicating the region of occurrence of the species. Merodon tangerensis sp. n. (figs. 41 a-d, 87) Type-material. — d Holotype ‘Environs de Tanger [Morocco], Fevrier 1859’ (MNHN). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apex rounded, antennal ratio 3.3; pubescence pale yellow with dark hair-tuft in ocellar region; vertex angle 30°, ocellar angle 35°, touchline incomplete, tl-v ratio 0.15; face slightly, diffusely pruinose, occiput densely pruinose down sides, me- tallic lustrous; compound eyes showing whitish pu- bescence. Thorax: Dark, slightly metallic lustrous on lateral dorsum and scutellum; katepisternum, anepisternum and anepimeron steel grey lustrous; dorsum with double paired anteriorly coalescent whitish pruinose bands; pubescence erect, yellow, moderately dense, distinctly uneven throughout, paler on sides. Wings clear, pale brown veined; halteres, squamae and anti- squamae yellow. Legs: Dark but apices of femora and bases and api- ces of tibiae lighter; trochanter 3 smooth, femur 3 swollen, black metallic lustrous, bearing normal tri- angular processus holding 6 bristles on serrate distal margin; tibiae normal, showing drawn-out distal me- dial angle bearing dense somewhat lengthened pubes- cence (this might be termed a subfloccus); basitarsi 3 swollen, with dense, yellowish, erect pubescence (densest medially); other tarsi 3 normal. Abdomen: Dark, slender, tergites rectangular, slightly transverse, with white, slightly arcuate mod- erately wide pruinose bands, interrupted on T II, continuous on T III-IV; pubescence short, white on bands, dark elsewhere; S IV strongly keeled, deeply emarginate posteriorly. Genitalia: The anterior surstyle lobe elongate, with apical part bearing short, even, yellow, dense pubes- cence, more scattered elsewhere; separated from pos- terior surstyle lobe by deep sulcus; posterior surstyle lobe high, apically curving medially; with marked an- tero-ventral angle, posteriorly rounded, there with moderately dense rather long yellow pubescence. Cercus inconspicuous, bearing long dense yellow pu- bescence. Aedeagus slender, fringed plates recum- bent, subapical cavity well marked. Body length 8.5 mm. Female. — Unknown. Diagnosis. — M. tangerensis is distinguished from other species in this group by its very long antenna 3, the relatively small projection of S IV and the relative- ly large somewhat dorsally curving anterior surstyle lobe. Period of flight and distribution (fig. 87). — This species is known only from the holotype, collected in northern Morocco in February. Etymology. — The species is named after the site where the holotype was collected. The epitheton is to be used as an adjective. Merodon tarsatus Sack (figs. 42 a-e, 83). Merodon tarsatus Sack, 1913a: 437. Holotype d: ‘Pamir 49409/ Mer. tarsatus Sack det. Sack’ (ZMHB) [examined]. Merodon smirnovi Paramonov, 1926b: 320. Holotype d: ‘Ak-Tash, Prope Tashkent, Turkestan’ [not examined, see discussion]. Syn. nov. Merodon tarsatus, Paramonov 1926b: 320; Paramonov 1927: 76, 78; Peck 1988: 174. Lampetia tarsata, Sack 1931: 328. Merodon smirnovi, Paramonov 1927: 76, 78; Liepa 1969: 20; Peck 1988: 173. Lampetia smirnovi, Sack 1931: 326. Material examined. — U. S. S. R.: 16 ‘Ak-Tash [41° 37° N, 69° 40° E], 9 vii 1923, Tashkent A.Zecholovtsev/ Lampetia smirnovi Param., S. Paramonov det. mm/ coll. J. Villeneuve’ (KBIN); 12 ‘Berkana, Turkestan, 24.vi.1925, Kuznetsov leg.” (ZMAS); 19 ‘Outsj. Koydara, dol. R. Warzob, Tadzj. [river Warzob valley in Tadzhikistan, North of Dushanbe in 68° 50’ E, between 38° 20’ and 38° 55’ N] (ZMAS). Description Male. — Head: Antennae brown, antenna 3 with upper margin edged, straight, apex subacute, anten- nal ratio 1.9; pubescence yellow, with dark hair-tuft in ocellar region; vertex angle 40°, ocellar angle 70°; d-v ratio 0.3; face and occiput metallic greenish lus- trous; central ocellus only half as large as the lateral ones; compound eyes with whitish pubescence. Thorax: Strongly lustrous on lateral dorsum and scutellum, katepisternum, anepisternum and anepim- eron; dorsum and scutellum with diffuse golden pu- bescence and scattered pruinosity, with 4 faint prui- nose bands also showing. Wings tinged brown on central portions, clear marginally, veins very dark; Hurkmans: Monograph of Merodon halteres, squamae and antisquamae yellow. Legs: All joints and tibial bases yellow, otherwise dark; trochanter 3 sharply keeled, femur 3 swollen, metallic lustrous, triangular processus normal, bear- ing 6-8 bristles on serrate distal margin; tibiae 3 rath- er strongly curved, bearing large lamella on distal me- dial face with ruddy floccus inserted apically; floccus extends over half the length of basitarsi 3; these swol- len, asymmetrical, with lower face showing large flat metallic lustrous area bearing stout bristle-like pubes- cence, bordered by very stout long pubescence; inner face of basitarsi 3 showing dense orange-yellow pu- bescence. Abdomen: Dark, greenish yellowish metallic lus- trous, rather slender; pubescence dense, moderately long, pale orange, partially obliterating roughly punc- tated surface; no pruinose bands; S IV modified, bearing large, slender paired appendages. Genitalia: The anterior and posterior surstyle lobes separated by shallow sulcus, of about equal size; ante- rior surstyle lobe apically with short, dense, erect, yel- low pubescence, of rectangular outline; posterior sur- style lobe moderately high, regularly rounded, bearing erect, long, yellow, uneven pubescence. Cer- cus of moderate size, with similar pubescence, slight- ly angular. Aedeagus robust, heavily sclerotized, api- cal shaft part short, fringed plates recumbent. Body length 10.5-13 mm. Female. — Rather different from the males, as not- ed by e.g. Sack, 1931; therefore a full description is given. Head: Antenna 3 dark orange, subacute; antennal ratio 2-2.3; pubescence whitish; ocellar angle 70°; the central ocellus clearly smaller than the lateral ones; frons diffusely pruinose laterally, lustrous for median one-third; metallic lustre absent. Thorax: Weakly greenish metallic lustrous on later- al dorsum and scutellum, katepisternum, anepister- num and anepimeron; pubescence rather short, erect, even, pale yellow. Wings as in male, slightly less tinged. Legs: All femora dark, tibiae orange with dark dis- tal band, femur 3 slightly swollen, triangular proces- sus bearing 6 bristles on serrate distal margin; tibiae 3 short, otherwise normal; basitarsi 3 clearly swollen, not deformed, bearing dense golden pubescence; other tarsi 3 normal. Abdomen: Dark, slightly metallic lustrous; prui- nose bands on T II-IV narrow, whitish, arcuate, well defined, interrupted; pubescence yellow to white, deepest coloured and densest laterally on T II. Body length 12 mm. Diagnosis. — M. tarsatus can be distinguished in the males, by its deformed basitarsi 3 with the lustrous 173 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 area at the lower face which at once separates it from all other species; in the females, by the clearly swollen basitarsi 3 and the great antennal ratio. Period of flight and distribution (fig. 83). — M. tar- satus occurs in Tadzhikistan, and adjacent regions, where it flies in June and July. Discussion. — In the original description the colour of the pubescence of the holotype is said to be foxy- red, its size as 12 mm; the colour has apparently faded, while the specimen was measured recently at 10.5 mm only. The synonymy of smirnovi with tarsa- tus was established after examination of one of the specimens (originating from the type-locality of smir- novi) identified as smirnovi by Paramonov, and matching the description as well as the holotype of tarsatus. Moreover, the females, all identified previ- ously as smirnovi, match the description of the © tar- satus as provided by Sack. Furthermore Sack (1931) noted the specialized setaceous area on the basitarsus 3 of smirnovi which he failed to observe in tarsatus, but which is easily seen on the holotype of tarsatus. Sack gave a difference in the tl-v ratios between tarsa- tus and smirnovi (the ratio of tarsatus being greater) in his key to the species (Sack, 1931), but checking of the holotype of tarsatus showed that the tl-v ratio in that species is about 0.3, whereas the ratio given by Paramonov for smirnovi is 0.5. Therefore Sack’ s interpretation that two species are involved is not fol- lowed here; in view of the above arguments smirnovi is considered synonymous with tarsatus. Merodon turkestanicus Paramonov (figs. 43 a-e, 86) Merodon turkestanicus Paramonov, 1926b: 319. Holotype à : ‘Tashkent, Tadzhikistan, 12.v.1915’ [U. S. S. R.] [not ex- amined]. Merodon turkestanicus, Paramonov 1927: 75; Liepa 1969: 20; Peck 1988: 175. Lampetia turkestanica; Sack 1931: 330. Material examined. — Iran: 1d ‘Kuh-e-Hazaran, S. Rayen Kerman [29° 24’ N, 57° 22’ E], 3800 m, 25.v.1978 leg. K. Warncke’ (LR); U. S. S. R.: 24 ‘Outsj. Koydara, dol. R. Warzob Tadzj. [68° 50’ E, 38° 20’ to 38° 50’ N], Warzob river, Tadzhikistan ] Stackelberg 6.v.43’ (ZMAS); two 9 from same locality and collector, dated 2.v.44 (ZMAS, BMNH); 19 ‘Outsj. Kwak. Werch. Koydarye Tadzh. [in same region as holotype] Stackelberg 27.v.43’ (RMNH); 1d ‘Hissar mountains Tadzhikistan [same region], Stackelberg 3.v.44’ (RMNH). Description Male. — Head: Antennae dark, antenna 3 with up- per margin straight, apex obtuse, antennal ratio 2.1; 174 vertex conspicuously large; vertex angle 45°, ocellar angle 50°, ocelli large, tl-v ratio 0.15, touchline in- complete; pubescence pale yellow, dark in ocellar re- gion, on compound eyes sparse, yellowish. Thorax: Moderately strong metallic lustrous on lat- eral dorsum, scutellum and sides, background colour dark; pubescence rather sparse, uneven, pale yellow. Wings clear throughout; halteres, squamae and anti- squamae pale yellow. Legs: All dark, tibiae and tarsi 1-2 with dense yel- low pubescence giving false impression of light colour; trochanter 3 with small rounded knob, femur 3 swollen, strongly metallic lustrous, with triangular processus strongly serrate apically, tibiae 3 with con- spicuous apical spur bearing strong floccus; basitarsi 3 very much swollen, bearing dense, short, yellow, even pubescence medially; other tarsi 3 normal. Abdomen: Outline rather stout, background colour dark, obliterated by strong metallic lustre and moderately dense, yellowish, even pubescence; no traces of pruinosity or lateral spots. S IV bears con- spicuous paired appendages. Genitalia: Anterior and posterior surstyle lobes sep- arated by deep narrow sulcus; anterior surstyle lobe bearing rather dense, yellow, erect, even pubescence apically; posterior surstyle lobe showing scattered, long, yellow pubescence, densest posteriorly. Cercus rectangular, well protruding, bearing long, erect, yel- low pubescence. Aedeagus stout, apical shaft part short, fringed plates on thecal apex recumbent. Body length 9 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Eyes separate widely, ocellar angle 55°, frons lustrous throughout, partially dulled laterally by light pruinosity. Legs: The basitar- si 3 slightly but distinctly swollen, with a length/ width ratio about 3.0. Abdomen: Appendages on S IV lacking. Body length 9 mm. Diagnosis. — The males of turkestanicus can be dis- tinguished from other species in this group by the large vertex, all dark legs, structure of tibiae 3 and basitarsi 3 and strong metallic lustre; the females by the swollen basitarsi 3 and dark legs. Period of flight and distribution (fig. 86). — M. turkestanicus occurs in Tadzhikistan and Turkestan, and Iran, in mountain areas, flying there during May and June. Discussion. — The identification of the specimens, some by Paramonov, seems to be correct; all speci- mens match the original description. Merodon xanthipous sp. n. (figs. 44 a-c, 85) Type-material. — & Holotype ‘Iran, Chiraz, 11.iv.1937, coll. F. H. Brandt / L. crassicornis Sack, det. v. Doesburg’ (ZMAN). 19 Paratype ‘Iran, Rd. Chiraz Kazeroun, Ft. Sine- Sefid, 19.iv.1937, coll. F. H. Brandt / L. crassicornis Sack, det. v. Doesburg’ (ZMAN) [both localities at about 29° 30° N, 52° E]. Description Male. — Head: Antennae brown, antenna 3 orange, upper margin slightly concave, apex acute, antennal ratio 2.1; vertex angle = 35°, ocellar angle = 45°; pu- bescence pale yellow, with hardly any dark pubescen- ce in ocellar region; tl-v ratio 0.15, touchline incom- plete; head without metallic lustre. Thorax: Dark, lacking metallic lustre; katepister- num, anepisternum and anepimeron with some dull- ish blue tinge; pubescence sparse, brown, uneven, moderately long; some diffuse pruinosity present on sutures; vestigial pruinose bands can be seen at certain angles on posterior dorsum. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: All pale brown; trochanter 3 with low knob bearing yellow hair-tuft, femur 3 slightly curved, swollen, with rather high triangular processus show- ing apical spur, bearing one strong and 5-6 lesser bris- tles; tibiae 3 with apical floccus reaching down basal half of basitarsi 3, inserted on drawn-out medial dis- tal angle; basitarsi 3 swollen, bearing rather dense, brownish yellow, erect, short, even pubescence throughout, densest on medial face; other tarsi nor- mal. Abdomen: Very dark brownish red, tapering poste- riorly, especially T II; slightly arcuate, faint, narrowly interrupted whitish pruinose bands present on T II- IV; pubescence pale brown, erect, uneven; S IV bear- ing robust large paired appendages that are smaller than in other species of this group, but still very con- spicuous. Genitalia: The anterior surstyle lobe rather spathu- late, with dense, yellow, erect, even pubescence dor- sally and apically, more scattered on other parts; sep- arated from posterior surstyle lobe by shallow sulcus; posterior surstyle lobe steeply rising, anteriorly high, with distinct slight anteroventral angle and rather dense, erect, yellow, uneven pubescence posteriorly; cercus somewhat elongate, rather wide, with dense, long, yellow pubescence; aedeagus moderately slen- der, with marked subapical cavity, fringed plates on thecal apex suberect, apical shaft part slightly length- ened. Body length 7 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Antenna 3 rounded, Hurkmans: Monograph of Merodon more obtuse; frons lustrous (not metallic) through- out, eyes widely separate, ocellar angle 60°. Thorax: Showing dull bluish-greenish lustre on dorsum, scu- tellum and sides. Legs: Deeper brown than in male. Abdomen: Stouter than in males, darker, pruinose bands more marked; T II-V with a micaceous lustre. Body length 7 mm. Diagnosis. — The species can be distinguished in both sexes by the relatively pale, uniformly coloured legs; the males are also distinguished by the relatively smaller appendages on S IV and the steeply rising posterior surstyle lobe as well as by the absence of me- tallic lustre. The former identification as crassicornis Sack, pre- viously applied to the holotype and paratype, is incor- rect in view of contradictions of characters in the types of xanthipous with the description of crassicor- nis. In the females of the latter species the abdominal pruinose bands are stated to be wide and clear, the an- tennae are dark, the compound eyes bearing very dense pubescence, the legs are black with yellow markings; moreover bands of long white pubescence are stated to be present on the posterior tergite mar- gins. The male of crassicornis has not been described. Period of flight and distribution (fig. 85). — The species, known only from the types, occurs in south- western Iran in April. Etymology. — The epitheton xanthipous, to be treated as a noun in apposition, refers to the golden brown leg coloration, and is derived from the greek xanthos, meaning blonde, and pous, meaning foot. The clavipes group Apomorphies: Anterior surstyle lobe elongate, den- sely pubescent and strongly to enormously enlarged. Description. — Head: Antennae with antennal ratio 1.4 to 2.0, antenna 3 mostly subacute; pubescence on head whitish to yellow, usually dense on frons; face often bluish lustrous; tl-v ratio in the males 0.3 to 0.5. Thorax: Dark, lustrous on dorsum and scutellum, mostly with strong blue lustre on katepisternum, an- episternum and anepimeron; pubescence most vari- able in colour density and length; some species have slightly to strongly infuscate wings. Legs: Mostly all dark, in some cases tibiae 1 and 2 basally with orange markings; femur 3 usually much swollen, curved, in some cases straight and slender; triangular processus, tibiae 3 and tarsi 3 normally shaped. Abdomen: Moderately slender to slender or stout; T II tapering (less strongly in the females); arcuate pruinose bands present on T II-IV which may in some species be dif- ficult to see through the dense abdominal pubes- 175 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 cence. Genitalia: The anterior surstyle lobe very large, rounded-rectangular, ridged laterally, usually with short even pubescence throughout; posterior surstyle lobe variously shaped, with dense pubescence, with specialized hair-tufts in warnckei, aberrans, hamifer and brevis, cercus variable; aedeagus mostly slender, apical shaft part short to slightly lengthened, fringed plates recumbent to oblique. Merodon aberrans aberrans Egger (figs. 45 a-d, 50 b, 89) Merodon aberrans Egger, 1860: 10. Lectotype d (here desig- nated): ‘Austria Alte Sammlung/ Merodon aberrans Egger’ (NHMW) [examined]. Merodon obscuripennis Palma, 1863: 47. [No type material mentioned in original description]. Merodon kneri Mik, 1867: 415. Syntypes: “Halicia’ [not ex- amined]. ; Merodon aberrans, Mik 1867: 417, 1883: 182; Bezzi 1900: 89; Sack 1913a: 435; Paramonov 1925: 153; Liepa 1969: 20; Gaunitz 1969: 82, 86; Van der Goot 1981: 215, 218; Peck 1988: 166. Lampetia aberrans, Oldenberg 1919: 388; Sack 1931, 302; Séguy 1961: 174. Merodon obscuripennis, Peck 1988: 166. Merodon knerit; Mik 1883: 182; Peck 1988: 166. Lampetia kneri; [sic!] Sack 1931: 302. Material examined. — Albania: 8¢ (NHMW, RMNH); Austria: 86 paralectotypes and 2? paralectotypes of aber- rans, with same label as lectotype; Czechoslovakia: 19 (ZMAN); France: 476, 89 (MNHN); Greece: 236, 119 (KBIN, BMNH, JLR, ZMUC, WH, RMNH, ZMAN); Hungary: 3d (NHMW, RMNH); Italy: 646, 462 (NHMW, BMNH, JLR, ZMAN); Lebanon: 1d, 1% (KBIN, NHMW); Poland: 14, 19 (BMNH); Romania: 2d (BMNH); Turkey: 1366, 459 (BMNH, JLR, WH); U. S. S. R.: 156, 142 (NHMW, ZMAS); Yugoslavia: 236 , 59 (NHMW, JLR, BMNH, ZMAN, RMNH). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apex obtuse to subacute, antennal ratio 1.6; pubescence yellow, with dark tuft in ocellar region; vertex angle 40°, ocellar angle 50°, tl-v ratio 0.4; compound eyes showing white pubescence; face variably bluish lustrous, mostly only weakly. Thorax: Blue lustrous on lateral dorsum and scutel- lum, katepisternum, anepisternum and anepimeron; pubescence yellow, moderately dense, with dark interalar band mostly clear. Wings troubled evenly in distal half, anterior veins pale; halteres, squamae and antisquamae pale yellow. Legs: All dark, all femora slender; femur 3 straight; trochanter 3 smooth, femur 3 bearing normally shaped triangular processus bearing 6-11 bristles; bas- itarsi 3, tibiae 3 and tarsi 3 normal. in most specimens tibiae and tarsi with bright yellow pubescence. Abdomen: Dark, with characteristic rather strong bluish overall lustre; T II strongly tapering; T II-IV 176 with slightly arcuate, well interrupted pruinose bands; vague mainly yellow pruinose lateral spots on T II present in some specimens; S IV moderately arched, deeply, narrowly emarginate posteriorly. Genitalia: Anterior surstyle lobe large, anteriorly bearing dense, short, even, yellow pubescence, with scattered pubescence throughout, showing lateral ridge; coalescent with posterior surstyle lobe; the lat- ter bearing low anteroventral projection with dense, long, yellow pubescence; posterior surstyle lobe with long, dense, yellow pubescence. Cercus conspicuous, rather quadrate with rounded corners. Aedeagus smooth on outer face, moderately slender, apical shaft part short, fringed plates on thecal apex recumbent. Body length 12-16 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Antenna 3 more ob- tuse; frons laterally pruinose, lustrous on median one- third or just over; ocellar angle = 40°, pubescence on frons less dense. Thorax: Pubescence brownish, inter- alar band less clear; vague longitudinal pruinose bands often present; wings in some specimens more clearly infuscate distally. Abdomen: T Il less tapering, pubescence very dense in some specimens; T V rela- tively narrow, posteriorly more or less acute. Body length 12-15 mm. Diagnosis. — As the slender femora 3 are not found in other species and the blue lustre is uncommon in a species of this large size, aberrans is hardly ever con- fused with other species; only hamifer is very similar but can be separated at once by its deformed triangu- lar processus; brevis is superficially similar but stouter, lacks the interalar band in the males which moreover have slightly different genitalia (see fig. 45); splendens has a wider, shorter abdomen, denser pubescence and the femora 3 are much more swollen than in aber- rans. Period of flight and distribution (fig. 89). — The species occurs from May through July in central Europe and the Mediterranean, in southern parts of Russia, Ukraine and in Asia Minor. Merodon aberrans ssp. flavitibius Paramonov Merodon aberrans ssp. flavitibius Paramonov, 1925: 153. Syntypes: ‘Armenia’ [not examined]. Merodon aberrans ssp. flavitibius, Liepa 1969: 20; Peck 1988: 166. Lampetia aberrans ssp. flavitibius, Sack 1931: 302. Description Subspecies flavitibius differs from the nominate subspecies only in the colour of the legs, which in the present subspecies show yellow markings on the basal parts of tibiae 1 and 2. Period of flight and distribution. — This subspecies occurs in Armenia. Merodon aberrans isperensis ssp. n. (fig. 90) Type-material. — d Holotype “Turkiye, Rize, Ovit Pass [near Ispir], S side 1800 m, 16.vii.1987, leg. J. A. W. Lucas’ (ZMAN). 436 , 249 paratypes, all from Turkey, all in (LR) and all leg. Lucas, except where noted: Topotypic 19, 31.vii.1983; 1d, 19, 16.vii.1986; 15 ‘Turkey Hakkari, Suvarihalil pass 1250 m W side nr. Habul Deresi 13.vi.1984’; 49 Turkey Hakkari, Tanin-Tanin pass W side 1700 m, 12.vi.1984; 12 ‘Turkey, Kars, Handere 2100 2200 m, 20 km W of Sarıkamıs, 1.vili.1983’; 26, 19 ‘Turkey, Siirt 21 km W of Uludere, 12.vi.1984’; 3d ‘Turkey, Kars, 25 km W. of Sarikamis 5.viii.1985, 2100 m, C. J. Zwakhals leg.’; 36 “Turkey, Kars, Bahnstat. Soganli W. Sarıkamıs 2100 m 5.vii.1985 W. Schacht’; 19 “Turkey, Hakkari, Sat Dagi, Varegös, SW Yiiksekova 1700 m 29.vi.1985 W. Schacht’; 134, 49 ‘Turkey, Bingöl, Buglan Gegidi 1640 m 5.vii.1986’; 16, 19 “Turkey, Kars 11 km E of Karakurt 1450 m 6.vii.1986; 6, 29 “Turkey, Kars, 8 km W Sarikamis, 2000 2050 m 6-9.vii.1986’; 2d, 19 “Turkey, Kars, Handere 20 km W Sarıkamıs 2000 m 7.vii.1986; 18,1% “Turkey, Kars, 12 km S Sarikamis 2050 m, 7.vii.1986; 106, 62 ‘Turkey, Erzurum (Kayak) Palandöken Dagi 2200-2300 m 1.vii.1986’. See map show- ing Turkish localities (fig. 79). Description Male. — Head: Pubescence more grey than in no- minate subspecies, otherwise similar. Thorax: Slenderer than in nominate subspecies, bearing less dense pubescence; interalar band less clear. Legs: Similar to nominate subspecies, but pubes- cence pale yellow, not golden. Abdomen: Clearly less wide than in nominate sub- species, showing slate grey instead of bluish lustre; pubescence slightly less dense. Genitalia: completely similar to nominate subspe- cies. Body length 10-15 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Thorax: The infuscation of the apical wing portions is less strong than in females of the nominate subspecies. Period of flight and distribution (fig. 90). — This subspecies occurs in June through August in the mountains of eastern Turkey. The distributions of the nominate form of aberrans and of the ssp. isperen- sis are shown in figs. 89-90. Hurkmans: Monograph of Merodon Etymology. — The adjective isperensis refers to the type-locality in the vicinity of the town of Ispir in north-eastern Turkey. Merodon brevis Paramonov (figs. 46 a-c, 86) Merodon brevis Paramonov, 1925: 157. Holotype d: ‘Inaklu, Bez. Etschmiadzin, Gouv. Erivan, (Armenien), Siidlicher Abhang des Alagéz, 4500’ [Gora Aragac, 40° 29’ E, 44° 12’ E]’ [Armenia] [not examined, considered lost]. Merodon brevis, Peck 1988: 168. Lampetia brevis, Sack 1931: 308. Material examined. — U. S. S. R.: 1¢, Armenia, Abaran [Aparan, 40° 34’ N, 44° 21 E], M. Karny-larich, 8000’, 10.vii.26. A. Shelk / Lampetia brevis Param. d, S. Paramonoy det./ Coll. J. Villeneuve, Lampetia brevis Param. (KBIN). Description Male. — Head: Antenna with articles 1-2 brown, 3 conspicuously orange, with upper margin convex, ap- ex subacute, antennal ratio 1.7; pubescence white, long, dense on frons, sparser on face, golden on occi- put, darker on vertex; face blue lustrous, partially with dense, pale pruinosity, vertex angle 35°, ocellar angle 50°, tl-v ratio 0.5. Thorax: Dark, lateral dorsum and scutellum, kate- pisternum, anepisternum and anepimeron greenish lustrous, showing rather dense, golden, erect pubes- cence; two faint longitudinal pruinose bands present on dorsum; no interalar band. Wings rather clear, very pale brown veined; halteres, squamae and anti- squamae yellow, halteres deepest coloured. Legs: Dark, but tibiae 1 and 2 orange on basal half. Trochanters 3 smooth, femora 3 swollen, steel grey lustrous, bearing rather large triangular processus with strongly serrate distal margin bearing 5-6 bris- tles; tibiae and tarsi normally shaped, tarsi 3 with golden pubescence laterally, dark on upper face. Abdomen: Relatively small, rather stout; dark, with interrupted well-defined arcuate interrupted pruinose bands; pubescence pale anterior and dark posterior, longest laterally; no lateral spots; S dark, S IV vaulted, moderately deep emarginate posteriorly. Genitalia: Anterior surstyle lobe elongate, recurved dorsally, apically wide, stalked, with clear lateral ridge, with dense, short, erect, yellow pubescence throughout, bearing small dorsal basal projection; posterior surstyle lobe rather quadrate, two more scle- rotized areas on ventral margin bearing dark bristle- like pubescence; pubescence otherwise as on anterior surstyle lobe but longer laterally. Cercus rather small, bearing short, dense, erect, yellow, uneven pubes- cence. Aedeagus moderately slender, apical shaft part short, fringed plates recumbent. 177 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Body length 12 mm. Female. — Unknown. Diagnosis. — M. brevis most closely resembles aber- rans from which it is distinguished by the shorter body, more elongate anterior surstyle lobe and uni- formly coloured thoracic pubescence; the areas of spe- cialized pubescence on the posterior surstyle lobe are not found in aberrans. Moreover, this species differs from aberrans by the orange markings on the legs. In this respect brevis resembles aberrans ssp. flavitibius. No material of the latter taxon has been examined, but the other characters mentioned should be suffi- cient to distinguish these taxa. Period of flight and distribution (fig. 86). — M. bre- vis has been found in the mountains of Armenia [ap- prox. 40° 30° N, 44° 20° E], where it was collected in July. Discussion. — The identification as brevis, by Paramonov, is considered reliable; the specimen ex- amined originates from a site less than 20 km from the locus typicus. Merodon clavipes (Fabricius) (figs. 47 a-c, 89) Syrphus clavipes Fabricius, 1781: 427. Lectotype d (here designated): ‘clavipes in Sehestedt & Tonder Lund col- lection (ZMUC) [examined]. Merodon clavipes var. albus Paramonov, 1927: 90. Syntypes: ‘Bezirk Balta, Gouv. Odessa’ [Ukraine, U. S. S. R.] [not examined]. Merodon clavipes var. ater Paramonov, 1927: 91. Syntypes: locality unknown [not examined]. Merodon clavipes var. niger Paramonov, 1927: 90. Syntypes: ‘Bezirk Balta, Gouv. Odessa [not examined]. Merodon canipilus Rondani, 1865: 131. Lectotype d (here designated): ‘52’ [number refers to description of canipi- lus] in coll. Rondani (LSF) [examined]. Musca clauda Villers, 1789: 463. Syntype(s): ‘Gallia Australiori’ [the southernmost of France] [not examined]. Musca curvipes Gmelin, 1790: 2871. Published in synonymy with Syrphus clavipes Fabricius, 1781. Syrphus gravipes Rossi, 1790: 286. Syntypes: ‘in provinciis Florentina et Pisana’ [Toscana, Italy] [not examined]. Lampetia sacki Paramonov, 1937a: 3. Lectotype © (here designated): ‘Chiklana’ [Chiklana de la Frontera, near Jerez, southern Spain] (ZBSM) [examined]. Syn. n. Merodon senilis Meigen, 1822: 356. Lectotype 9 (here des- ignated): ‘senilis’ (NHMW) [examined]. Syrphus clavipes, Fabricius 1787: 337; Rossi 1790: 286; Fabricius 1794: 292. Lampetia clavipes, Meigen 1800: 34; Coquillett 1910: 557; Oldenberg 1919: 388; Sack 1931: 310; Séguy 1961: 176. Lampetia clavipes var. alba; Sack 1931: 311. Lampetia clavipes var. atra; Sack 1931: 311. Lampetia clavipes var. nigra; Sack 1931: 311. 178 Merodon clavipes Fabricius 1805: 195; Latreille 1804: 443; Meigen 1822: 351; Macquart 1828: 291; Macquart 1834: 513; Westwood 1840: 137; Macquart 1842: 70; Rondani 1845: 256, 259; Macquart 1849: 466; Walker 1849: 597; Rondani 1857: 54, 62; Schiner 1857: 410; Schiner 1862: 344; Palma 1863: 46; Portschinsky 1877: 184; Strobl 1893: 76; Bezzi 1895: 16; Bezzi 1900: 89; Strobl 1900: 593; Verrall 1901: 559; Villeneuve 1903: 115; Sack 1913a: 433; Paramonov 1925: 146; Gil Collado 1930: 243, 246; Gaunitz 1969: 83, 86; Delfinado & Hardy 1975: 343; Marcos Garcia 1985: 197; Hurkmans 1985: 69; Bradescu 1986: 123; Peck 1988: 165, 168 (see for additional references Kertész, 1907: 274). Merodon clavipes var. albus, Liepa 1969: 20; Peck 1988: 169. Merodon clavipes var. ater. Liepa 1969: 20; Peck 1988: 169. Merodon clavipes var. niger. Liepa 1969: 20; Peck 1988: 169. Milesia clavipes, Latreille 1810: 331. Merodon canipilus, Peck 1988: 169. Lampetia canipila, Sack 1931: 310 (as syn. of clavipes). Musca clauda; Kertész 1907: 274; Peck 1988: 168. Musca curvipes, Peck 1988: 168. Merodon curvipes, Meigen 1803: 74 (erroneous); Peck 1988: 168. Lampetia curvipes, Sack 1931: 310 (as syn. of clavipes). Syrphus gravipes, Peck 1988: 168. Merodon senilis, Peck 1988: 168. Material examined. — Algeria: 328, 59 (MNHN, BMNH, ZMAN, ZMUC); Austria: 106, 22 (NHMW); France: 1436, 612 (MNHN, NHMW,Bonn, BMNH, ZMUC, ZMAN); Greece: 1178, 372 (ZMUC, den Hollander, NHMW, Bonn, BMNH, Thessaloniki, ZMAN, JLR, WH); Hungary: 5d, 42 (BMNH, ZMUC); Italy: 1168, 579 (ZMUC, JLR, NHMW, BMNH, ZMAN, WH); Morocco: 19 (BMNH); Spain: 2 Paralectotype of Lampetia sackî: ‘Chiklana’ (ZSBM) and 206, 52 (ER, ZMUC, BMNH, Bonn, ZMAN); Sweden: 12, Bohuslan, leg. Malm, 1862 (MNHN); Switzerland: 16, 22 (ZMUC); Yugoslavia: 31d, 52 (BMNH, NHMW, JLR, ZMAN). Description Male. — Head: Antennae dark, antenna 3 with up- per margin concave, apex subacute, vertex angle 50°, ocellar angle 60°, antennal ratio 1.8; pubescence yel- lowish, darker in ocellar region; face dense white pruinose; tl-v ratio 0.4. Thorax: Dark, slightly metallic lustrous on lateral dorsum, scutellum, katepisternum, anepisternum and anepimeron, pubescence yellow to brown, erect, rath- er dense, with dark interalar band. Wings troubled evenly, halteres, squamae and antisquamae pale yel- low. Legs: All dark, trochanters 3 with high ridge shaped like chisel edge; femora 3 curved, very strong- ly swollen, often with oily lustre, and golden pubes- cence; triangular processus normally shaped, tibiae 3 and tarsi 3 normal. Abdomen: Dark; T II-IV with interrupted arcuate wide pruinose bands and dense pubescence, yellow on T II and III (anteriorly) and orange on posterior abdomen; T II clearly but not very strongly tapering, normally without lateral spots; S IV strongly vaulted, posterior margin widely and deeply emarginate. Genitalia: The anterior surstyle lobe extremely large, bearing short, dense, yellow, even pubescence throughout, laterally ridged; posterior surstyle lobe coalescent, steeply rising, rather high, long, uneven, with less dense pubescence. Cercus well protruding, angular, with dense, long, yellow pubescence. Aedeagus slender, smooth on outer face, somewhat concave subbasally, apical shaft part slightly length- ened, fringed plates on thecal apex recumbent- oblique. Body length 17-23 mm. Female. — Quite different from the males, as they lack the dense abdominal pubescence. Apart from sexual dimorphism the main other differences are: Head: Frons whitish pruinose, leaving lustrous mid- stripe occupying one-fifth of width. Thorax: Pubescence greyish, shorter than in males, rather even, no interalar band. Legs: The femur 3 slightly less swollen, often pubescence duller; compared with other Merodon females, still very much swollen. Abdomen: Pubescence overall much less conspicu- ous; dense only in anterolateral areas of T II, and on pruinose bands; in some specimens large portions of the tergites bald altogether; colour of the pubescence mainly dark yellow to pale; in many specimens pale orange lateral spots can be seen. Body length 15-22 mm. Varieties of clavipes and variation. — In the var. 4/- bus the pubescence is white throughout, except for a black interalar band in both sexes with blue back- ground lustre. In the var. ater the thoracic dorsum shows only black pubescence; of this variety only the males are known. In the var. niger the pubescence is black throughout, except on the face, while the prui- nose abdominal bands are almost or completely ab- sent in both sexes. No types of either of the varieties have been examined; they are considered to be lost (Liepa 1969: 4). Apart from the described varieties and size varia- tion the specimens of this species are remarkably uni- form. A few aberrant individuals totally lack the pu- bescence on the abdomen and have strangely vaulted posterior tergite margins. These specimens are not considered to belong to a separate variety. Possibly they have suffered from frost damage, known to pro- duce aberrances in insects. Diagnosis. — M. clavipes is easily distinguished from other species in this group by its size. Moreover, the combination of enormously swollen femora 3 and chisel-edged trochanters 3 does not occur in males of any other species; the degree of swelling in the femo- Hurkmans: Monograph of Merodon ra 3 in the females far exceeds that of any other Merodon currently known. Period of flight and distribution (fig. 89). — M. cla- vipes occurs from late April through August in south- ern Europe, the Mediterranean and Asia Minor, though it may occasionally occur more northward; the specimen from Sweden seems unquestionable; re- ports of clavipes from Great Britain have been serious- ly doubted by Verrall (1901). Biology. — The territorial behaviour of the males has been described by Hurkmans (1985). In the field the males visit flowers, mainly of Umbelliferae, and are conspicuous, but the females fly close to the soil through the vegetation. Discussion. — The types of canipilus, sacki and sen- ilis have been examined, and found conspecific with clavipes. The types of clauda, curvipes and gravipes have not been examined. The original description of clauda leaves hardly any doubt that this is a synonym of clavipes this synonymy has been considered by Kertész (1907: 274), while Peck (1988: 168) also names clauda as synonym of clavipes. Musca curvipes was originally published as a synonym of Syrphus cla- vipes. Syrphus gravipes refers to the publication of the female of clavipes, a possibility already considered in the original description. Merodon cupreus sp. n. (figs. 48 a-c, 88) Type-material. - & Holotype “Turkey, Kars, Handere 2100-2200 m, 20 km W of Sarıkamıs, 1.viii.1983, leg. J. A. W. Lucas’ (ZMAN). Paratypes: 298, 59: 244,49 topotyp- ic, with dates ‘1.viii.1983’ and ‘2.viii.1983’ (LR), 12 with same data (ZMAN); 36 ‘Turkey, Hakkari S. of Yiiksekova 28.vi.1986 W. Schacht’ (LR); 26 “Turkey, Kars, 8 km W. of Sarıkamıs 2000-2050 m, 9.vii.1986' leg. J. A. W. Lucas (JLR). Description Male. — Head: Antennae dark, antenna 3 cordate, apex subacute; antennal ratio 1.4; vertex angle 30°, ocellar angle 35°, pubescence yellow, often with dark tuft in ocellar region; face and occiput white pruino- se, somewhat bluish lustrous; tl-v ratio 0.35. Thorax: Dark, metallic lustrous on lateral dorsum and scutellum, katepisternum, anepisternum and an- epimeron; pubescence deep yellow to brown, even, rather dense, with dark interalar band. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: Dark, but tarsi laterally with golden pubes- cence, contrasting; trochanter 3 sharply keeled, with chisel edge, femur 3 rather swollen, slightly lustrous, with normally shaped triangular processus bearing 6- 179 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 7 bristles on very strongly serrate distal margin; tibiae 3 normal, basitarsi 3 slightly swollen. Abdomen: Dark, no lateral spots on T II; T II-IV with white pruinose bands, straight on T II, arcuate on T III-IV; interrupted on T II-III, continuous on T IV; pubescence golden to brassy or orange, rather dense, recumbent; T II tapering; S IV vaulted, deep- ly, rather narrowly emarginate posteriorly. Genitalia: The anterior surstyle lobe large, round- ed, with basal projection on ventral margin, and dense, short, even, yellowish pubescence anteriorly; laterally weakly ridged; the posterior surstyle lobe sep- arated by shallow sulcus, low, elongate, bearing yel- low, moderately long, uneven pubescence along ven- tral and posterior margins. Cercus small but well protruding, with dense, long, yellow pubescence. Aedeagus moderately slender, fringed plates on thecal apex oblique, apical shaft part short. Body length 12-15.5 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons tapering poste- riorly, sides white pruinose, lustrous midstripe opccu- pying one-third of width; pubescence grey, but dark tuft present in ocellar region; ocellar angle 60°. Thorax: Dorsum dull, pubescence grey, dark interalar band present. Legs: Dark, with grey pubescence; on tibiae and tarsi yellow-grey pubescence with some contrast as in d but weaker; trochanter 3 smooth; tri- angular processus as in males. Abdomen: Stouter than in male, with rather dense grey pubescence, pruinose bands moderately wide, continuous, arcuate; tergite V dark, with scattered grey pubescence. Body length 13 mm. Diagnosis. — At first sight, cupreus seems to be a small clavipes. Differences between those species are: antennal ratio in cupreus 1.4, in clavipes 1.8; antenna 3 with upper margin convex in cupreus, concave in clavipes, triangular processus stronger serrate in cupre- us than in clavipes, in the males of clavipes the anteri- or surstyle lobe is much larger than in those of cupre- us, in the female cupreus lateral abdominal spots lacking in, present in clavipes, size in clavipes 15-23 mm, as opposed to 12-15.5 mm in cupreus. The dif- ferences between dzhalitae (discussed hereafter) and cupreus are: antenna 3 with upper margin convex in cupreus, concave in dzhalitae, on the hind legs the tar- si Il in dzhalitae are almost as long as the basitarsi; in cupreus they are less than half as long. Moreover, in the males of cupreus the S IV is deeply emarginate posteriorly while this element is entire in dzhalitae. Differences with the description of M. clavipes var. albus, stated to have all black and white pubescence in both sexes, are: The size in clavipes var. albus is great- er, in cupreus the pubescence is not black and white in 180 both sexes. Moreover, the differences (with respect to clavipes) already mentioned apply. The 2 cupreus are of the same size and habitus as those of femoratus. They are distinguished by their lack of pruinose bands on the thorax and much denser overall pubes- cence; also the shape of the antenna 3 is different: ovoid to cordate in cupreus, more elongate in femora- tus. Period of flight and distribution (fig. 88). — M. cu- preus occurs in July and August in eastern Turkey. Etymology. — The adjective cupreus refers to the colour of the larger portion of the abdominal pubes- cence in the male of this species. Merodon dzhalitae Paramonov (fig. 99) Merodon dzhalitae Paramonov, 1927: 89. Holotype d: ‘Hissar, Yalta’ [south slope of the Crimea mountains, Crimea, Ukraine] [not examined, considered lost]. Merodon dzhalitae, Liepa 1969: 20; Van der Goot 1981: 215; Peck 1988: 169; Zimina 1989: 24. Lampetia dzhalitae, Sack 1931: 313. Description The description of dzhalitae presented here is based on the original description, on Sack (1931: 313) and on Zimina (1989: 24) who described recently collec- ted material of both sexes. Male. — Head: Antennae dark, antenna 3 yellow- brownish, upper margin concave, lower margin con- vex, apex acute; face, frons and vertex with yellow pu- bescence, face sparsely pruinose; compound eyes with yellow pubescence; tl-v ratio 0.5. Thorax: Dorsum dark with some weak blue lustre, with conspicuous orange pubescence throughout, scutellum concolorous, duller; wings slightly trou- bled; reddish pubescence clear, interalar band some- times present. Legs: All dark, with yellow pubescence, femur 3 normal, resembling this element in avidus [therefore less swollen than in e.g. clavipes|, triangular processus with strong apical pedestal (cf. Zimina 1989: fig. 2); on hind legs the second tarsi almost as long as basitar- Si. Abdomen: T II slightly tapering only, clearly bluish lustrous, T III-IV with white pruinose bands, just interrupted on T III, just continuous on T IV; pubes- cence reddish, but black, short and recumbent on posterior margins of T II-III. S IV with posterior margin entire, keeled. Body length 13 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Habitus: Pubescence overall paler than in male. Head: With broad lustrous mid- stripe on frons. Antenna 3 less long than in d, ovoid, dark with paired yellow spots ventrally. Legs: Tibiae brownish basally. Abdomen: T II showing nearly tri- angular greenish metallic lustrous lateral spots which are reddish translucent mediad. Pubescence dark, short and recumbent on T II-III, reddish elsewhere. Body length 13 mm. Diagnosis. — The differences with respect to cupre- us, which seems to resemble dzhalitae most closely, are as follows. Third antennal article concave in dzhalitae, convex in cupreus tarsi II on hind leg as long as basitarsi in dzhalitae, only half as long in cu- preus, and S IV entire in dzhalitae, deeply emarginate in cupreus. Period of flight and distribution (fig. 99). — The species occurs in August and probably is endemic to the Crimea. Discussion. — In view of the extensive original de- scription and the full account by Zimina (1989) it seems clear that dzhalitae is a distinct species. The de- scription of the recently collected material by Zimina (1989) matches the original description on all signifi- cant points. M. dzhalitae has been tentatively assigned to the clavipes group on account of its external simi- larities with e.g. clavipes and cupreus. Merodon hamifer Sack (figs. 49 a-b, 88) Merodon hamifer Sack, 1913a: 436. Lectotype d (here des- ignated): ‘Anatolien Ak-chehir [Aksehir] Korb 1900/ Mer. hamifer Sack det. Sack/ coll. Lichtwardt/ coll. D. E. I. Eberswalde’ (DEIC) [examined]. Merodon hamifer, Paramonov 1926b: 322; Paramonov 1927: 78; Peck 1988 : 171. Lampetia hamifera; Sack 1931: 318. Material examined. — Greece: 1d ‘Samos, Ampelos 22.vi.1932, Werner (NHMW); Turkey: Paralectotype 9, same data as lectotype. Description M. hamifer is similar to aberrans except in the fol- lowing details. Male. — Head: Antennal ratio 2.0; antennae all orange. Thorax: Pubescence less dense. Legs: All tib- iae and tarsi orange, tibiae 3 basally brown; femora 3 bearing modified triangular processus, basitarsi 3 showing apical projection on medial face. Genitalia: Pubescence on posterior surstyle lobe denser. Body length 13.5-14.5 mm. Female. — Except for sexual dimorphism, differing HURKMANS: Monograph of Merodon from the male as follows. Head: Compound eyes sep- arate, frons lustrous, slightly dulled down sides; an- tennae orange, antennal ratio 2.8. Legs: Similar to those of hamifer male, triangular processus slightly smaller. Abdomen: Pruinose bands wider; tergite V narrower. Body length 14 mm. Diagnosis. — This close relative of aberrans can be distinguished from all other species by the shape of the triangular processus in both sexes. Period of flight and distribution (fig. 88). — M. ha- mifer occurs in the Greek Aegean islands and western Turkey, where it flies in June. Merodon karadaghensis Zimina Merodon karadaghensis Zimina, 1989: 24. Holotype d: ‘Karadagh, Crimea, 5. ix. 1984, leg. Zimina’. Moscow Zoological Institute [not examined]. It appears that this species may be related to M. dzhalitae, with which it is compared by Zimina (1989). The differences are stated to be in the relative length of the tarsal joints, the outline of the triangular processus on the hind femora, while the abdominal pruinose bands also slightly differ. Diagnosis. — After Zimina (1989), M. karadaghen- sis may be distinguished from M. dzhalitae by the length ratio of the basal tarsal joints (II:I) on tarsus 3, which is about 0.5 in karadaghensis and 0.8 in dzhal- itae,and also by the abdominal pruinose band on T IV being chevron-shaped in karadaghensis while this is more rounded in dzhalitae. Discussion. — This species is provisionally regarded as belonging to the c/avipes group, on the grounds of resemblance to M. dzhalitae. Merodon lusitanicus sp. n. (figs. 50 a, 93) Type-material. — 2 Holotype ‘Portugal, Algarve, Quarteira 27.iv.1985, J. A. W. Lucas’ (ZMAN); paratype 9, same data, but locality: ‘Vilamoura’ (JLR). Description Female. — Head: Antennae black, antenna 3 with upper margin concave, apex rounded, antennal ratio 2.8; pubescence white on face, black on frons and ver- tex, pale brown on occiput; frontal sides whitish prui- nose, leaving lustrous midstripe on median one-third; ocellar angle 60°. Thorax: Dark, dorsum with 4 vague pruinose 181 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 bands and yellow pubescence; pubescence duller on scutellum and sides, katepisternum, anepisternum and anepimeron steel grey lustrous. Wings clear; hal- teres, squamae and antisquamae yellow. Legs: All dark, joints lighter; femora 3 extremely swollen for a female, triangular processus showing strongly serrate distal margin bearing 10 bristles. Abdomen: Dark with some purplish lustre, trian- gular in outline, with clear lateral spots on T II com- bined with dense erect white pubescence; pubescence otherwise sparse; T II-IV bearing rather narrow, con- spicuous widely interrupted somewhat arcuate prui- nose bands; T V acute, all dark. Body length 11-12 mm. Male. — Unknown. Diagnosis. — M. lusitanicus can be separated from aberrans by the lateral spots on T II and the swollen femora 3; /usitanicus is too dark to be confused with females of avidus, nigritarsis or distinctus, its habitus can be compared with that of alexeji from which it differs in larger size, longer antennae and more swol- len femora 3. Period of flight and distribution (fig. 93). — This species, known only from the types, has been found in Portugal in April. Discussion. — In view of the shape, width and posi- tion of the abdominal pruinose bands, relative bald- ness of the abdomen with purplish lustre, and gener- al habitus, /usitanicus is provisionally regarded as closely related to aberrans, and therefore assigned to the clavipes group. Etymology. — The adjective /usitanicus refers to the classical name for Portugal, Lusitania. Merodon quadrinotatus (Sack) (figs. 51, 85) Lampetia quadrinotata Sack, 1931: 324. Holotype 2: ‘Mesopotamia’ [Iraq] [not examined, considered lost]. Merodon quadrinotatus, Peck 1988: 173. Material examined. — Turkey: 19 ‘Hakkari, Suvarihalil Pass W side 2300-2400 m 11.viii.1983 leg. J. A. W. Lucas’ (ZMAN). Description Female. — Similar to the female of clavipes, but with the following differences. Head: Frons much tapering posteriorly, laterally whitish pruinose leaving a lus- trous midstripe occupying a quarter of the width; pu- _ bescence white, but grey on occiput. Thorax: Grey pubescence on dorsum, with conspicuous black inter- 182 alar band. Legs: Pubescence grey to black, back- ground colour deep black. Abdomen: Black, pubescence dense and black throughout but with two pairs of very conspicuous white spots, one each on T HI-IV, and weak latero- posterior triangular spots of grey pubescence on same tergites. Body length 14 mm. Diagnosis. — The dense black, white-dotted pubes- cence of the abdomen is unique and at once separates quadrinotatus from all other species currently known. Period of flight and distribution (fig. 85). — The species is known from ‘Mesopotamia’ [Iraq] (month not reported by Sack) and the eastern Turkish moun- tains where it was collected in August. Discussion. — Sack (1931) did not indicate a de- pository of the holotype. He did provide a reasonably clear description of guadrinotatus. Given the unique features of the species the identification of the Turkish specimen is unquestionable. Since quadri- notatus is believed to be a close relative of clavipes the species is assigned to the c/avipes group. Merodon splendens sp. n. (fig. 47 d, 91) Type-material. — ¢ Holotype ‘Sardaigne St. Ussassai 16.v.1977 P. Goeldlin’ (LAU). Description Male. — Head: Antennae blackish, antennae 3 rounded-subacute, upper margin about straight, ver- tex angle 30°, ocellar angle 40°, tl-v ratio 0.4; pubes- cence pale yellow, face greenish lustrous, yellow prui- nose; slight dark tuft present in ocellar region; oral margin weakly protruding. Thorax: Dorsum and sides dark, weakly bluish lus- trous on anepisternum, katepisternum and anepime- ron; pubescence moderately dense, erect, pale yellow, with dark interalar band; no pruinose bands present. Wings clear, halteres, squama and antisquama pale yellow. Legs: Black, with yellow pubescence; trochanters 3 with curved low chisel-edge; femora 3 very much swollen, with some oily lustre, curved. Abdomen: T II strongly tapering, T III-IV cylin- drical. Lateral spots on T II weak, micaceous; prui- nose bands on T II-IV very clear, interrupted, arcu- ate; pubescence moderately dense, yellow on T II, deep orange on T III-IV. Genitalia: Anterior surstyle lobe reniform, pubes- cence, although dense, so extremely short that it is in- conspicuous; posterior surstyle lobe coalescent, much bulging outward anteriorly, rounded, with short, erect, moderately dense pubescence. Cercus slightly protruding, bearing short, pale pubescence. Aedeagus slender, with smooth outer face, subapical cavity well marked, somewhat concave subbasally; fringed plates on thecal apex recumbent. Body length 14.5 mm. Female. — Unknown. Diagnosis. — M. splendens is a close ally of clavipes and velox. From the latter it is distinguished by the clear wings. The differences with clavipes, apart from the smaller size, are: Antennae more slender, pubes- cence on head, thorax and abdomen less dense and shorter, T II more tapering, III-IV cylindrical. Differences with both species are: Anterior surstyle lobe more curving dorsad, differently shaped (cf. figs. 46 a and d), pubescence much less dense and very much shorter on anterior and posterior surstyle lobes and cercus. M. splendens can be distinguished from cupreus and dzhalitae by the swollen femora 3. The same character also separates sp/endens from brevis, to which it is slightly similar on account of its habitus. Moreover, the anterior surstyle lobe is quite different- ly shaped in brevis. It differs from aberrans by the strongly swollen femora 3 and denser pubescence. Period of flight and distribution (fig. 91). — M. splendens is known only from the holotype, collected in Sardinia, in May. Etymology. — The adjective splendens is latin and means splendid or beautiful. It was given in view of the colourful pubescence of this species. Merodon velox Loew (figs. 47 a-c, 91) Merodon velox Loew, 1869: 253. Lectotype d (here desig- nated): ‘Rhodus/ Alte Sammlung’ (NHMW) [examined] . Merodon velox var. anathema Paramonov, 1925: 149. Holotype 2: ‘Mons Takalta, prope Kulp, Transkaukasien’ [in Turkey, approx. 38° 30 N, 41° 05 E] [not examined, probably lost]. Merodon velox var. armeniacus Paramonov, 1925: 147. Syntypes: 126 and 19 ‘Eriwan’ and ‘Ordubad (Erivan)’ [Armenia] [not examined, probably lost]. Merodon velox, Sack 1913a: 434; Paramonov 1925: 147; Paramonov 1926a: 15; Peck 1988: 175. Merodon velox var. anathema, Liepa 1969: 20. Merodon velox var. anathemus, Peck 1988: 175. Merodon velox var. armeniacus, Liepa 1969: 20; Peck 1988: 175. Lampetia velox, Sack 1931: 331. Material examined. — Greece: 2 Paralectotype with same label as lectotype; 1006, 732 (BMNH, NHMW, RMNH, ZMAN, JLR, WH); Italy: 26, 12 (NHMW); Turkey: 16d and 32 paralectotypes ‘As. Minor/ Alte Sammlung’ (NHMW); Hurkmans: Monograph of Merodon 496, 372 (MNHN, RMNH, NHMW, BMNH, JLR); U. S. S. R.: 38,12 (NHMW, RMNH); Yugoslavia: 25,32 (NHMW). Description Male. — Head: Antennae dark, antenna 3 with up- per margin concave, apex subacute, antennal ratio 1.8; pubescence pale yellow, with few dark hairs in ocellar region; face and occiput densely white to yel- low pruinose; vertex angle 40°, ocellar angle 70°. to 85°, tl-v ratio 0.5. Thorax: Dark, lateral dorsum and scutellum, kate- pisternum, anepisternum and anepimeron weakly blue lustrous; pubescence rather dense, erect, brown- ish, dark interalar band present. Wings strongly in- fuscate, except apical-posterior margin; halteres, squamae and antisquamae yellow. Legs: All dark, with yellow pubescence; trochanter 3 sharply ridged, femur 3 strongly swollen, triangular processus normally shaped, tibiae 3 and tarsi 3 nor- mal; Abdomen: Rather slender; all dark; T II tapering, bearing vague pale orange lateral spots, T I-IV with widely interrupted arcuate whitish pruinose bands, pubescence moderately dense, orange; S IV vaulted, posteriorly deeply, widely emarginate. Genitalia: Completely similar to those of clavipes. Body length 17-23 mm. Female. — Except for the rather marked sexual di- morphism, differing from the male as follows. Head: Frons with lustrous midstripe occupying one-third of width, laterally white pruinose. Thorax: Incon- spicuous longitudinal pruinose bands present, pubes- cence brown to grey throughout; no interalar band; wings less infuscate, strongest on surface bordering veins. Legs: The trochanter 3 smooth, femur 3 slen- derer than in & but still much swollen, pubescence duller. Abdomen: Pruinose bands conspicuous, stronger arcuate, less interrupted, rather variable between specimens; pubescence almost restricted to T II where dense; lateral spots clearer. Body length 17-18 mm. Diagnosis. — M. velox is easily distinguished by the infuscate wings in the male; in the female it could be confused with clavipes and pruni from which it is dis- tinguished by the veins bordered with infuscation (in clavipes wings clear; in pruni wings more evenly trou- bled), from pruni also by the all dark legs, less slender, much darker coloured abdomen and larger size. Moreover, in pruni the genitalia are quite different. Period of flight and distribution (fig. 90). — M. ve- lox occurs in the eastern Mediterranean where the im- agines are found from May through August; the spe- cies may be bivoltine. 183 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Biology. — The males of velox, observed by the au- thor in May of 1984 in several Greek localities, were flying close to the soil on rather dry sunny hillsides with short vegetation. Discussion. — Although the similarity of the geni- talia with respect to clavipes is clear, the following ar- guments against synonymizing velox can be listed: (1) there are clear external differences; (2) the distribu- tion of velox is quite restricted and purely Mediterranean, whereas clavipes occurs over all of southern and central Europe; (3) the flight period of velox is more restricted; (4) there is some evidence that velox, where it occurs together with clavipes, pre- fers more open, drier biotopes (author’s observations; J. Lucas, pers. comm.); therefore, the ecological nich- es might be different; (5) behavioural differences seem to exist. When the velox males were observed in Greece, they did not show territorial behaviour. Males of clavipes observed at the same locality and date displayed territorial behaviour (cf. Hurkmans, 1988). This may indicate that either velox shows no territorial behaviour, or this behaviour takes place in a different period of the season. Merodon velox vat. anathemus Paramonov Material examined. — Greece: 12 (WH); Turkey: 19 (WH). Diagnosis. — Completely similar to the nominate form, except for the all dark pubescence. Only fe- males are known. Merodon velox var. armeniacus Paramonov Material examined. — Turkey: 12 (wr) Diagnosis. — Completely similar to the nominate form except in the abdomen where lateral spots are lacking and pruinose bands are reduced, in both sex- es. Discussion. — No recent male material is known; of the syntypes, all but one were males. Possibly some of the material might be present in the collection of the Museum of Armenia, Erivan. Merodon warnckei sp. n. (figs. 52 a-d, 88) Type-material. — d Holotype ‘Turkey, Hakkari, 5.vili.1983, Gavaruk-lake Mt. Sat, 2900 m, leg. Warncke’ [37° 20° N, 43° 35° E] (ZMAN). Paratypes: 48, 19; 1d ‘Turkey, Hakkari, Sat Mountains north of Mt. Gavaruk 184 2900 m, 7.viii.1983 leg. Warncke’ (LR); 2d, 19 “Turkey, Hakkari, Sat Mountains, south of Sat Lake, 2800 m, 7.viii.1983, leg. Warncke’ (LR); 16 ‘Turkey, Van, Mengene Dagi north of Baskale, 2700-3000 m, 27/28.vi.1986, leg. Hurkmans’ (wr). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apax acute, antennal ratio 2.0; pubescence yellow throughout, pruinosity slight on face, dense on occiput; face strongly blue lustrous, oc- ciput greenish lustrous; vertex angle 25°, ocellar ang- le 60°, tl-v ratio 0.3. Thorax: Dark, greenish to blue lustrous on lateral dorsum and scutellum, katepisternum, anepisternum and anepimeron; slightly pruinose on sides; pubes- cence rather even, moderately dense, dull yellow. Wings slightly tinged; halteres, squamae and anti- squamae yellow. Legs: Dark except joints on legs 1-2, leg 3 all dark; trochanters 3 smooth, femora 3 bluish lustrous, swol- len, with normally shaped triangular processus bear- ing 6-9 bristles, tarsi 3 normally shaped; pubescence yellow. Abdomen: All tergites dark with some micaceous purple lustre; vague lateral pruinose spots present on T II; T ILIV with arcuate widely interrupted moder- ately wide yellowish pruinose bands; pubescence dull yellow, moderately dense throughout; S IV shallowly emarginate posteriorly, rather flat. Genitalia: The anterior surstyle lobe very large, rather stalked, bearing dense, even, yellow pubes- cence apically, coalescent with posterior surstyle lobe; the latter with anterior low elongate projection bear- ing yellow, rather dense pubescence, posteriorly with long, dense, yellow pubescence. Cercus rather large, showing dense, very long, yellow pubescence. Aedeagus moderately slender, slightly widening api- cally, apical shaft part short, fringed plates on thecal apex recumbent. Body length 14-15 mm. Female. — Except for the conspicuous sexual di- morphism, differing from the male as follows. Head: Lustrous midstripe on frons occupying one-third of width; slight dark hair-tuft in ocellar region; face weakly lustrous, apex of antenna 3 subacute; pubes- cence grey. Thorax: Pruinose bands on dorsum more distinct; pubescence greyish yellow. Legs: With grey pubescence. Abdomen: Pruinosity more extensive on T II, weaker on T III-IV; pubescence grey, less dense than in male; lustre weaker; T V tapering, apex rounded. Body length 14 mm. Diagnosis. — M. warnckei males can be distin- guished from those of other species in this group by its relatively stout build, the stalked anterior surstyle lobe and the strong metallic lustre of the face; the fe- males of warnckei are stouter and smaller, and have less swollen femora 3 than clavipes or velox, are much stouter than pruni or the species in e.g. the avidus and alagoezicus groups from which they differ as well in having much less conspicuous pruinose bands. Period of flight and distribution (fig. 88). — The species is known from eastern Turkey; it has been col- lected in high altitudes only, where it occurs in June and August; in view of the few specimens collected, the split occurrence does not necessarily indicate that warnckei is bivoltine. Etymology. — The species has been named after the German entomologist K. Warncke, who collected the holotype and most paratypes. A noun in genitive case. The pruni group Apomorphies: The d genitalia with surstyle sulca- te; vertex angle very small; yellow coloration on abdo- men extensive. Merodon pruni (Rossi) (figs. 53 a-d, 94) Syrphus pruni Rossi, 1790: 293. Syntypes: ‘in provinciis Florentina et Pisana’ [Toscana, Italy] [not examined]. Merodon pruni var. obscurus Gil Collado, 1929: 407. Holotype d: ‘Tanger, Mz. Escalera/ M. pruni var. ob- scurus Gil Tipo, Gil Collado det./ M. N. C. N. Madrid’ (IEE) [examined]. Merodon fulvus Macquart, 1834: 514. Syntypes ‘France Meridionale’ [not examined]. Merodon fuscinervis Von Réder, 1887: 73. Syntypes: ‘bei Elos [Elous, 35° 20’ N, 23° 36° E] in der Nahe von Kisamos auf der Insel Kreta’ [not examined]. Merodon pallidus Macquart, 1842: 70. Lectotype d (here designated): ‘Baghdad’ [examined]. Merodon sicanus Rondani, 1845: 258, 264. Lectotype 9 (here designated): ‘58’ [number referring to description of M. sicanus in catalogue by Rondani] [examined]. Merodon pruni, Schiner 1857: 415; Bezzi 1900: 90; Becker 1912: 602; Sack 1913a: 434; Paramonov 1925: 152; Paramonov 1926a: 15; Hurkmans 1985: 69. Merodon pruni var. obscurus, Peck 1988: 173. Lampetia pruni, Sack 1931: 323; Séguy 1961: 181; Van der Goot 1964a: 431. Merodon pallidus, Brunetti 1923: 217. Lampetia pallida; Sack 1931: 323. Merodon sicanus, Rondani 1857: 55, 65; Schiner 1857: 411; Schiner 1862: 347; Palma 1863: 46; Strobl 1893: 76; Strobl 1900: 593. Lampetia sicana; Sack 1931: 323. Material examined. — Algeria: 6d, 22 (BMNH, MNHN); Cyprus: 104, 102 (BMNH, NHMW, KBIN); France: 1d, 12 (MNHN); Greece: 236, 112 (MNHN, NHMW, BMNH, JLR, Hurkmans: Monograph of Merodon Den Hollander, WH); Iran: ® paralectotype of pallidus. ‘Baghdad’ (MNHN); 1d, 12 (RMNH); Italy: 9. paralectotype of sicanus: ‘58’; 796, 322 (BMNH, NHMW, ZMAN, RMNH, JLR, WH); Israel: 166, 119 (KBIN, NHMW, MNHN, NHMW, BMNH); Lybia: 6d, 72 (BMNH); Morocco: 5d (MNHN, RMNH); Pakistan: 116, 109 (BMNH); Syria: 1d (NHMW); Turkey: 88, 142 (JLR, ZMUC, MNHN). Description Male. — Head: Antennae brown to orange, antenna 3 with upper margin convex, apex subacute, antennal ratio 1.4; pubescence yellowish, darker in ocellar re- gion; face and occiput rather densely pale yellow prui- nose, weakly blue metallic lustrous; vertex angle 25°, ocellar angle 35°, tl-v ratio 0.4. Thorax: Dark; katepisternum, anepisternum and anepimeron weakly blue lustrous; pubescence on dor- sum and scutellum rather short, very even, yellow, in some specimens with more or less clear interalar band and/ or with more or less vague pruinose longitudinal bands; pubescence on sides concolorous, less even. Wings troubled, strongest in anterio-basal portion; halteres, squamae and antisquamae pale yellow. Legs: The femora dark with yellow apices, tibiae yellow with broad dark distal band, tarsi 1 and 2 yel- low, tarsi 3 dark with striking golden pubescence on sides contrasting dark upper face; trochanters 3 bear- ing acute projection, femora 3 strongly swollen, bear- ing much projecting triangular processus holding 6-9 bristles on serrate distal margin; all tarsi normal. Abdomen: Rather slender; T II strongly tapering, with large yellow lateral spots; background colour dark brown, but in most specimens replaced with yel- low to a large extent, in some specimens yellow throughout; pruinose bands pale yellow, arcuate, in- complete on T II, wide and hardly interrupted on T III-IV where they may occupy most of the tergite sur- face; pubescence rather dense, predominantly yellow, longest laterally; sternites yellow, S I-II pale, III-IV darker, the latter strongly vaulted, deeply, narrowly emarginate posteriorly. Genitalia: Anterior surstyle lobe rounded, bearing short, erect, yellowish pubescence throughout except basally; separated from posterior surstyle lobe by shal- low sulcus; posterior surstyle lobe bearing long, un- even, erect, yellow pubescence. Cercus conspicuous, wide, with long, dense, yellow pubescence, somewe- hat angular. Aedeagus rather short, moderately slen- der, apical shaft part short, fringed plates on thecal apex recumbent. Body length 8-23 mm. Female. — Very similar to the male. Except for sex- ual dimorphism, differing as follows. Head: Frons with lustrous midstripe occupying less than 0.2 of frontal width; ocellar angle 45°. Thorax: Narrower than in male; Pruinose bands often vaguely present 185 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 on dorsum; wings stronger infuscate. Legs: Femora 3 less swollen, but strongly swollen compared with fe- males of other species of Merodon; trochanters 3 smooth. Abdomen: Pruinose bands wider medially, narrower laterally compared with males; T V taper- ing, all dark. Body length 10-21 mm. Variation. — Merodon pruni shows an immense variation in size, which does not seem to be correlat- ed with locality or flight period. The specimens as- signed to pruni can be arranged in a continuous se- ries; the expression of several characters seems to be correlated with specimen size, e.g. relative size of pro- jection on trochanters 3 and triangular processus. This phenomenon is also found in other species, most strongly in kawamurae. The differences between spe- cimens may be the consequence of differing food availability in the larval stage. Diagnosis. — The usual size in both sexes (normal range 17-21 mm) and predominantly yellow abdo- men combined will distinguish pruni from all other species except clavipes and velox, from these two the males are easily distinguished by the more slender ab- domen with less dense pubescence, moreover from velox by the absence of deep infuscation on wings; the females of pruni are paler and have more slender ab- domina than those of clavipes and velox. Small speci- mens can be distinguished from other Merodon, in the males by the nearly all yellow abdomen and the genitalia; the dwarf females may cause some problems but can be distinguished by their triangular yellow abdomina and darkened wings. In pruni females of all sizes the darkening of the wings is black, without pur- ple reflections. Period of flight and distribution (fig. 94). — M. pruni has been recorded by Sack (1931: 324) to fly from May to August in southern Europe, north Africa and Asia Minor. The species is common in Toscana, Italy and in central Greece. From the speci- mens in collections it appears that pruni may be bi- voltine at least in Italy and Greece. M. pruni seems to be rare in southern France and absent from Spain. Biology. — M. pruni occurs in rather open habitats with scattered tall plants, mainly Umbelliferae and Liliaceae, on hillsides exposed to the sun (Hurkmans 1985); the males are territorial and aggressively expel invaders; the females are much less conspicuous in the field than the males as they fly close to ground level (Hurkmans 1985). This difference in flight behavi- our might explain the sex ratio found among pre- — served material in museum collections studied, which is about 1.7 males to a female. 186 Discussion. — Type-material of fulvus Macquart, 1834 and fuscinervis Von Réder, 1887 could not be examined. To judge from the descriptions both taxa seem to fall well within the variation range of pruni. The synonymy with pruni was already given by Sack (1913a, 1931) and Peck (1988). Merodon pruni var. obscurus Gil Collado Diagnosis. — Rather similar to the nominal form, except for the following differences in the male (fe- males unknown). Head: Antennae 3 dark brown [not black as indicated in description], truncate. Thorax: Pubescence very short, even, yellow, but darker in middle of dorsum, transition gradual; no pruinose bands present. Wings darkened along anterior mar- gin, the brown veins bordered with slight infuscation. Body length 18 mm. The longicornis group Apomorphies: Extreme lengthening of antenna 3; male genitalia with surstyle deeply sulcate and very slender aedeagus; vertex angle very small. Description. — Head: Antennae 3 with upper mar- gin straight, apex subacute to rounded, antennal ratio 2.2 to 4.8; vertex angle 25° to 30° in males, ocellar angle some 35° in males, 60° in females; in females lustrous midstripe on frons occupying 0.35 to 0.6 of width. Thorax: Dark, rather narrow, bearing 4 longi- tudinal, mostly clear pruinoise bands. Wings clear. Legs: All dark, sometimes joints paler, trochanters 3 smooth, femora 3, triangular processus and basitarsi 3 normally shaped. Abdomen: Rather slender; T II somewhat tapering, T III-IV cylindrical in male, tapering in female; in er- ivanicus and longicornis dark background coloration partially replaced by crimson red colour typical of this group; pruinose bands narrow, arcuate and conspicu- ous. Genitalia; Anterior surstyle lobe quadrate to rec- tangular, anteriorly showing dense, yellow pubes- cence, separated from posterior surstyle lobe by deep sulcus, stalked; the posterior surstyle lobe low, short to elongate. Cercus well protruding. Aedeagus very slender, fringed plates on thecal apex recumbent to oblique. Body length in both sexes 9-13 mm. Merodon erivanicus Paramonov (figs. 54 a-b, 55 a, 92) Merodon erivanicus Paramonov, 1925: 151. ‘Erivan, Armenien [not examined]. Merodon erivanicus, Liepa 1969: 20; Peck 1988: 170. Syntypes: Lampetia erivanica; Sack 1931: 315; Sack 1938: 22. Material examined. — Greece: 29 (NHMW); Israel: 19 (BMNH); Turkey: 192 (BMNH, JLR, WH); Yugoslavia: 19 (NHMW) . Description Female. — Head: Antennae dark brown, antenna 3 with upper margin convex to straight, apex obtuse, antennal ratio 2.2 to 2.8; ocellar angle 60°, pubescen- ce yellow, face and occiput with slight metallic lustre obliterated by dense whitish pruinosity; frons densely pruinose laterally, midstripe occupying 0.4 of width. Thorax: Dark, dorsum with well marked pruinose bands, pubescence greyish yellow, short, dense, even; katepisternum, anepisternum and anepimeron green- ish to bluish metallic lustrous, hardly pruinose, pu- bescence concolorous. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: Dark, tibiae 1 and 2 basally yellow; dense yel- low pubescence on sides of tarsi, contrasting dark pu- bescence on upper faces; trochanters 3 smooth, femo- ra 3 slightly swollen, triangular processus normal. Abdomen: Slender, dark coloration partially re- placed by crimson; the extent of the crimson colour is much variable between specimens; T II-IV with inter- rupted arcuate white pruinose bands; pubescence very short, inconspicuous, black on dark surface, yellowish on crimson surface; S I-II red, III-IV dark red to black. Body length 10-12 mm. Male. — Unknown. Diagnosis. — M. erivanicus is distinguished from fe- males of other Merodon species by the long antenna 3 and crimson colour on the abdomen; from longicornis by the shorter antenna 3 (antennal ratio in longicornis is 4.1 to 4.8). Period of flight and distribution (fig. 92). — M. er- ivanicus occurs in June and July, in the eastern Mediterranean, in eastern Turkey and Armenia. Merodon kaloceros sp. n. (figs. 54 d, 55 c, 56 a-c, 87) Type-material. — d Holotype “Turkey, Antalya, H. Coene, J. Lucas & B. van Oorschot / Irmesan Gedigi 12 km N of Akseki 1600 m 24.vii.1981’ (ZMAN). Paratypes: 24d, 19:19, same data as holotype (LR); 48 “Turkey, Hakkari, Suvarihalil pass 1250 m, W side near Habul Deresi 13.vi.1984 leg. J. A. W. Lucas’ (LR); 2d “Turkey Hakkari, Chilo Daglari N of Oramar 1400 m, 16/17.vi.1984, leg. J. A. W. Lucas’ (LR); 28 “Turkey, Hakkari, Sat Dag, SW Yüksekova 1600 m 18.vi.1984 J. A. W. Lucas’ (LR); 8d ‘Turkey pr. Hakkari, Sat Dag, Varegös SW Yiiksekova 28. and 29. vi.1985’ (LR); 18 ‘Turkey, Van, 40 km SW Hurkmans: Monograph of Merodon Akdamar 1720 m, 17.vii.1986, leg. P. van Ooijen’ (ZMAN); 16 ‘Dalmatien [Yugoslavia], Gravosa vi.28 [1928] Zerny (NHMW); 1d ‘Greece, Ithaca 6/11.vi.1965 F. J. Francois’ (KBIN); 24 ‘Hill Scrub 500 ft above lake/ Macedonia Prespa geul Otesevo 20/27.vi.1958/ Yugoslavia R. L. Coe’ (BMNH); 1d ‘Yugoslavia Kroatie Bribir (50 100) 10.viii.1964 H. J. P. Lambeck’ (JLR); 16 ‘Corfu, Erber 218’ [Greece] (JLR). Description Male. — Head: Antennae dark, antenna 3 brown with base and apex paler, upper margin straight to convex, apex subacute to obtuse, antennal ratio 2.3 to 2.7; pubescence greyish yellow with dark tuft in ocel- lar region ; vertex angle 25-30°, ocellar angle 40°, tl- v ratio 0.55. Thorax: Dark, dorsum with inconspicuous prui- nose bands, katepisternum, anepisternum and ane- pimeron blue to green metallic lustrous; pubescence yellowish, rather even and short on dorsum and scu- tellum, longer, uneven on sides. Wings clear; halteres, squamae and antisquamae yellow. Legs: Dark, tarsi with contrasting golden pubes- cence on sides and dark upper faces; trochanters 3 smooth, femora 3 slightly swollen, triangular proces- sus normally shaped, distal margin much serrate; tib- iae 3 and tarsi 3 normal. Abdomen: Dark, slender, with conspicuous orange lateral spots on T II; T II-IV showing white conspic- uous narrowly interrupted arcuate pruinose bands; pubescence concolorous with surface, yellow on later- al spots; S brown, S IV darkest, flat, posterior margin entire. Genitalia: Anterior surstyle lobe large, more or less rectangular, rounded anteriorly, on margin with dense, yellow, even pubescence; the posterior surstyle lobe separated by deep sulcus, rather short, low, somewhat angular, showing dense, uneven, yellow pubescence. Cercus rather short, wide, with dense yellow pubescence. Aedeagus bearing paired humps basally on outer face, apical shaft part short, fringed plates on thecal apex recumbent to oblique. Body length 13 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons with lustrous midstripe occupying 0.3 of width. Thorax: Pruinose bands more conspicuous; pubescence denser, grey, even as if shaven. Legs: Pubescence greyish through- out. Abdomen: Outline more triangular; pubescence denser, greyish; pruinose bands wider, continuous; lateral spots extended over all of T II and anterior margin of T III; T V tapering, dark, with grey pubes- cence. Body length 13 mm. Diagnosis. — M. kaloceros can be distinguished from other Merodon except erivanicus and longicornis by the large antennal ratio and structure of the male 187 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 genitalia; it is distinguished from the two species mentioned by the coloration on the abdomen which is orange in kaloceros but crimson in erivanicus and longicornis. Period of flight and distribution (fig. 87). — M. kaloceros occurs in the eastern Mediterranean region and Turkey in June through August. Given the col- lection dates from eastern Turkey, where most of the type material was collected, the species is univoltine. Discussion. — In view of apomorphies shared with longicornis, e.g. the shape of the d genitalia and large antennal ratio, kaloceros is assigned to the longicornis group. Etymology. — The epitheton kaloceros is derived from the greek words kalon, meaning good, beautiful, and ceros, meaning horn; it refers to the large anten- nae (not as extremely long as in longicornis). The epitheton is to be treated as a noun in apposition. Merodon longicornis Sack (figs. 54 c, 55 b, 57 a-c, 93) Merodon longicornis Sack, 1913a: 447. Lectotype © (here designated): ‘longicornis det. Sack/ Merodon longicornis Sack Type Turkestan/ longicornis det. Hermann’ (NHMW) [examined]. Merodon longicornis, Paramonov 1925: 154, 1926a: 15; Van der Goot 1981: 214, 218; Peck 1988: 171. Lampetia longicornis, Sack 1931: 320. Material examined. — Greece: 26, 259 (KBIN, JLR, Coll. Thessaloniki, ZMAN, BMNH, NHMW); Italy: 306, 249 (RMNH, JLR, ZMAN); Israel: 12 (RMNH); Morocco: 19 (BMNH, doubtful); Syria: 12 (KBIN); Turkey: 16d, 249 (BMNH, JLR, NHMW); Yugoslavia: 42 (BMNH, RMNH). Description Male. — Head: Antennae dark, antenna 3 with up- per margin rather straight, apex rounded, antennal ra- tio 4.1 to 4.8; pubescence pale yellow, dark in large ocellar region; ocelli placed far forward; pruinosity on face and occiput dense, whitish; vertex angle 30°, ocellar angle 35°; tl-v ratio 0.3. Thorax: Dark; dorsum with pruinose bands, pu- bescence rather sparse, even, moderately long; surface of katepisternum, anepisternum and anepimeron weakly lustrous, with some silky lustre produced by yellow recumbent pubescence. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: Dark, joints paler in specimens; trochanters 3 smooth, femora 3 rather strongly swollen, straight; triangular processus and tarsi normally shaped. Abdomen: Dark; moderately slender; T II taper- ing; T II-IV showing conspicuous slightly arcuate white narrow pruinose bands; pubescence concolor- 188 ous with surface; S IV moderately vaulted, posterior- ly narrowly emarginate. Genitalia: The anterior surstyle lobe rather rectan- gular, with short, even, yellow pubescence, separated from the rather long, low posterior surstyle lobe by a deep sulcus; the posterior surstyle lobe posteriorly bearing moderately long, yellow, uneven pubescence, its lateral face with several ridges and furrows. Cercus rather large, well protruding, showing long, dense, slightly wavy, yellow pubescence. Aedeagus normally wide in ventral view, extremely slender in lateral view, apical shaft part short, fringed plates on thecal apex oblique. Body length 9-10 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Lustrous midstripe on frons occupying 0.6 of width; pubescence dark on midstripe, grey laterally. Thorax: Pubescence shorter than in male. Legs: Yellow pubescence on basitarsi 3 found in some 2. Abdomen: Strong crimson colora- tion always present in @, restricted to T II or extend- ed over most of abdomen; T II more tapering than in male; T V acute, dark, scattered pale pubescence present. Body length 9-10 mm. Diagnosis. — The extremely large antennal ratio distinguishes /ongicornis from all other known species. Period of flight and distribution (fig. 93). — The species occurs in May through July in the Mediterranean region, Turkey, Ukraine and southern Russia. It appears to be rare in the western, common in the eastern part of the range and in Turkey and adjacent areas. The vandergooti group Apomorphies: Lengthening of anterior surstyle lo- be; extreme swelling of femora 3; extensive yellow co- loration on legs. Merodon vandergooti sp. n. (figs. 58 a-c, 88) Type-material: 4 Holotype “Turkey, Hakkari, Suvarihalil pass 1250 m W side near Habul Deresı, 13.vi.1984 leg. J. A. W. Lucas’ (ZMAN). Paratypes: 20d; 183 with same data as holotype (16 JLR, 2 WH); 1d [Turkey] ‘Ankara 650 m, 1.vi.1971 leg. KI. Warncke’ (JLR); 16 [Turkey] ‘Ankara, South of Ankara, 24.vi.1984 leg. J. A. W. Lucas (LR). Description Male. — Head: Antennae conspicuously orange, an- tenna 3 with convex upper margin, apex acute, an- tennal ratio 1.5 to 1.7; pubescence golden yellow, rat- her dense, producing silky lustre on frons; face blue metallic lustrous, moderately dense white pruinose; vertex angle 40°, ocellar angle 40°, tl-v ratio 0.4. Thorax: Dark, with vestigial pruinose bands on dark dorsum; katepisternum, anepisternum and ane- pimeron weakly blue lustrous; pubescence golden on dorsum and scutellum, greyish-golden on sides; in many specimens some dark hairs present between wing insertions giving suggestion of vague intaralar band; posterior fringe of scutellum bearing dense yel- low pubescence. Wings clear; halteres, squamae and antisquamae yellow. Legs: Bases of femora 1 and 2 dark, femora 3 all dark, otherwise legs fiery orange throughout, with orange pubescence; trochanters 3 showing distinct ridge ending laterally in a low but marked hump; fe- mora 3 enormously swollen, rather strongly curved; triangular processus large, bearing 10-13 delicate bris- tles on hardly serrate distal margin. Abdomen: Dark; T II relatively large, tapering, bearing vague yellow lateral spots; T II-IV with oblique slightly arcuate, yellow pruinose bands, pu- bescence concolorous with surface; S IV moderately vaulted, narrowly and deeply emarginate posteriorly. Genitalia: Anterior surstyle lobe moderately wide, much elongate, the anterior part showing dense yel- low pubescence throughout, ventral margin similarly pubescent; separated from posterior surstyle lobe by deep wide sulcus; posterior surstyle lobe low, elon- gate, bearing dense, long, yellow pubescence on ven- tral and posterior margins; there is a secondary lateral lobe, bearing moderately long yellow pubescence. Cercus rounded, well protruding, with dense yellow pubescence. Aedeagus moderately slender with short apical shaft part and fringed plates on thecal apex re- cumbent. Body length 13-14 mm. Female. — Unknown Diagnosis. — M. vandergooti is easily separated from other Merodon species by the structure of the genital- ia and the enormously swollen femora 3; the colour pattern of the legs is very unusual as nearly all species with partially yellow legs have dark bands on the tib- iae. The colour pattern on the legs and orange anten- nae are also found in aureotibia females. In vandergoo- ti the antenna 3 is acute, in aureotibia obtuse; the abdominal pruinose bands are much more arcuate in aureotibia, where the triangular processus has a strongly serrate distal margin with 6-7 robust bristles; in vandergooti the triangular processus bears 10-13 fine bristles on a hardly serrate distal margin. These differences cannot be accounted for by explaining them as sexual dimorphism and are thus considered Hurkmans: Monograph of Merodon valid distinctions on the species level. Period of flight and distribution (fig. 88). — M. vandergooti occurs in June and has been found in cen- tral and south-eastern Turkey. Etymology. — It is a pleasure to dedicate this species to Mr Volkert van der Goot who, over the last decades, has largely stimulated research work on Syrphidae in the Netherlands. A noun in genitive case. The nigritarsis group Apomorphies: Aedeagus with apical shaft part lengthened; two pairs of spines on outer face of aede- agus. Description. — Head: Antennae dark, antennal ra- tio over 1.5, tl-v ratio about 0.4 in male; face bluish lustrous, strongest in male; in female, frons lustrous on median 0.25 of width.Thorax: Inconspicuous pruinose bands present; pubescence rather dense, short. Legs: Dark with yellow markings on tibiae, tar- si 1 and 2 in normal (not melanistic) specimens; tro- chanters 3 much variable between specimens, usually humped; golden pubescence on sides of tarsi 3 con- trasting dark upper face. Abdomen: Moderately slen- der; T IV bulging in male; pruinose bands arcuate, clear; S IV in male strongly vaulted, deeply emargi- nate posteriorly. Genitalia: Anterior surstyle lobe el- ongate, stalked, diamond-shaped, posterior surstyle lobe elongate; aedeagus with 2 pairs of chitinous spines on outer face and lengthened apical shaft part. Merodon femoratoides stat. nov. (figs. 59 a-d, 95) Merodon spinipes var. femoratoides Paramonov, 1925: 158. Lectotype d (here designated): ‘Karadagh bei Theodosia, Krym, 3.vi.1919° (ZMHB) [examined]. Merodon spinipes var. femoratoides, Liepa 1969: 20; Peck 1988: 174. i Lampetia spinipes var. femoratoides, Sack 1931: 328. Material examined. — Algeria: 16 (BMNH); Greece: 2d, 12 (LR); Turkey: 126 (LR, BMNH, NHMW, WH); U. S. S. R.: 2d Paralectotypes of spinipes var. femoratoides, ‘Warnutka, Sevastopol, Krym 11.vi.1923’ (ZMHB); 1d, Krym, Karadagh 24.vi.25 [1925] Paramonov leg./ Typus Merodon spinipes var. femoratoides 4 / B. Kurodke Pinob [in Russian script] (ZMHB) [This specimen cannot be consid- ered as a type specimen since it is not mentioned in the orig- inal publication]. Description Male. — Head: Antennae dark, antenna 3 with up- per margin concex, apex subacute, antennal ratio 1.5; pubescence yellow, face strongly white pruinose; ver- 189 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 tex angle 30°, ocellar angle 45°, tl-v ratio 0.4. Thorax: Dark, with four clear pruinose bands and a fifth anteromedian fainter one; pubescence dark yel- low, rather short, even; katepisternum, anepisternum and anepimeron blue lustrous. Wings slightly evenly troubled; halteres, squamae and antisquamae yellow. Legs: The tibiae basally and femora apically orange, otherwise dark; trochanters 3 smooth, femora 3 mod- erately swollen, triangular processus bearing 6-10 strong bristles on moderately serrate distal margin; tibiae 3 bearing dense golden pubescence, tarsi 3 with same pubescence on sides, contrasting dark upper face. Abdomen: Moderately slender; T II much taper- ing; T II-IV with arcuate off-white pruinose bands; T II bearing clear to almost absent yellow lateral spots which may extend over part of T III; pubescence con- colorous with background; S IV vaulted, acutely, nar- rowly emarginate posteriorly up to half its length. Genitalia: Anterior surstyle lobe elongate, clearly stalked, the apical portion triangular, anteriorly showing dense yellow pubescence; posterior surstyle lobe with small anterior, accessory lobe bearing dense, long, yellow pubescence; itself low and elongate, showing dense yellow pubescence posteriorly. Cercus inconspicuous and hidden between projecting poste- rior surstyle lobes. Aedeagus rather slender, bearing two pairs of spines on outer face, apical shaft part lengthened, fringed plates on thecal apex oblique. Body length 13-15 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons lustrous on median one-third, whitish pruinose laterally. Thorax: Pubescence as if shaven, golden; four well marked pruinose bands present. Abdomen: T II tapering, lat- eral spots conspicuous, extended over part of T III; pruinose bands arcuate, widely interrupted and con- spicuous. Body length 12.5 mm. Diagnosis. — M. femoratoides may be easily distin- guished from avidus by the very different genitalia in the & and the contrast of hair colour on tarsi 3 in both sexes. The distinction with nigritarsis is in the differently shaped anterior surstyle lobe in the males and the less dark abdomen with larger lateral spots in the females. Period of flight and distribution (fig. 95). — The species occurs in the Mediterranean, Asia Minor and southern Russia and Ukraine from May through July; its distribution range is large, but the species is un- common. 190 Merodon nigritarsis Rondani (figs. 60 a-g, 97) Merodon nigritarsis Rondani, 1845: 258, 264. Lectotype d (here designated): ‘68/ Torino D. Ghiliani’ in coll. Rondani (LSF) [the number 68 refers to the description of this species in Rondani’s catalogue] [examined]. Merodon nigritarsis, Rondani 1857: 55, 65; Schiner 1857: 415; Rondani 1873: 295; Strobl 1893: 76; Bezzi 1895: 16; Strobl 1900: 91; Sack 1913a: 447; Paramonov 1925: 157, 1926a: 15; Gil Collado 1930: 254; Peck 1988: 174. Lampetia nigritarsis Sack 1931: 327; Séguy 1961: 177. Material examined. — Austria: 46 (BMNH, NHMW, ZMUC); France: 966, 322 (MNHN, KBIN, RMNH, BMNH); Greece: 296, 192 (NHMW, ZMUC, den Hollander, KBIN, Bonn, ZMAN, RMNH, Thessaloniki, BMNH); Hungary: 2d, 42 (RMNH, Bonn, ZMUC); Italy: 38, 39 paralectotypes, ‘68’ in coll. Rondani (LSF); one 9 paralectotype, ‘68/ Susa, D. Ghiliani’ in coll. Rondani (LsF) and 136, 5® (LSF, KBIN, ZMAN, RMNH); Poland: 1 2 (BMNH); Spain: 98,52 (MNHN, BMNH, JLR, NHMW); Turkey: 96, 62 (LR, BMNH, NHMW, ZMAN, RMNH); Yugoslavia: 496, 199 (BMNH, NHMW, Bonn, ZMAN, RMNH). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apex subacute, antennal ratio 1.6; pubescence yellow with dark tuft in ocellar region; vertex angle 30°, ocellar angle 40°, tl-v ratio 0.4. Thorax: Dark, lustrous on lateral dorsum, scutel- lum and sides, bearing rather dense yellow pubes- cence. Wings clear; halteres, squamae and antisqua- mae yellow. Legs: All femora dark, paler apically; tibiae yellow with broad dark distal band; trochanters 3 with much variable hump, usually well marked, femora 3 slight- ly swollen, with normal triangular processus bearing 6-9 bristles on serrate distal margin; tibiae 3 and tarsi 3 normally shaped; the basitarsi 3 with strong con- trast between dark pubescence on upper face and golden pubescence on sides; this is a characteristic that can be of use in the field as it is conspicuous to the naked eye at short range. Abdomen: Dark; T II with orange to yellow lateral spots of variable size and clearness; T II] in many spec- imens with yellow anterior lunules; T II-IV bearing ar- cuate, narrowly interrupted pruinose bands; S IV very deeply, narrowly emarginate, strongly vaulted. Genitalia: Anterior surstyle lobe elongate, widened subapically, more or less rounded apically, with dense yellow even short pubescence there; posterior surstyle lobe separated by small sulcus; rather elongate, mod- erately high, posteriorly bearing dense, long, yellow pubescence. Cercus rather inconspicuous, rounded, pubescence as on posterior surstyle lobe. Aedeagus bearing two pairs of spines on outer face; apical shaft part clearly lengthened; fringed plates on thecal apex recumbent. Body length 12-16 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons lustrous on median quarter, sides white pruinose, pubescence concolorous; dark tuft in ocellar region less clear than in male; ocellar angle 45 to 50°. Thorax: Shorter pu- bescence; pruinose bands clearer. Legs: Trochanters 3 always smooth. Body length: 12-17 mm. Variation. — Merodon nigritarsis is one of the more variable species of the genus; the hump on the tro- chanters 3 and the shape of the anterior surstyle lobe are notoriously variable between specimens, even those collected together at the same site. There are many local forms, each with their own peculiar aber- rations, notably in the shape of the anterior surstyle lobe. Since a fluent, continuous series can be arranged using these specimens, I have refrained from describ- ing local subspecies. Diagnosis. — M. nigritarsis can be distinguished from other Merodon except femoratoides in the males by the general morphology of the genitalia and the spines on the aedeagus and the contrast on the basi- tarsi 3 in both sexes; from femoratoides it may be dis- tinguished in the males by the differently shaped an- terior surstyle lobe and basally convex aedeagus, smaller lateral spots and less wide interrupted prui- nose bands; in the females by the darker abdomen. Period of flight and distribution (fig. 97). — M. nig- ritarsis occurs in the Mediterranean and the southern parts of central Europe from May through August; possibly the species is bivoltine. Discussion. — M. nigritarsis has been considered a variety or subspecies of avidus. In view of the com- pletely different genitalia a close relation between av- idus and nigritarsis is out of the question. M. nigritar- sis was named for the dark basitarsi 3; however, this feature also occurs in other species externally resem- bling nigritarsis, such as alagoezicus and its allies; the lengthened apical shaft part of the aedeagus is another shared feature. The avidus group Apomorphies: Anterior surstyle lobe anteriorly much widened; T II extremely tapering. Merodon avidus (Rossi) (figs. 61 a-g, 96) Syrphus avidus Rossi, 1790: 292. Syntypes: ‘in provinciis Hurkmans: Monograph of Merodon Florentina et Pisana’ [Toscana, Italy] [not examined]. Syrphus spinipes Fabricius, 1794: 296. Lectotype d (here designated): ‘spinipes in coll. J. C. Fabricius (ZMUC) [seri- ously damaged] [examined] Merodon serrulatus Wiedemann in Meigen, 1822: 360. Lectotype d : (here designated) ‘Portugall coll. Winthem/ serrulatus Wied.’ (NHMW) [examined] Merodon italicus Rondani, 1845: 257, 259. Lectotype 9 (here designated): ‘67b’ in coll. Rondani (LSF); the ‘67b’ number refers to the description of ttalicus in Rondani’s catalogue. [examined] syn. n. Merodon rufitibius Rondani, 1845: 259, 265. Lectotype d (here designated): ‘72’ in coll. Rondani (LSF) [examined]; the ‘72’ number refers to the description of rufitibius in Rondani’s catalogue. Merodon graecus Walker, 1852: 243. Lectotype ? (here des- ignated): ‘Albania/ graecus Walker’ (BMNH) [examined]. Merodon aurifer Loew, 1862a: 83. Lectotype d (here desig- nated): ‘Melatia [Malatya, Turkey] Erber/ coll. H. Loew/ Zool. Mus. Berlin’; on separate pin: ‘... Merodon aurifer n. Berl. Zeit. vi.1862 pg 82 ...’ (ZMHB) [examined]. Merodon quadrilineatus Lioy, 1864: 473. [Type-material probably never existed. Lioy published only theoretical works and had no intention to describe new species.] Merodon avidus, Rondani 1845: 259; Schiner 1857: 415; Strobl 1893: 76; Bezzi 1895: 16, 1900: 91; Strobl 1900: 593; Villeneuve 1903: 115; Gil Collado 1930: 254; Van der Goot 1971: 106, 1981: 216, 218; Barendregt 1982: 80; Marcos Garcia 1985: 196; Bradescu 1986: 123; Peck 1988: 167; Hurkmans 1988: 107. Lampetia avida; Séguy 1961: 176; Van der Goot 1964a: 431. Milesia spinipes, Latreille 1804: 331. Merodon spinipes, Fabricius 1805: 197; Meigen 1822: 361; Macquart 1828: 295, 1834: 516, 1842: 70; Rondani 1845: 258, 264, 1857: 55, 66; Schiner 1857: 414; Strobl 1893: 76; Bezzi 1895: 16, 1900: 91; Strobl 1900: 593; Villeneuve 1903: 115; Czerny & Strobl 1909: 204; Sack 1913a: 444; Paramonov 1925: 146, 1926a: 15; Gil Collado 1929: 409; Gil Collado 1930: 243; Bankowska 1980: 38, 40, 83; Marcos Garcia 1985: 137; Peck 1988: 174. Lampetia spinipes, Oldenberg 1919: 388; Sack 1930: 96, 1931: 327, 1938: 21; Suster 1959: 243; Séguy 1961: 176. Merodon serrulatus; Schiner 1857: 413, 1862: 348; Strobl 1893: 76; Bezzi 1895: 16; Strobl 1900: 593; Czerny & Strobl 1909: 203, 204; Gil Collado 1930: 254. Lampetia serrulata; Sack 1931: 322. Merodon italicus, Rondani 1857: 54, 62; Schiner 1857: 413, 1862, 347. Lampetia italica; Sack 1931: 322. Merodon rufitibius, Rondani 1857: 55, 66; Palma 1863: 46; Becker 1907: 253. Lampetia rufitibia; Sack 1931: 327. Merodon graecus, Schiner 1857; Schiner 1862: 347; Van der Goot 1964b: 218; Peck 1988: 167. Merodon aurifer, Peck 1988: 174. Lampetia aurifera; Sack 1931: 327. Merodon quadrilineatus, Peck 1988: 174. Lampetia quadrilineata; Sack 1931: 327. Material examined. — Albania: 1d, 19 (NHMw); Algeria: 26, 242 (MNHN, KBIN); Austria: 534, 632 (NHMW, MNHN, BMNH, RMNH); Belgium: 3d (MNHN, KBIN); Cyprus: 136, 49 (KBIN, BMNH); Czechoslovakia: 16, 19 (BMNH, 191 TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 JLR); France: 2666, 2109 (MNHN, BMNH, NHMW, RMNH, JLR, ZMAN, WH); Germany: 36 (MENS, JLR); Greece: 966, 622 (BMNH, RMNH, NHMW, KBIN, MNHN, JLR, den Hollander, ZMUC, Bonn, Coll. Thessaloniki, WH); Hungary: 293, 172 (Bonn, ZMUC, NHMW, BTM); Italy: 29 Paralectotypes of ztalicus ‘67b’ in coll. Rondani (LSF); 2d, 7% Paralectotypes of rufitibius ‘72’ in coll. Rondani (LSE); 3196, 1229 (ZMUC, KBIN, NHMW, JLR, RMNH, ZMAN, BMNH, LSF, Coll. Marogna (Verona, Italy), WH); Lebanon: 19 (KBIN); Lybia: 48, 22 (BMNH); Morocco: 56, 89 (BMNH, MNHN, JLR, NHMW); Netherlands: 6d, 22 (RMNH, JLR); Poland: 1d (ZMUC); Romania: 26, 12 (BMNH); Spain: 708, 209 (ZMUC, ZMAN, RMNH, JLR, WH); Switzerland: 56, 62 (RMNH, ZMAN, JLR); Turkey: 576, 302 (LR, WH, BMNH,NHMW); U. S. S. R.: 4d, 82 (MNHN); Yugoslavia: 706, 409 (NHMW, BMNH, RMNH, JLR, MNHN, Bonn, ZMAN). Description Male. — Head: Antennae with antenna 3 showing subacute apex and convex upper margin, antennal ra- tio 1.5, pubescence yellow, dark hair-tuft often pre- sent in ocellar region; face rather densely white prui- nose, partially obliterating blue lustrous surface; occiput concolorous; vertex angle 35°, ocellar angle 40°-45°, tl-v ratio 0.5. Thorax: Moderately wide, dark, rather dulled by diffuse yellowish pruinosity, dorsum bearing longitu- dinal pruinose bands of most variable clearness, kate- pisternum, anepisternum and anepimeron very weak- ly to conspicuously blue lustrous; pubescence erect, yellowish, moderately dense. Wings clear, veins pale to dark brown; halteres, squamae and antisquamae yellow. Legs: Dark on trochanters and femora, pale on tib- iae and tarsi, but both ends of femora paler and dark band present on tibiae is the usual pattern; there is an immense variation resulting in relatively pale or all dark legs; the femora always mainly dark; trochanters 3 may be acuminate; femora 3 moderately swollen, bearing 6-9 bristles on serrate distal margin of trian- gular processus. Abdomen: Slender; dark; T II much tapering, usu- ally with clear yellow lateral spots; pruinose bands on T II-IV clear, arcuate, interrupted, of variable width and colour; pubescence often concolorous with sur- face, erect, longest anterolaterally; S IV strongly vaulted, deeply acutely emarginate posteriorly. Genitalia: Anterior surstyle lobe stalked, widening to semicircular apical margin with short even yellow pubescence; posterior surstyle lobe moderately high, slightly elongate, with moderately dense, long, yellow pubescence. Cercus rather large, bearing dense, long, yellow, often wavy pubescence. Aedeagus moderately slender, apical shaft part short, pipes plates on the- cal apex recumbent. Body length 11.5-18 mm. 192 Female. — Except for sexual dimorphism, differing ferom the males as follows. Head: Lustrous midstripe on frons occupying 1/4 to 3/4 of width, the lateral pruinose strips usually well defined; face and occiput less lustrous. Thorax: Pruinose bands clearer, slightly wider. Abdomen: Lateral spots often reddish, con- spicuous and extending over part of T III; in some cases however virtually absent; pruinose bands less clear, less wide. T V dark. Body length 12-17 mm. Variation. — There is much variation in the colora- tion in both sexes. Head: The antennae may range from black to bright yellow. Thorax: Lustre most var- iable, density and colour of pubescence more con- stant. Legs: The colour is extremely variable; all joints, the tarsi and both ends of the tibiae are paler than the rest of the legs, but the colour may be all black or all yellow throughout. The pubescence is whitish to yellow even in dark specimens. Abdomen: The extent of the yellow colour is variable; the fe- males often show a deeper colour than the males, and are less variable in this respect. Genitalia: The male genitalia are rather constant. The main variation oc- curs in the length and density of the pubescence on the posterior surstyle lobe and the cercus. The large degree of variation found in avidus is not randomly distributed over the specimens. Many series collected at one site on one date, show uniform char- acteristics. Size and colour are probably correlated with quantity and quality of food ingested during the larval stages. There is no clear trend in size, morphol- ogy or colour of the various local forms over the en- tire range. Therefore, to present descriptions of local forms, varieties or subspecies is considered inappro- priate. Diagnosis. — M. avidus males could possibly be confused with toscanus from which they are distin- guished by the longer stalk of the anterior surstyle lobe (which in zoscanus is bifid), the shorter apical shaft part of the aedeagus and the usually much dark- er abdomen. The unique genitalia are sufficient to distinguish avidus males from any other Merodon males at first sight. The females are posing many problems since they cannot always be separated from females of the species in the alagoezicus group. The best separating characteristic is the pruinosity on the thorax where avidus females always show four longi- tudinal bands, whereas in the females of the alagoezi- cus group there usually is a fifth, anteromedian band as well. Moreover, the abdomen of females in the a/- agoezicus group is usually narrower overall. Period of flight and distribution (fig. 96). — M. av- idus occurs in Europe, except the northern parts, in north Africa, the Middle East and Asia Minor from March through October; the species is very likely multivoltine in the southern parts of its range (cf. Marcos Garcia, 1985). The distribution does not ap- pear to be associated with human activity. As avidus is not particularly easy to collect and seems to occur in rather the same habitats as other Merodon the relative abundance in collections probably reflects abundance in the field. Biology. — Few details are known (if the great dis- tribution and abundance is borne in mind). This sec- ond most common Merodon occurs both in generally wet, and in dry areas, such as the Vosges mountains of France and the inland steppe region of Turkey. There are indications that in the driest parts of its range av- idus prefers relatively wet biotopes such as lush strips along streams or marshes. Moreover imagines occur early in the season in the hot, dry parts of its range, probably to avoid water shortage later on. Water shortage influences the vegetation and therefore the possibilities for the larvae which are probably phy- tophagous (all known Merodon larvae are phytopha- gous). There seems to be a preference for flowers of Umbelliferae, as recorded by Suster (1959) and con- firmed by Mr. Lucas’ s (Pers. Comm.) and the author’s observations in e.g. France, Italy, Greece and Turkey. Some notes on the behaviour of the species are provided by Hurkmans (1988). Discussion. — No type-material of avidus has been found; it might be supposed to be in the Rossi collec- tion at ZMHB (Horn & Kahle 1936), but is not cur- rently present there (dr. H. Schumann, in litt.). The types of spinipes are mentioned purely formally as they are too much decayed to be of any value what- soever. There are considerable difficulties in distin- guishing avidus from externally similar species, if the genitalia are not used for identification. The existence of the type-specimens in their present state does re- grettably forestall designation of a neotype, which would seem to be necessary, given the identification problems mentioned. The original description by Rossi (1790) and following emendations are detailed enough to define avidus with respect to the external characteristics. No examination of the genitalia had been made until comparatively recently; especially in avidus the use of genital characters for separation from externally resembling species is essential. It is however considered fair to rule that as the over- whelming majority of museum specimens identified as avidus examined fulfil the requirements of the orig- inal description these specimens must be regarded as the true avidus. Moreover most other species external- ly resembling avidus can be (and have been) separated by external characteristics as well, e.g. nigritarsis Rondani and alagoezicus Paramonov. Hurkmans: Monograph of Merodon The crassifemoris group Apomorphies: Posterior surstyle lobe very high, with large anteriorly projecting lobe; cercus reduced. Merodon crassifemoris Paramonov stat. n. (figs. 62 a-c, 95) Merodon spinipes (Fabricius) var. crassifemoris Paramonov, 1925: 158. Holotype d: ‘Theodosia, Krym, 13.vi.1923 [Crimea, U. S. S. R.] [not examined]. Merodon spinipes var. crassifemoris, Liepa 1969: 20; Peck 1988: 174. Lampetia spinipes var. crassifemoris, Sack 1931: 328. Material examined. — France: 16 (ZMAN); Greece: 26 (ZMAN); Turkey: 88 (WH, JLR); U. S. S. R.: 16, labelled ‘Karadagh Krym 16 vii 1929 S. Paramonov. leg. / Typus Merodon spinipes Fbr. var crassifemoris var. nov. d | v. Karodke Pinab’ (wrongly labelled as a type!) (ZMHB); Yugoslavia: 16 (ZMAN). Description Male. — Head: Antennae dark, antenna 3 with up- per margin convex, apex very obtuse, antennal ratio 1.5, vertex angle = 30°, ocellar angle = 40°, tl-v ratio 0.5; pubescence creamy white with dark tuft in ocel- lar region, face dense whitish pruinose, compound eyes bearing dense white pubescence. Thorax: Dark, metallic blue lustrous on lateral dor- sum and scutellum, katepisternum, anepisternum and anepimeron, longitudinal pruinose bands vesti- gial to absent; pubescence dark yellow to brownish with vague dark interalar band. Wings clear; halteres, squamae and antisquamae yellow. Legs: All dark except joints; trochanter 3 angular, femora 3 markedly curved and swollen, with smallish rather steeply rising triangular processus bearing 6-10 bristles on weakly serrate distal margin; tibiae 3 with distal outer angle drawn out into small spur; tarsi 3 with golden pubescence on sides contrasting dark pu- bescence on upper face, as in nigritarsis. Abdomen: Dark; T II with vague brownish lateral spots, distinctly tapering; T II-IV bearing arcuate conspicuous greyish interrupted pruinose bands; pu- bescence yellowish grey on bands, dark elsewhere; S IV keeled throughout, posteriorly deeply emarginate. Genitalia: The anterior surstyle lobe relatively small, reniform, showing dense, short, yellow, even pubescence apically and on margins; coalescent with enormously elongate posterior surstyle lobe which bears rather dense, yellow, uneven pubescence ven- trally; posterior part of posterior surstyle lobe with free anteriorly projecting cuneiform part, with dense yellow pubescence laterally and postero-ventrally. Cercus very small, bearing dense, long, yellow pubes- cence. Aedeagus rather stout, apical shaft part short, fringed plates on thecal apex oblique to erect. 193 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Body length 12-14 mm. Female. — Unknown. Diagnosis. — M. crassifemoris can be easily separat- ed from other Merodon by the anteriorly projecting posterior surstyle lobe; externally the species resem- bles avidus, nigritarsis, femoratoides and the species in the alagoezicus group. Period of flight and distribution (fig. 95). — The species occurs in the Mediterranean and Black Sea re- gions in June and July. Discussion. — Because type-material of crassifemor- is was not available, identification of this species is largely based on the original description and the spe- cimen in ZMHB, identified as crassifemoris by Pa- ramonov himself. Paramonov described var. crassif- emoris as a variety of spinipes Fabricius (= avidus Rossi) in view of several external similarities. The genitalia however are rather different and do not indi- cate a close relationship between avidus and crassifem- oris. The elegans group Apomorphy: Thin chitinous flanges running along outer face of aedeagus. Description. — Head: Antennae orange to dark, an- tennal ratio 1.5-1.8, vertex angle 25-35°, ocellar angle 35-55°, tl-v ratio 0.35-0.6; pubescence white to yel- low. Thorax: Pubescence rather short, even; whitish to pale brown. Legs: Dark, usually with yellow mark- ings on apices of femora and at both ends of tibiae; trochanter 3 often edged; femora 3 swollen, strongly curved. Abdomen: Slender to rather stout, T II strongly to moderately tapering; T II-IV bearing ar- cuate pruinose bands, lateral spots variable. Genitalia: The anterior surstyle lobelarge, elongate with concave dorsal margin except manicatus, anterior and posteri- or surstyle lobe separated by sulcus; aedeagus with chitinous flanges on outer face. Merodon bequaerti sp. n. (figs. 63 a-c, 98) Type-material: & Holotype ‘Noiseux Oran Algeria Dr J Bequaert, 23.1V.10’ (written on lower face) / Merodon parie- tum Mg, 3” (MNHN). 19 paratype, with label as holotype (MNHN). Description Male. — Head: Antennae dark, antenna 3 somew- hat paler, densely yellowish pruinose, upper margin slightly concave, apex acute, antennal ratio 1.8; vertex 194 angle 30°, ocellar angle 40°, tl-v ratio 0.6; pubescen- ce pale yellow with strong dark tuft in ocellar region, occiput deep yellow pubescent; face rather bald, black-bluish lustrous; compound eyes with rather sparse, pale, yellow, short pubescence. Thorax: Rather dull, some bluish lustre on sides; pubescence even, short, pale brown, erect. Wings tinged evenly, halteres, squamae and antisquamae yel- low. Legs: All dark; the tarsi, especially tarsi 3 with dense recumbent golden pubescence; trochanters 3 with ridge on lower face, femora 3 quite swollen, rather curved, with triangular processus bearing 10 bristles; tibiae 3 rather slender, tarsi 3 normally shaped. Abdomen: Dark, moderately slender, with hardly conspicuous interrupted arcuate whitish pruinose bands; T II somewhat tapering, with orange lateral spots; pubescence rather dense, orangish brown (par- tially worn off on holotype); T III-IV laterally with cupreous lustre; all S dark, bearing scattered pale erect pubescence; S IV slightly vaulted, its posterior margin entire. Genitalia: The anterior surstyle lobe ovoid, bearing yellowish pubescence anteriorly and along ventral margin and showing ventro-posterior angle; deep sul- cus present between surstyle lobes, connected by a stalk, visible as a broad chitinous plate in top view; the posterior surstyle lobes square, with strong cen- tro-posterior and lesser ventral accessory lobe; pubes- cence golden, longest, erect on ventral margin and lobes; the centro-posterior lobe bears a single black bristle. Cercus short, wide, showing pale, yellow, long, erect pubescence. Aedeagus with outer face ex- cavate subbasally on outer face, there with paired free chitinous plates, and bearing paired flanges running along all of the outer face; apical shaft part slightly lengthened, fringed plates on thecal apex suberect to recumbent. Body length 13.5 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Antenna 3 rounded apically, though clearly concave on upper margin; frons dark, slightly greenish metallic lustrous, sparse- ly pubescent (probably artifact; head has been glued on to the body at some time) ocellar angle 50°. Thorax: Pubescence shorter than in male. Legs: Trochanters 3 smooth. Abdomen: Less dense pubes- cent except on lateral spots; T II less tapering; T V dark, tapering. Body length 13 mm. Diagnosis. — M. bequaerti can be easily recognized by the stout build of both sexes in comparison with other species of the e/egans group. The females may pose problems but the combination of swollen femo- ra 3 and arcuate pruinose bands on the abdomen will separate this species from e.g. ruficornis and its allies which have more conspicuous, less arcuate pruinose bands. Period of flight and distribution (fig. 98). — This species is known only from the types, collected in Algeria in April. Etymology. — The epitheton bequaerti has been de- rived from the collector s name on the labels of both type specimens. À noun in genitive case. Merodon elegans sp. n. (figs. 64 a-c, 99) Type-material: 4 Holotype ‘Italia Sicilia V. S. v. d. Goot/ Etna rif. Filiciusa 1400-1500 m 22-28.vii.1961 / Lampetia spinipes det. V. S. v.d. Goot 1963 / Type A’ (ZMAN). — Paratypes: 52d, 249; 198, 10? with same data as holo- type but collectors V. S. v.d. Goot & J. A. W. Lucas (14d JLR, 58, 102 RMNH); 16 ‘Italia, S. Castaldo vi.1937, A. Loken leg.’ (ZMUC); 38 ‘Sicilia Schiodte’ (ZMUC); 16 ‘Italia, Calabria 18.v.1954 F. F. Tippmann’ (RMNH); 1d ‘France Aveyron 19.vii.1980 D. Prins’ (Coll. D. Prins, Rhenen, The Netherlands); 38 ‘France Vaucluse Cheval Blanc 7.vii.1973 H. I. Meuffels (JLR); 19, same data, col- lected 9.vii.1973 LR); 28 ‘France Estagel 22.vi.1982 exc. LH Wageningen’ (LHW); 2d ‘France, Chateau de Peypertuse 25.vi.1982 exc. LH Wageningen (LHW); 1d ‘France, Mts. de Tauch, exc. LH Wageningen 23.vi.1982’ (LHW); 1d ‘S. F. Chaines des Alpilles, Les Baux 21.vii.1980 Barkemeyer (Coll. C. Claussen, Flensburg); 3d ‘Espana Torla Huesca 1035 m, 8/26 vii.1974 J. Wolschrijn’ (JLR); 54 Espana Valencia Moraira 16/30.v.1981 van Aartsen (3 JLR, 1 RMNH, 1 ZMAN); 42 ‘Espana, Guadalajara, Molina 1100 m 11.vii.1977 W. Schacht (LR); 19 ‘Espana Tragacete 1300 m, 17.vii.1974 W. Schacht’ (LR); 2d ‘Espana Catalonia, Espluga de F., 3.vii.1920 Codina’ (RMNH); 1d ‘Espana Catalonia Carretera [=road] de Espluga de F. a Massies’ (RMNH); 1d, 19 ‘Espana Teruel M. J. & J. P. Duffels / Rafales 35 km ZO v. Alcaniz 600 m 12.vii.1972° (ZMAN); 38 ‘Espana, Lebena (Santander) 5.vii.1986 Ma. A. Marcos Garcia’ (SAL); 16, 19 “Algerie, Alger Dr. Bequaert, Foret de Buinen (MNHN); 1d ‘Algerie, Alger Dr. Bequaert Benandreit’ (MNHN); 1d ‘Algeria, Le Tarf 27.vii.1896 A. E. Eaton’ (BMNH); 38 ‘Maroc Meh- dia (Ft. Lyautey) 31.v 1.vi.1950 P. M. F. Verhoeff 3d (- RMNH). Description Male. — Head: The antennae orange to brown, an- tenna | darkest, antenna 3 with upper margin slight- ly convex, apex obtuse, antennal ratio 1.6; pubescen- ce dull yellow, erect, even, with rather large dark tuft in ocellar region; vertex angle 30°, ocellar angle 50°, tl-v ratio 0,6; face and occiput diffusely whitish prui- nose, blue metallic lustrous. Thorax: Dark, with inconspicuous longitudinal Hurkmans: Monograph of Merodon pruinose bands; lateral dorsum and scutellum as well as sides blue metallic lustrous; pubescence brownish to yellow, erect, short, rather even, a dark interalar band present in some specimens; pubescence on pos- terior margin of scutellum longer; some diffuse prui- nosity on dorsum and sides. Wings clear to evenly slightly tinged throughout; halteres, squamae and antisquamae yellow. Legs: Dark with yellow markings apically on femo- ra, both ends of tibiae and most of tarsi; trochanter 3 smooth, femora 3 quite swollen, metallic lustrous, with normally shaped triangular processus bearing 6- 9 bristles on moderately serrate distal margin. Abdomen: Dark; T II strongly tapering, with large pale yellow to deep orange vaguely bordered lateral spots; pruinose bands white, interrupted, moderately wide, arcuate; pubescence yellowish, densest laterally; S IV rather vaulted, narrowly emarginate posteriorly. Genitalia: The anterior surstyle lobe large, reni- form, with short, dense, even, yellow pubescence; separated from posterior surstyle lobe by deep sulcus; posterior surstyle lobe high, large, bearing rather long, dense, yellow pubescence laterally and on ven- tral-posterior margin, with lateral accessory lobe bear- ing dense concolorous pubescence. Cercus angular, well protruding, densely long yellow pubescent. Aedeagus rather slender, with paired high chitinous flanges on outer face and bearing paired basal humps; apical shaft part short, fringed plates on thecal apex oblique to recumbent. Body length 13-15.5 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons with lustrous midstripe occupying less than one-third of width; clear frontal depression of ovoid outline present. Thorax: Pruinose bands clearer, in some specimens a fifth anteromedial band can be found. Legs: Orange markings on basal femora more extensive, femora 1 and 2 often apically widely yellow. Abdomen: T II conspicuously bright orange; pruinose bands on T III IV continuous; pubescence sparser than in male. Body length 13-15 mm. Diagnosis. — The conspicuous flanges on the ae- deagus distinguish e/egans from species outside the elegans group. M. elegans may be separated from be- quaerti by the longer posterior surstyle lobe, the slen- derer habitus and greater size; from manicatus by the different shape of the surstyle and possession of an ac- cessory lobe on the posterior surstyle lobe in elegans, from testaceus also in the surstyle shape, the height of the aedeagal flanges, the overall size and the absence of typical red abdominal coloration as in testaceus. The females of elegans are similar to the females of the alagoezicus group, avidus and many other species, 195 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 from which they can be distinguished by their frontal depression. Period of flight and distribution (fig. 99). — M. ele- gans occurs from May to July in the western part of the Mediterranean. Discussion. — Van der Goot (1964a) noticed differ- ences among sets of specimens then considered to be- long to avidus, and distinguished between types A and B. Examination of the genitalia showed that type B corresponded to avidus whereas type A was shown to be a new species, described here. The flight period coincides with that of avidus in the mediterranean re- gion; the distribution of e/egans is much more restrict- ed than that of avidus and seems to be Mediterranean. As the species are very similar externally (elegans has longer, rougher pubescence on the thorax and more metallic lustre on face and femora 3, which is more swollen than in avidus) and their flight period over- laps, some of the avidus records in literature probably pertain to elegans. Etymology. — The adjective elegans is latin, mean- ing well-formed, and refers to the slender habitus of the species. Merodon manicatus (Sack) (figs. 65 a-c, 98) Lampetia manicata Sack, 1938: 21. Holotype 6: ‘Stip, v.1937, leg. R. Meyer/ Lampetia manicata Sack det. Sack’ (HLMD) [examined]. Lampetia manicata; Sack 1938: 21. Merodon manicatus, Peck 1988: 171. Material examined. — Macedonia: 9 Paratype with same labels as holotype (HLMD). Description Male. — Head: Antennae dark, antenna 3 ovoid, upper margin strongly convex, apex obtuse, antennal ratio 1.6; pubescence pale yellow with some dark pu- bescence in ocellar region; vertex angle 30°, ocellar angle 55°, tl-v ratio 0.35; face rather densely white pruinose; compound eyes bearing rather short, dense, white pubescence. Thorax: The dorsum and scutellum showing weak olivaceous lustre laterally; same lustre stronger on kat- episternum, anepisternum and anepimeron; longitu- dinal pruinose bands weak; pubescence golden, erect, moderately dense, even, slightly longer and paler lat- erally. Wings tinged overall; halteres, squamae and antisquamae pale yellow. Legs: Dark, with some olivaceous lustre on femora 3; tibiae with yellow markings at both ends; tarsi dark, but with recumbent dense golden pubescence 196 on lower face, strongest on tarsi 3; trochanter 3 with distinct sharp ridge, femora 3 strongly swollen, much curved, bearing normally shaped triangular processus bearing 7 bristles on moderately serrate distal margin. Abdomen: Dark; T II with large orange lateral spots, continued on T III; T II-IV showing rather faint white pruinose arcuate bands best observed from behind, hardly interrupted medially; all S orange, bearing erect white pubescence; S IV darker, slightly keeled, deeply narrowly emarginate posteriorly; ab- dominal pubescence in general very sparse compared to other Merodon, concolorous with surface and very short medially. Genitalia: The anterior surstyle lobe reniform, bearing dense short even yellowish pubescence anteri- orly and ventrally, separated from the posterior sur- style lobe by a moderately deep sulcus; the posterior surstyle lobe with yellow pubescence, bearing distinct anteromedial accessory lobe with pubescence erect and long; main disc of posterior surstyle lobe bearing shorter pubescence, somewhat diamond-shaped. Cercus angular, with same pubescence as posterior surstyle lobe. Aedeagus moderately slender, bearing paired translucent, twice interrupted (artifact?) chi- tinous flange on outer face, apical shaft part slightly lengthened, fringed plates on thecal apex oblique; paired humps present basally on outer face. Body length 11.5 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons with lustrous midstripe occupying rather less than a quarter of the frontal width just above the antennae, but wider pos- teriorly; ocellar angle 60°. Thorax: Pruinose bands more clear due to shorter, less dense pubescence; sides much less lustrous than in male. Legs: The femora 3 less swollen, less curved, less lustrous. Abdomen: Predominantly orange; T V with dark median zone; pruinose bands widely interrupted; pubescence very sparse, even shorter than in male. Body length 11 mm. Diagnosis. — The males manicatus are distinguished from those of bequaerti by the much slenderer habit- us and more rounded anterior surstyle lobe as well as the sparse pubescence; from those of elegans also by the rounded anterior surstyle lobe, and much smaller size; from those of testaceus by the less slender habitus of manicatus and its less tapering T II, and the more rounded anterior surstyle lobe; from all other species by the aedeagal flanges. The females of manicatus are slenderer than those of elegans and bequaerti, but are hard to separate from those of testaceus, they can how- ever be distinguished by the more acute ocellar trian- gle with ocellar angle 60°, (70° in testaceus). From all other species they differ in the combination of ex- tremely slender build, moderate size and red abdomi- nal colour, which is of a quite different type than found in the longicornis group where the colour is deep crimson. Period of flight and distribution (fig. 98). — The species occurs in Macedonia in May, and is currently known only from the types. Merodon testaceus Sack (figs. 66 a-c, 98) Merodon testaceus Sack, 1913a: 445. Lectotype d (here des- ignated): “Asia minor/ Alte Sammlung/ Merodon testaceus type Sack/ testaceus det. Sack’ (NHMW) [examined]. Merodon testaceus, Paramonov 1925: 154; Peck 1988: 175. Lampetia testacea; Sack 1931: 329, 1938: 21. Material examined. — 3d, 19 Paralectotypes, ‘Asia Minor/ idae det. Schiner/ Alte Sammlung/ testaceus det. Sack’ (NHMw); 29 Paralectotypes, ‘Mann, Brussa [=Bursa, Turkey], 1863/ idae det. Schiner/ Alte Sammlung/ testaceus det. Sack’ (NHMW). Description Male. — Head: Antennae orange to brown, antenna 3 with upper margin convex, apex obtuse, antennal ratio 1.8; vertex angle 25°, ocellar angle 35°, pubes- cence pale yellow, tl-v ratio 0.35. Thorax: Dark, with pruinose bands inconspicuous; dorsum and scutellum dark, katepisternum, anepis- ternum and anepimeron blue metallic lustrous, pu- bescence rather sparse, short, whitish. Wings clear, halteres, squamae and antisquamae yellow. Legs: The femora dark, tibiae dark with yellow markings at both ends, tarsi brown to yellow, in some specimens with faint dark band; trochanters 3 with conspicuous edge, femora 3 strongly swollen, curved, with normally shaped triangular processus bearing 6 stout bristles on rather weakly serrate distal margin with relatively large apical spurlet. Abdomen: Slender; T II strongly tapering; pre- dominantly orange to reddish brown, with clear arcu- ate narrowly interrupted white pruinose bands on T II-IV; S IV slightly keeled, deeply emarginate posteri- orly. Genitalia: The anterior surstyle lobe bearing dense, short, yellow, even pubescence apically and on margins, with incision on dorsal margin, ridged later- ally, separated from the posterior surstyle lobe; the posterior surstyle lobe with anterior accessory lobe bearing dense long yellow pubescence; main disc rather elongate, protruding far posteriorly, bearing rather dense yellow pubescence posteriorly. Cercus angular, with dense long yellow pubescence. Aedeagus moderately slender, bearing large paired humps subbasally on outer face as well as paired chi- tinous flanges; apical shaft part short, fringed plates Hurkmans: Monograph of Merodon on thecal apex recumbent. Body length 10 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Frons black, lustrous on median one-third, sides diffusely whitish prui- nose, pubescence concolorous, ocellar angle 70°. Thorax: Pubescence notably short, pruinose bands more conspicuous.Legs: Femora 3 less swollen. Abdomen: Slightly stouter than in male, orange to brown throughout, pruinose bands slightly less con- spicuous. Body length 10.5 mm. Diagnosis. — The males of testaceus can be distin- guished from those of bequaerti by the less stout hab- itus, from males of elegans and manicatus by the low- er aedeagal flanges, and from those of elegans by smaller size as well. Moreover, they differ from males of manicatus also by slenderer abdomen, less rounder antenna 3 and shorter thoracic pubescence. From all other Merodon they differ by the aedeagal flanges. The females can be distinguished by their red abdom- inal coloration, size and slender habitus from all spe- cies. The differences with females of manicatus are: the larger ocellar angle (70° here, 60° in manicatus) and the slightly greater antennal ratio (1.8 versus 1.6 in manicatus). Period of flight and distribution (fig. 98). — M. tes- taceus occurs in Turkey; the flight period is unknown. The alagoezicus group Apomorphies: Apical shaft part lengthened in ae- deagus of the d; fringed plates on thecal apex of ae- deagus oblique to erect. Description Head: Antennae variably coloured but antenna 3 often paler than rest; face with often strong blue me- tallic lustre; pubescence yellow to whitish; vertex ang- le in the males 25-35°, ocellar angle in both sexes ap- proximately 45°. Thorax: Dark, with longitudinal pruinose bands on dorsum more or less clear; katepisternum, anepis- ternum and anepimeron often with very strong blue lustre. The females show 5 longitudinal pruinose bands. Legs: The usual pattern of coloration is: femora dark, tibiae yellow with wide dark band and some- times with distal spur, tarsi dark; the tarsi 3, especial- ly basitarsi 3 often with dense golden recumbent pu- bescence on sides, contrasting dark pubescence on upper face; this contrast is less strong in the females. 197 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Abdomen: Slender, T II strongly tapering, T IV of- ten bulging to hold the relatively large genitalia in the males; usually slender, triangular in the females; prui- nose bands strongly arcuate, the one on T IV in the males often recurving laterally; lateral portions of ter- gites often metallic lustrous. Genitalia: The anterior surstyle lobe lengthened, extremely lengthened in some species; accessory lobes may be present on both main surstyle lobes. Cercus usually large, always well protruding. Aedeagus slen- der, invariably with extremely lengthened apical shaft part, outer face often with humps or plates; fringed plates on thecal apex suberect to erect. Identification of females in the alagoezicus group. — Apart from the fentales of satdagensis which have been so assigned on the grounds of associated capture, the females in this group cannot be assigned to any spe- cies yet. As a group these females can be distinguished from those of other groups by the presence of 5 prui- nose bands on the thorax (4 bands in other groups), relatively large size and slender, usually triangular ab- domen. Merodon alagoezicus Paramonov (figs. 67 a, 68 a-d, 99) Merodon alagoezicus Paramonov, 1925: 151. Syntypes d: ‘Inaklu, Etschmiadzin, Erivan district, Armenia, Southern slope of Mount Alagéz’ [not examined]. Merodon alagoezicus, Paramonov 1926a: 15;1926b: 318; 1927: 74; Liepa 1969: 20; Peck 1988: 166. Lampetia alagoécica; Sack 1931: 303 [orthographic error]. Material examined. — Greece: 38 (ZMUC); Turkey: 81 4 (LR, WH, BMNH). Description Male. — Head: Antennae brown to pale brown, an- tenna 3 palest, with upper margin convex, apex obtu- se, antennal ratio 1.7; pubescence pale yellow, conspi- cuously dense on frons; face strongly blue metallic lustrous, sparsely pruinose throughout; vertex angle 30°, ocellar angle 40°, tl-v ratio 0.65; compound eyes bearing inconspicuous pale pubescence. Thorax: Dark; blue lustrous on lateral dorsum and scutellum, katepisternum anepisternum and anepim- eron; pubescence rather even, moderately long, pale yellow on dorsum, longer and less even on sides; in many specimens vague dark interalar band present; longitudinal pruinose bands vestigial to well-defined. Wings clear; halteres, squamae and antisquamae yel- low. Legs: The femora dark with narrow paler apices; tibiae dark with yellow apices and, in most speci- mens, yellow bases; tarsi dark, golden pubescence on 198 sides and lower face, upper face dark; trochanter 3 with inconspicuous low ridge, femora 3 swollen, slightly curved, with normally shaped triangular pro- cessus bearing 6-9 bristles on serrate distal margin; tibiae 3 with distal outer angle much projecting, spur- like; tarsi 3 normally shaped. Abdomen: Dark; T II tapering, bearing yellow to orange lateral spots with deep lanceolate postero-lat- eral dark indention; T II-IV with arcuate interrupted moderately wide pale pruinose bands, on T IV re- curving laterally; pubescence mainly pale, with some dark pubescence on dark surface in most specimens; S IV widely, rather acutely emarginate posteriorly up to two-thirds of its length. Genitalia: Anterior surstyle lobe ventro-posteriorly recurved, enormously elongate, its apical portion showing dense, even, yellow, short pubescence; the posterior surstyle lobe coalescent, rather low, with moderately dense, yellow, erect pubescence along en- tire medio-apical margin and posteriorly. Cercus large, rounded, showing dense, long, erect, yellow pu- bescence. Aedeagus smooth, with paired recumbent chitinous plates on outer face guarding the subapical cavity; apical shaft part long, the fringed plates on thecal apex erect. Body length 12-15 mm. Female. — Unknown (see preceding observation). Diagnosis. — In view of the curious genitalia ala- goezicus can be confused with /ucasi only; from that species it is distinguished by the projection at the dis- tal outer angle of tibiae 3 and the lack of a semicircu- lar projection on the posterior surstyle lobe; alagoezi- cus can be easily distinguished from the other members of this group by the much longer anterior surstyle lobe. Period of flight and distribution (fig. 99). — The species occurs from June through August in south- eastern Europe, Turkey and Armenia. Biology. — Specimens have been collected in wet grassland on white- and yellow-flowering Umbelli- ferae; the majority of the specimens was collected in mountains at altitudes over 1500 metres. Merodon lucasi sp. n. (figs. 67 d, 69 a-c, 102) Type-material: 8 Holotype ‘Turkey, Hakkari, Suvarihalil pass 2300-2400 m, 11.viii.1983, leg. J. A. W. Lucas’ (ZMAN). Paratypes: 22d: 8d with same data as holo- _ type (LR); 48 ‘Turkey, Konya 1300 m, 5/6.viii.1981, leg. H. Coene et al.’ (LR); 16 ‘Turkey, Hakkari, Suvarihalil pass 2.viii.1982 M. Kuhbandner (LR); 26 “Turkey Pr. Hakkari Suvarihalil Pass SE Beytisebap 2300 m 2.viii.1982 and 27.vi.1985 W. Schacht (LR); 16 “Turkey, Kars, Handere 20 km W. Sarıkamıs 2100 2200 m, 1.viii.1983 leg. J. A. W. Lucas’ (JLR); 28 “Turkey, Hakkari, Suvarihalil pass 2300 m 27.vi.1985 C. J. Zwakhals (LR); 16 ‘Turkey, Hakkari S of Yiiksekova, Sat Daglari 1700 m 28.vi.1985 leg. W. Schacht (LR); 16 “Turkey, Hakkari, Varegös Sat Daglari 1700 m 29.vi.1985 W. Schacht (LR); 16 ‘Turkey, Hakkari, Habero Deresi valley 1200 m 25.vi.1985 W. Schacht (LR); Id ‘Turkey, Erzurum, Palandöken Dagi 2300 m 11.vii.1986 leg. J. A. W. Lucas’ (JLR). Description Male. — Head: Antennae dark, antenna 3 with up- per margin concex, apex obtuse to subacute, antennal ratio 1.4; vertex angle 35°, ocellar angle 45°, tl-v ratio 0.6, pubescence pale yellow, denser on frons where causing silky lustre; face bronze to blue lustrous, spar- sely white pruinose. Thorax: Dark; blue lustrous on katepisternum, an- episternum, anepimeron and (less strong) on lateral scutellum and dorsum; pubescence pale brown to yel- low, moderately dense; pruinose bands inconspicu- ous, the median bands clearest; a fifth anteromedian band often present; these bands often cross-connect- ed along dorsal suture. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: Dark with yellow markings on femoral apices and both ends of tibiae; trochanter 3 with low but well marked sharp ridge; femora 3 swollen, curved, triangular processus bearing 7-9 bristles on weakly serrate distal margin with clear apical pedestal; tibiae 3 with apical outer angle projecting slightly, tarsi 3 normally shaped. Abdomen: Dark; slender; T II strongly tapering, T IV bulging; T II with conspicuous orange lateral spots, T II-IV with arcuate interrupted moderately wide, well bordered white pruinose bands; pubes- cence concolorous but many pale yellow hairs mixed in laterally and posteriorly on T III-IV, densest on lateral spots; S IV widely acutely emarginate up to half its length. Genitalia: Anterior surstyle lobe ventro-posteriorly recurved, enormously elongate, apically with dense even short yellow pubescence; the posterior surstyle lobe coalescent, anteriorly showing semicircular ven- tral projection and bearing yellow moderately dense erect pubescence. Cercus large, quadrate, with dense, long, yellow pubescence. Aedeagus rather slender, with paired humps on basal outer face and bearing chitinous plates guarding subapical cavity, apical shaft part long, fringed plates on thecal apex narrow, long, erect. Body length 11-15 mm. Female. — Unknown (see preceding observation). Diagnosis. — M. lucasi can be distinguished from Hurkmans: Monograph of Merodon the other members of the alagoezicus group by the much longer anterior surstyle lobe, and moreover from alagoezicus by the semicircular plate on the pos- terior surstyle lobe and by having no large spur on tib- iae 3. Period of flight and distribution (fig. 102). — The species is known from Turkey where it was collected in June and August. Etymology. — It is a pleasure to dedicate this species to Jan Lucas who collected an enormous amount of specimens which were generously placed at the authors disposal. A noun in genitive case. Merodon nitidifrons sp. n. (figs. 70 a-d, 101) Type-material: & Holotype “Turkey, Hakkari, Suvarihalil pass 2300 m [37° 25’ N, 43° 03’ E], 13.vi.1984, leg. J. A. W. Lucas’ (ZMAN). Description Male. — Head: Antennae orange, antenna 3 with convex darkened upper margin, apex subacute, an- tennal ratio 1.3; vertex angle 25°, ocellar angle 35°, tl-v ratio 0.55; pubescence yellow, densest and re- cumbent on frons where causing silky lustre; face densely white pruinose over blue metallic surface; some dark pubescence present in ocellar region. Thorax: Dark, blue lustrous on katepisternum, an- episternum and anepimeron, lateral dorsum and scu- tellum; conspicuous longitudinal bands on dorsum in two pairs and a fainter anteromedian band; pubes- cence dull golden, moderately dense, most even. Wings clear; halteres, squamae and antisquamae yel- low. Legs: Dark but femoral apices and both ends of tib- iae orange; trochanters 3 smooth, femora 3 slightly swollen, triangular processus rather low, bearing 4-6 rather weak bristles on hardly serrate distal margin; tibiae 3 with distal outer angle somewhat projecting. Abdomen: Slender, T II much tapering, T IV bulg- ing; T II-III with pale orange lateral spots, T I-IV with arcuate rather wide pale pruinose bands (inter- rupted on II-III, continuous and laterally recurved on IV), pubescence concolorous; on closer examination T IV shows small orange anterolateral spots. Genitalia: The anterior surstyle lobe much elon- gate, narrow, evenly wide throughout, apically re- curved medially and bearing dense, short, even, yel- low pubescence; basally bearing accessory lobe on medial face; this lobe is visible as a ridge in lateral view and has same pubescence as apical anterior sur- style lobe part; the posterior surstyle lobe coalescent, moderately high, elliptical, bearing moderately dense 199 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 long yellow pubescence in strip along margins. Cercus wide, with long, dense, somewhat wavy, yel- low pubescence. Aedeagus very slender, bearing paired chitinous plates guarding subapical cavity, ap- ical shaft part long, fringed plates on thecal apex sub- erect. Body length 15.5 mm. Female. — Unknown. (see preceding observation) Diagnosis. — M. nitidifrons is easily distinguished from other members of the alagoezicus group by the ribbon-shaped, medially recurved anterior surstyle lobe. Period of flight and distribution (fig. 101). — This species is known only from the holotype, collected in eastern Turkey in June. Etymology. The epitheton nitidifrons derives from the latin nitere, to shine, and frons, a part of the head so indicated, and refers to the silky lustrous frons. It is to be treated as a noun in apposition. Merodon satdagensis sp. n. (figs. 67 c, 71 a-c, 101) Type-material: & Holotype ‘Turkey, Hakkari, Suvarihalil pass 2300 m, SE Beytisebap 2.viii.1982 leg. W. Schacht’ (ZMAN). Paratypes: 31d, 22; 2d with same data as holotype (LR); 54 topotypic, 4.viii.1982, leg. J. A. W. Lucas (LR) 5d topotypic, 1 1.viii.1983, leg. J. A. W. Lucas (LR); 36 “Turkey, Hakkari, Varegôs S. of Yüksekova 1700 m 29.vi.1983 leg. W. Schacht’ (LR); 12d from same locality, 5-8.viii.1983, leg. Schacht (JLR), 16, 29 from same locality, 5-8.viii.1983, leg. Lucas (JLR); 16 “Turkey, Hakkari, Sat Dag, North of Mt. Gavaruk 2900 m, 7.viii.1983 leg. Kl. Warncke’ (LR); 1d ‘Turkey, Hakkari, Habur Deresi S. of Yüksekova, 26.vi.1985, leg. Schacht’ (LR); 18 “Turkey, Hakkari, S of Yüksekova 1700 m, 28.vi.1985, leg. W. Schacht’ (LR). Description Male. — Head: Antennae dark orange to brown, an- tenna 3 paler, upper margin convex, apex obtuse, an- tennal ratio 1.3, vertex angle 35°, ocellar angle 45°; pubescence yellowish white, rather dense on face and frons; face and vertex blue metallic lustrous, modera- tely dense white pruinose; tl-v ratio 0.3. Thorax: Dark, katepisternum, anepisternum ane- pimeron, lateral dorsum and scutellum blue to green or slate grey lustrous; two pairs of narrow longitudi- nal pruinose bands present; pubescence pale brown to yellow, moderately dense, rather even. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: Dark, but tibiae yellow with wide dark distal band, femora apically yellow in most specimens; tro- chanters 3 smooth, femora 3 swollen, curved, trian- gular processus normally shaped, bearing 7-9 bristles 200 on serrate distal margin; tibiae 3 apically with spurlike outer angle. Abdomen: Dark, slender, T II strongly tapering,- with large pale yellow lateral spots; T II-IV with mod- erately wide interrupted (continuous on T IV) whit- ish strongly arcuate well bordered pruinose bands, sometimes combined with red surface coloration; T IV bulging; pubescence concolorous but many pale hairs prsent throughout; S IV deeply widely emargi- nate, strongly vaulted; T IV posteriorly pruinose. Genitalia: The anterior surstyle lobe much elon- gate, apical part recurved, dorso-ventrally aligned, with subbasal ventral accessory lobe; both apical part and this lobe short dense erect even yellow pubes- cence; the anterior and posterior surstyle lobes separ- ated by shallow sulcus; posterior surstyle lobe elon- gate, twisted, with long, yellow pubescence posteriorly and along medial distal margin. Cercus large, elongate, with denser pubescence than posteri- or surstyle lobe. Aedeagus extremely long, slender, bearing paired chitinous flanges guarding subapical cavity which due to extreme length of apical shaft part is situated halfway up the aedeagus; fringed plates on thecal apex erect, rather long and narrow. Body length 12-15 mm. Female. — Except for sexual dimorphism, differing from the male as follows. Head: Lustrous midstripe on frons occupying 1/4 of width; ocellar angle 45°, antennae with third article darkest. Thorax: Pubescence shorter, more even, a fifth anteromedian pruinose band present. Legs: the spurlet on tibiae 3 lacking. Abdomen: T II less tapering, lateral spots larger, pruinose bands less wide; T IV posteriorly pruinose. Body length 13 mm. Diagnosis. — M. satdagensis males will be easily dis- tinguished by the long anterior surstyle lobe with sub- basal accessory lobe and twisted posterior surstyle lobe. The females are very similar to other females of species in this group; they can preliminarily be distin- guished from those of other species by the strongly serrate triangular processus; see preceding observa- tion. Period of flight and distribution (fig. 101). — M. satdagensis is known from south-eastern Turkey where it occurs late in June and in August. The known range of the species is restricted to the Sat Mountains in the Turkish province of Hakkari near the Iranian and Iraqi borders. Discussion. — The females have been assigned to satdagensis given their concurrent flight in the same biotope at Varegös in the Sat mountains of eastern Turkey; moreover the much serrate triangular proces- sus in these females occurs in males of only two spe- cies of the alagoezicus group, viz. satdagensis and tani- niensis, of which the latter is stouter, much lighter coloured and in the possession of much wider ab- dominal pruinose bands; the females concerned being relatively dark and slender were assigned to satdagen- sis. Etymology. — The epitheton is an adjective, and named after the Sat Daglari, a mountain range in south-eastern Turkey. Merodon schachti sp. n. (figs. 67 b, 72 a-c, 100) Type-material: 8 Holotype ‘Turkey, Hakkari, Suvarihalil pass 2300 m, 2.viii.1982 leg. W. Schacht’ (ZMAN). Paratypes: 9 ; 24 topotypic, 11.viii.1983 leg. J. A. W. Lucas (JLR); 16 topotypic, 13.vi.1984 leg. Lucas (LR); 16 ‘Turkey, Hakkari, Tanin-Tanin pass 1700 m (W side near Habul Deresi) 12.vi.1984 leg. Lucas’ (LR); 3d from same locality, 25.vi.1985 leg. W. Schacht (LR), 1d from same locality, 25.vi.1985 leg. C. J. Zwakhals (LR); 1d ‘Turkey, Hakkari, Habur Deresi valley 26.vi.1985 1700 m leg. Schacht’ (JLR). Description Male. — Head: Antennae orange-brown, antenna 3 slightly paler, upper margin convex, apex obtuse, not rounded, antennal ratio 1.3; pubescence whitish, densest on frons; face blue metallic lustrous, diffusely white pruinose (densest medially); vertex angle 25°, ocellar angle 40°, tl-v ratio 0.4. Thorax: Dark, blue lustrous on katepisternum, an- episternum and anepimeron, lateral dorsum and scu- tellum; pruinose bands on dorsum in anteriorly co- alescent pairs, narrow; pubescence sparse, pale brown to yellow. Wings clear; halteres, squamae and anti- squamae pale yellow. Legs: Dark with yellow markings on apices of fe- mora and both ends of tibiae; trochanter 3 smooth, femora 3 swollen, somewhat curved, triangular pro- cessus bearing 7-9 bristles on slightly serrate distal margin (less serrate than in satdagensis but more than in alagoezicus); tibiae 3 with distal outer angle extend- ed, tarsi 3 normally shaped. Abdomen: Dark: slender; T II strongly tapering, T IV bulging, T II-III with large and small orange later- al spots respectively; T II with weak, T III-IV with strong whitish interrupted arcuate pruinose bands, the one on T IV recurved laterally; posterior margins of T III-IV pruinose; pubescence pale, but strictly dark on posterior, dark surface of T III; S IV acutely emarginate posteriorly up to 3/4 its length or even more. Genitalia: Anterior surstyle lobe very elongate, re- curved apically, there dorso-ventrally aligned, apically Hurkmans: Monograph of Merodon bearing dense, yellow, short, even pubescence; anteri- or surstyle lobe coalescent with gradually rising poste- rior surstyle lobe elongate, low, and bearing moder- ately long, yellow pubescence along its distal margin and posteriorly; posterior surstyle lobe laterally guard- ed by a free chitinous plate basally attached to the sur- style base; this plate has a distinct antero-ventral angle and a lateral approximately dorso-ventral ridge and posteriorly shows dense yellow pubescence. Cercus large, with dense, long, erect, yellow pubescence. Aedeagus quite robust, apical shaft part lengthened much, fringed plates on thecal apex erect, long and narrow, some scattered short pubescence present on inner face. Body length 13 mm. Female — Unknown (see preceding observation). Diagnosis. — M. schachti can hardly be confused with any other species given the very conspicuous lat- eral guarding plate alongside the posterior surstyle lobe in the males; moreover, the robust aedeagus sep- arates it from other species in the alagoezicus group. Period of flight and distribution (fig. 100). — The species is known exclusively from the south-eastern- most region of Turkey where it has been collected in June and August. Etymology. — The species is named after the collec- tor of the type, the German entomologist Dr. Wolfgang Schacht, of Munich. A noun in genitive case. Merodon taniniensis sp. n. (figs. 73 a-c, 100) Type-material: d Holotype “Turkey, Hakkari, Tanin- Tanin pass W. side 1700 m, 12.vi.1984 leg. J. A. W. Lucas’ (ZMAN). Paratypes: 68; 2d with same data as holotype (LR); 26 ‘Turkey, Hakkari, Tanin-Tanin pass E Uludere 2300 m 25.vi.1985 leg. W. Schacht’ (LR); 18 “Turkey, Hakkari, Suvarihalil pass near Habul Deresı 1250 m 13.vi.1984 leg. Lucas’ (LR); 1d “Turkey, Malatya, Sarıhacı 45 km W. Malatya 21.vi.1984 leg. Lucas’ (JLR). Description Male. — Head: Antennae brown, antenna 3 with upper margin convex, apex obtuse, antennal ratio 1.6, pubescence whitish pale yellow throughout; vertex angle 25°, ocellar angle 45°, tl-v ratio 0.4; face blue metallic lustrous, densely white pruinose. Thorax: Dark, lustrous on katepisternum, anepis- ternum, anepimeron and lateral dorsum and scutel- lum; dorsum showing 5 pruinose bands; pubescence yellow, very even, moderately dense. Wings clear; hal- teres, squamae and antisquamae pale yellow. 201 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Legs: Dark but tibiae yellow with wide dark distal band; trochanter 3 with low sharp ridge, femora 3 swollen, curved, triangular processus bearing 6-8 bris- tles on somewhat serrate distal margin. Abdomen: Slender; T II strongly tapering, T IV bulging, predominantly orange, with connected ante- rior and posterior dark lunules on T II and weaker brown medial coloration on T III; posterior lunules on T III-IV pale brown to orange; T II-IV bearing rather wide strongly arcuate, interrupted to continu- ous whitish pruinose bands (recurved on T IV); pu- bescence pale yellow, moderately dense (also on dark surface); all S orange, S IV widely, obtusely emargi- nate, slightly vaulted. Genitalia: The anterior surstyle lobe large, rather quadrate, slightly bifid anteriorly, bearing dense short erect yellow even pubescence; connected by a slender stalk to the base of the posterior surstyle lobe, a deep narrow sulcus separating from the apical part; the posterior surstyle lobe twisted, bearing two apical ridges, both bearing long, erect, yellow, sparse pubes- cence, posterior part of posterior surstyle lobe with dense pubescence. Cercus large, mainly hidden between the elongate posterior parts of the posterior surstyle lobes, with dense, long, yellow pubescence. Aedeagus slender, apical shaft part lengthened, ring paired chitinous plates guarding subapical cavity (these plates do not resemble those found in the ele- gans group); fringed plates on apex aedeagi erect, long and rather narrow. Body length 14-16 mm. Female. — Unknown (see preceding observation). Diagnosis. — M. taniniensis can be distinguished from other species of the alagoezicus group by the rela- tively very large, rather quadrate anterior surstyle lobe and the double ridges on the posterior surstyle lobe. Period of flight and distribution (fig. 100). — Known from the central and southern parts of eastern Turkey where it has been collected in June. Etymology. — The epitheton taniniensis, an adjec- tive, refers to the Tanin-Tanin pass, the type locality of the species in south-eastern Turkey. Merodon toscanus sp. n. (figs. 74 a-c, 99) Type-material: d Holotype ‘Italia Toscana, Careggi (Firenze) 19-21.v.1986 W. Hurkmans’ (ZMAN). Description Male. — Head: Antennae brown, antennae 3 with upper margin convex, apex obtuse, antennal ratio 1.6, 202 vertex angle 20°, ocellar angle 35°; pubescence pale yellow, face metallic blue lustrous, densely whitish pruinose; a dark hair tuft present in ocellar region. Thorax: Dark, metallic lustrous on katepisternum, anepisternum, anepimeron, lateral dorsum and scutel- lum; pubescence dull yellow, even, rather sparse; two pairs of rather faint pruinose bands present. Wings clear, halteres, squamae and antisquamae pale yellow. Legs: All femora dark brown, tibiae orange with distal wide dark bands, tarsi deep orange to brownish; trochanter 3 smooth, femora 3 rather strongly swol- len, curved, triangular processus bearing 7 bristles on weakly serrate distal margin. Abdomen: Rather slender; T II tapering; T IV bulging; colour mainly deep orange, T II bearing connected anterior and posterior dark lunules, T III with deep orange posterior and paler anterior part, T IV deep orange throughout; T II-IV with arcuate moderately wide pruinose bands, just interrupted on T I-III, continuous and laterally recurved on T IV; all S orange, S IV with drop-shaped emargination, strongly vaulted. Genitalia: Anterior surstyle lobe elongate, round- ed, bifid anteriorly, bearing dense short erect yellow even pubescence throughout (sparser posteriorly); connected to base of posterior surstyle lobe by a short stalk; the posterior surstyle lobe low, elongate, bear- ing an accessory lobe with dense long erect pubes- cence medially; its anterior part bald, lustrous, ap- proximately ovoid, its posterior part with sparse yellow pubescence, but denser on margins and poste- riorly. Cercus large, with dense, erect, yellow, long pubescence. Aedeagus very slender, with large paired basal humps on outer face; apical shaft part much lengthened; fringed plates on thecal apex suberect, rather narrow. Body length 12 mm. Female. — Unknown (see preceding observation). Diagnosis. — M. toscanus has an accessory lobe on the posterior surstyle lobe and a bald anterior part of the posterior surstyle lobe not found in the other spe- cies of the alagoezicus group. The bifid anterior sur- style lobe is much stronger than in taniniensts. Period of flight and distribution (fig. 99). — The unique holotype was collected in May in Toscana, Italy. Etymology. — The species has been named after Toscana, the region where the holotype was collected. An adjective. REMAINING SPECIES Two of the three species discussed under this head- ing are known only after female specimens. Since these species share character states considered apo- morphic for some species treated in this paper, they are discussed here. Of the third species, M. viaticus no material was examined, but according to its descrip- tion it might occupy a position close to some of the species treated in this paper. Merodon affinis Gil Collado Merodon affinis Gil Collado, 1930: 243, 255. Holotype 2: ‘Escorial, Arias Encobet, vii.1905’ [probably El Escorial, Madrid] (IEE) [examined]. Merodon affinis, Peck 1988: 166. Description Female. — In many respects the female affinis is si- milar to avidus, with following differences. Head: According to the original description, antenna 3 slightly curved, antennal ratio 2.0, apex acute (anten- na 3 missing on the holotype on both sides). Legs: Femora 3 with distal margin of triangular processus much stronger serrate than in avidus. Abdomen: Outline triangular, pruinose bands narrowly inter- rupted, less arcuate than in avidus. Body length 12 mm. Male. — Unknown. Discussion. — Because of several differences in the females, affinis cannot simply be regarded a synonym of avidus, especially not in absence of the males. The status of this species remains uncertain. In view of the mentioned similarities with the females of avidus, af- finis could conceivably belong to the avidus group but to e.g. the alagoezicus or nigritarsis group as well. Period of flight and distribution. — Known only from the holotype, collected in central Spain in July. Merodon aureotibia sp. n. (fig. 106) Type-material: 2 Holotype ‘Turkey, Adiyaman, Nemrut Daglari, 1.vi.1983, leg. M. Kuhbandner (ZMAN). Paratypes: 32 ‘Tk [Turkey] 5-6-1988 10 km S. Ankara leg. Warncke’ (JLR). Description Female. — Head: Antennae bright orange, antenna 3 rather broad in outline, apex subacute, upper mar- gin straight, arista long, suddenly tapering, antennal ratio 2.8; pubescence golden; lustrous midstripe oc- cupying 1/3 of width, ocellar region showing dark Hurkmans: Monograph of Merodon pubescence; ocellar angle 70°, lateral ocelli larger than central ocellus; pubescence on compound eyes rather thin, whitish. Thorax: Rather narrow, black with weak bluish lustre on katepisternum, anepisternum, anepimeron, lateral scutellum and dorsum, with four longitudinal yellow pruinose bands on dorsum, coalescent anteri- orly; pubescence rather short, erect, uneven, golden yellow, but darker, brownish in interalar band. Wings clear; halteres, squamae and antisquamae pale yellow. Legs: All orange except coxae and trochanteres brownish; femora 3 swollen, rather straight, bearing 6 bristles on normal triangular processus. Abdomen: Rather slender, dark with micaceous lustre; T II bearing large orange lateral spots, T II-IV with slightly arcuate, moderately wide, interrupted, golden pruinose bands combined with golden pubes- cence; pubescence concolorous otherwise, but yellow laterally and on posterior fringe of T III-IV; T V far projecting, rounded posteriorly. Body length 11 mm. Male. — Unknown. Diagnosis. — M. aureotibia can be distinguished from all other females of Merodon by its extensively orange leg coloration. Period of flight and distribution (fig. 106). — This species occurs in June in central and south-eastern Turkey. Discussion. — In view of the unique leg coloration, the narrow thorax and striking golden pubescence on especially head and abdomen aureotibia has been de- scribed as a new species. It is reminiscent of females of aberrans but differs from it by the leg colour, shape and colour of the antennae, narrower thorax and con- spicuous orange lateral spots on T II. Moreover the femora 3 in aureotibia are more swollen. In the coloration of the.legs the females of aureoti- bia resemble vandergooti, known only after males. Reasons for not considering aureotibia as the female sex of vandergooti are: (1) aureotibia has obtuse anten- nae, orange and slightly swollen hind femora, arcuate pruinose bands not obliquely placed and a serrate dis- tal triangular processus margin with 6-7 bristles, (2) vandergooti has acute antennae, dark and enormously swollen hind femora, oblique bands and a hardly ser- rate distal triangular processus bearing 10-13 bristles. These differences are far too great and of a wrong type to be considered as sexual dimorphism. In view of the slender habitus, relatively long an- tenna 3 with suddenly tapering arista, aureotibia might possibly be related to the longicornis group. However, none of the species in that group has 203 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 orange legs and relatively wide pruinose bands, whereas aureotibia lacks the typical crimson abdomi- nal colour of the longicornis group. Merodon viaticus Fabricius Merodon viaticus Fabricius, 1805: 197. Lectotype, sex un- known (here designated): ‘viaticus’, [the upper half of a thorax with basal wing fragments attached] in Sehestedt & Tonder Lund coll. (ZMUC) [examined]. Merodon viaticus Wiedemann in Meigen 1822: 364; Macquart 1828: 296; Rondani 1845: 258; Walker 1849: 598; Schiner 1857: 414. Lampetia viatica, Sack 1931: 327. Material examined. — Paralectotype, sex unknown ‘viaticus [part of a thoracic dorsum] in Kieler coll. (ZMUC). Description The size of the thoracic dorsum indicates an overall size of about 12 mm. The background coloration is dark, pubescence largely rubbed off, but where remai- ning, erect and whitish; pruinose bands rather clear. Wing fragments clear, brown veined. Discussion. — The original description by Fabricius (1805) does not mention the sex of the fly described. As however a dark median line on the frons is men- tioned it must have been a female. The habitus should be similar to that of annulatus which is a syno- nym of natans. Stated differences with viaticus are the all dark abdomen in natans (with a red base in wiati- cus), the all dark legs and smaller size, averaging 9 mm for natans. Wiedemann (in Meigen 1822) gives an extensive description of viaticus, in which the abdomen is said to be red on T I II and anterior T III; size is given by Wiedemann as 5 to 6 lines, i. e. 10.5 to 12.2 mm. Wiedemann’s description is considered to include both sexes as he comments on differences in the colour of antennae and tarsi between sexes. Some specimens from Sicily, leg. Schigdte, labelled ‘viaticus’, without any further data (ZMUC) have been re-identified as elegans sp. n.; differences with the de- scription of viaticus are the size (14 mm), much swol- len femora 3 (not mentioned in either of the descrip- tions); legs yellow, not reddish or rusty brown-yellow as in the descriptions; abdomen dark brown with ex- tensive yellow coloration, not red, orange or purplish as in descriptions. M. viaticus is assigned a place in the section of Merodon containing the species with arcuate pruinose bands since these are mentioned in the original de- scription. As there are no females of species in this section of Merodon with a cupreous lustre on the abdomen and at the same time partially dark, partially orange legs, 204 viaticus cannot be synonymized with any of the spe- cies. It was listed by Sack (1931: 327) as a synonym of spinipes (=avidus); no reasons for this synonymy were given. ANNOTATED LIST OF SPECIES DISCUSSED IN THIS PAPER Abbreviations used to describe status: H = Holotype examined; ? = Type material not located, not examined; S = syntype(s) examined; P = paraty- pe(s) examined; L = lectotype examined (here de- signated); PL = paralectotype(s) examined. Deposition Status Ann distinctus assemblage biarcuatus Curran, 1939 AMNH H = clunipes Sack, 1913 ZMHB = distinctus Palma, 1863 ? ? 1 dimorphus (Szilady, 1940) syn. n. TMA [lost] 2 femoratus Sack, 1913 ? S? 3 maride sp. n. ZMAN H, P = ottomanus sp. n. ZMAN H, P = testaceoides sp. n. RMNH, ZMAN H,P = alexeji group alexeji Paramonov, 1925 PC [lost] 4 altinosus sp. n. ZMAN H hirsutus Sack, 1913 NHMW L, PL = hypochrysos sp. n. ZMAN H, P = kawamurae Matsumura, 1916 ? 5 micromegas (Hervé- Bazin, 1929) syn. n. MNHN, BMNH L, PL 5 marginicornis sp. N. ZMAN H rufitarsis Sack, 1913 stat. nov. ZMHB H = sophron sp. n. BMNH H = tener Sack, 1913 DEIC IL, JBL, 6 trizonus (Szilady, 1940) TMA [lost] 2 tarsatus group ankylogaster sp. n. RMNH lal, IP auronitens sp. n. ZMAN H - caudatus Sack, 1913 FIS [lost] 3 oidipous sp. n. ZMAN H, P = persicus sp. n. RMNH H = tangerensis sp. N. MNHN H = tarsatus Sack, 1913 ZMHB H = smirnovi Paramonov, 1926 syn. n. PC [lost] 4 turkestanicus Paramonov, 1926 PC [lost] 4 xanthipous sp. n. ZMAN H, P = clavipes group aberrans aberrans Egger, 1860 NHMW IE IRL, = knerii Mik, 1867 NHMW? S? = obscuripennis Palma, 1863 aberrans ssp. flavitibius Paramonoy, 1925 aberrans ssp. isperensis ssp. n. brevis Paramonov, 1925 clavipes (Fabricius, 1781) canipilus Rondani, 1865 clauda (Villers, 1789) curvipes (Gmelin, 1790) gravipes (Rossi, 1790) sacki Paramonov, 1937 syn. n. senilis Meigen, 1822 clavipes var. albus Paramonov, 1927 clavipes var. ater Paramonov, 1927 clavipes var. niger Paramonov, 1927 cupreus sp. n. dzhalitae Paramonov, 1926 hamifer Sack, 1913 karadaghensis Zimina, 1989 lusitanicus sp. n. quadrinotatus (Sack, 1931) splendens sp.n. velox Loew, 1862 velox var. anathema Paramonov, 1925 velox var. armeniacus Paramonov, 1925 warnckei sp. n. pruni group pruni (Rossi, 1790) fulvus Macquart, 1834 fuscinervis Von Réder, 1887 pallida Macquart, 1842 sicanus Rondani, 1845 pruni var. obscurus Gil Collado, 1929 longicornis group erivanicus Paramonov, 1925 kaloceros sp. n. longicornis Sack, 1913 vandergooti group vandergooti sp. n. nigritarsis group femoratoides (Paramonov, 1925) comb. n. nigritarsis Rondani, 1845 PC ZMAN, JLR PC NHMW, DEIC Moscow ZMAN, JLR PC PC ZMAN, JLR PC ZMAN, JLR NHMW ZMAN, JLR ZMHB LSF Hurkmans: Monograph of Merodon avidus group avidus (Rossi, 1790) ZMHB [lost] 9 spinipes (Fabricius, 1794) ZMUC IL il serrulatus Wiedemann in Meigen, 1822 NHMW It = italicus Rondani, 1845 LSE L, PL = rufitibius Rondani, 1845 LSF L, PL = graecus Walker, 1852 BMNH IL = aurifer Loew, 1862 ZMHB IL - quadrilineatus Lioy, 1864 nom. nud. = 12 crassifemoris group crassifemoris Paramonov, 1925 comb. n. PC [lost] 4, 13 elegans group bequaerti sp. n. MNHN H, P = elegans sp. n. ZMAN e. a. IND 14 manicatus (Sack, 1938) | HLMD EIND, = testaceus Sack, 1913 NHMW L, PL = alagoezicus group alagoezicus Paramonov, 1925 PC [lost] 4 lucasi sp. n. ZMAN, JLR EUD - nitidifrons sp. n. ZMAN H satdagensis sp. n. ZMAN, JLR lial, Je - schachti sp. n. ZMAN, JLR H,P = taniniensis sp. n. ZMAN, JLR lal, IP - toscanus sp. n. ZMAN H = Remaining species affinis Gil Collado, 1930 IEE H = aureotibia sp. n. ZMAN H = viaticus (Fabricius, 1805) ZMUC L, PL 15 Annotations 1. The types from the Palma collection have not been located. The descriptions and drawings provi- ded by Palma (1863) of obscuripennis and distinctus are extensive enough to be used for diagnostic purpo- ses. Sack (1931) has been followed in considering 0b- scuripennis synonymous with aberrans. 2. The type-material, formerly at TMA, is lost (Dr. A. Dély-Draskovits, in litt.). 3. The holotypes of caudatus and quadrinotatus, and the syntypes of M. femoratus, all formerly in Sack’s private collection, were bequeathed to the Forschungsinstitut Senckenberg, Frankfurt/ Main. According to Dr. W. Tobias, curator of Syrphidae in the said museum, no Sack types of any Merodon are currently there. When Sack left his collection to SMF the material was presumably still stored in his own house which was plundered shortly after World War II. All Sack types of Merodon left to the Senckenberg Museum are therefore to be considered lost. 205 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 4. Many type-specimens from the S. J. Paramonov collection have not been located. According to Liepa (1969) most of the types originally in that collection are lost. Some material which probably originated from that collection has been found in ZMHB, includ- ing type-material of Merodon femoratoides and M. crassifemoris. Dr O. G. Ovchinnikova of the USSR Academy of Sciences (St. Petersburg) informed me that some types from the Paramonov collection might still be preserved at the Kiev Zoological Museum. This is also the view of Dr. Y. Nekrutenko, Kiev (pers. comm.). Regrettably this cannot be confirmed. The type of M. smirnovi should be stored in the Zoological Institute, Moscow University but this also could not be confirmed. Reliably identified material, in some cases det. S. J. Paramonov, was present of the following species: ala- goezicus (ZMAN); alexeji (KBIN); brevis (KBIN); crassif- emoris (ZMHB); erivanicus (ZMAN); femoratoides (ZMHB) and turkestanicus (ZMAN). No material which can be reliably assigned to smirnovi is known; in view of its description this species is here considered syn- onymous with zarsatus. 5. The deposition of the type(s) is not mentioned in Matsumura’s original publication. After Horn & Kahle (1936) the Matsumura collection is at the University of Sapporo. Following enquiry to the de- partment of Entomology, Hokkaido University, Sapporo, the presence of the types of M. kawamurae there cannot be confirmed. The type(s) (sex and number unknown) originate from Kumamoto, Kyushu. However, this is the name of both a city and an extensive district. The description of the species presented here is based on 29, 2d from Harima [in- land sea of Japan at 34° 30° N, 134° 30° E], in BMNH and fully complying with the original description. 6. Only female type-specimens have been exam- ined, no male types have been found. The male from Saratow, formerly preserved at TMA, is lost (Dr. A. Dély-Draskovits, in litt.). The male and female syn- types from the Becker collection are not at ZMHB and have not been located elsewhere. A male type from Uralsk, stated in the original publication to be in NHMW, was not found there during a visit in 1987. The 2 female syntypes from which the lectotype has been designated are stored in DEIC, Eberswalde. 7. This species has been described without mention of material, although it is stated by Villers (1789) that Musca clauda occurs in Europe, in Bressia [? = Brescia]. 8. Musca curvipes Gmelin is directly referred to Fabricius’ original description of Syrphus clavipes by 206 Gmelin (1790). Therefore type-material of this taxon does not exist. 9. No type-material of Syrphus gravipes, Syrphus pruni or Syrphus avidus has been located. After Horn & Kahle (1936) the Rossi collection was transferred to ZMHB; after Dr. H. Schumann (in litt.) no Rossi types of Merodon are currently present there. 10. No material was examined of fulvus Macquart or fuscinervis Von Réder. Both original descriptions suggest a great similarity to pruni. Both species were listed by Peck (1988) as synonyms of pruni. This syn- onymy has been followed. 11. The type-material of spinipes consists of a sin- gle, quite completely decayed probable syntype in ZMUC. This taxon is here synonymized with avidus on the ground of complete compliance of the descrip- tions and figures throughout the literature, starting from the first publications. It must of course be borne in mind that Rossi or Fabricius may have been de- scribing any species of the large group externally re- sembling avidus. 12. No type-material of this taxon was probably ever in existence, as Lioy, after Horn & Kahle (1936) did only theoretic systematic work. 13. The holotype has not been located (see annota- tion 1). In ZMHB a specimen dating from 1929, with a label attached marking it as a type, was found. As this specimen was collected after the publication of var. crassifemoris it is not available as a type specimen; however the labels are in Paramonov’ s handwriting. The specimen fully complies with the description of the var. crassifemoris. It is therefore available as a check on identifications. 14. A large number of paratypes spread over many institutions was designated to facilitate genital study. This was done since elegans externally resembles avi- dus and could well be confused with it. 15. The two syntypes from ZMUC, one from the Kieler coll. and one from the Sehestedt and Tonder Lund coll., are so seriously damaged that no conclu- sion can be drawn as to the status of this species. No other material identified as viaticus and complying with the description of that species has been found. Therefore the descriptions given are the only base for identification. ANNOTATED LIST OF SPECIES EXCLUDED FROM MERODON The status of the species described by Walker (1849, 1852, 1857a, 1860) is not clear in all cases. As not all material was available, the specimens studied have been referred to as type-specimens. The present author does not intend to designate lectotypes of spe- cies excluded from Merodon. Merodon angustiventris Macquart, 1855: 90. Lectotype d (designated by Thompson 1988) (BMNH) [examined] not considered to belong to Merodon in view of morphology of femur 3 and wing venation. According to Thompson (1988) the lecto- type belongs to Quichuana angustiventris (Macquart), a species occurring in the Neotropical region. Merodon balanus Walker, 1849: 600. A d type- specimen (BMNH) [examined] belongs to the genus Mallota in view of femur 3 and wing venation. Specimen originates from New York [Nearctic re- gion, U. S. A.] where a natural occurrence of Merodon is unknown. Merodon bardus Say (in Walker 1849: 598). The d type-specimen [examined] does not belong to Merodon. After Osten-Sacken (1878: 135) this speci- men belongs in Mallota. Merodon bautias Walker, 1849: 600. Two syn- types, d and $ (BMNH) [examined]. Both syntypes originate from Georgia, United States. The facial knob and wing venation are not compatible with Merodon. After Osten-Sacken (1878: 135) this spe- cies belongs in Mallota. Merodon bicolor Walker, 1852: 243 [type-material not examined]. In the original description ‘feelers’ [interpreted as antennae] are stated to possess six joints, therefore the species is considered not to be- long to the Syrphidae. The type-material originated from Brazil [Neotropical region] where Merodon spe- cies do not naturally occur. Merodon bipartitus Walker, 1849: 599. The @ type-specimen (BMNH) originates from Georgia, United States [examined]. In view of the wing vena- tion, this species probably belongs to Mallota. See Osten-Sacken (1878: 135). Merodon chiragra Fabricius, 1805: 198 [type-mate- rial not examined]. Not included in a shipping of all other available Merodon types from ZMUC; this species belongs in Sphegina. Merodon coerulea Becker, 1912: 602 [type-material not examined]. On the basis of the original descrip- tion this species belongs in Eumerus. Merodon contrarius Walker, 1849: 599. Both d and © type-specimens (BMNH) [examined]. This ma- terial originated from New Holland and belongs to Orthoprosopa grisea (Walker). Merodon edentulus Macquart, 1855: 90. Holotype, Hurkmans: Monograph of Merodon ? from Cape of Good Hope [not examined]; a 2 non type (BMNH) dating from XIX century identified as edentulus [examined] belongs to Mallota. The orig- inal description mentions a facial knob, unknown in Merodon. Merodon interveniens Walker, 1860: 120. 3 Type- specimen (BMNH) [examined]. This specimen origi- nated from Macassar [Sulawesi] and does not belong in Merodon in view of wing venation, shape of leg 3 etc. Merodon melota Seguy, 1941: 13. 2 Holotype from Agadir, Morocco (MNHN) [examined]. This spe- cies belongs to Eumerus. Merodon morosus Walker, 1849: 599. A 2 type- specimen (BMNH) [examined]. This does not belong to Merodon but according to Osten-Sacken (1878: 135) 1s Polydontomyia curvipes O.-S. Merodon ornatus Brunetti, 1915: 232. Holotype in Indian Museum, Calcutta [not examined] could not be shipped in view of its conservation state. After original description the wings have a brown stigma, there is no triangular processus on the femur 3 and the face is projecting; therefore this species is consid- ered not to belong to Merodon. Merodon scutellaris Shiraki, 1968: 200. [Type-ma- terial not examined]. From the description and pho- tographed holotype it can be concluded that this spe- cies belongs to the genus Azpeytia. Merodon tenebricus Walker, 1849: 601. Lectotype 9 (designated by Thompson 1988) (BMNH) [exam- ined]. In view of the shape of femur 3 and wing vena- tion this species does not belong to Merodon. The identification as Palpoda furcata was given by Thompson (1988: 220). Merodon torpidus Walker, 1857a: 152. Lectotype d (designated by Thompson 1988) (BMNH) [exam- ined]. This does not belong to Merodon given the shape of femur 3 and wing venation. The identifica- tion as Orthoprosopa grisea Walker was given by Thompson (1988: 220). Merodon tuberculatus Brunetti, 1923: 214. The types, in the Indian Museum, Calcutta, [not exam- ined] were too fragile to be shipped; the description of this species was based on that of Helophilus tuber- culatus. The description and the figures (Brunetti, 1908) make clear that this species does not belong to Merodon. Merodon umbrifer Walker, 1849: 601. The d type- specimen from Sierra Leone does not belong to Merodon in view of leg morphology and wing vena- tion. Merodon varicolor Walker, 1857b: 122. The & type-specimen from Borneo, in BMNH, belongs to Mallota in view of the shape of femur 3 and the wing venation pattern. 207 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Proposal of Azpeytia shirakii nomen n. for Merodon scutellaris Shiraki, 1968 Since Merodon scutellaris Shiraki is incorrectly as- signed to Merodon (see above), it is here proposed to transfer this taxon to Azpeytia, where it clearly be- longs. As there already exists an Azpeytia scutellaris, published by Walker (1857b: 113), the epitheton of the species described by Shiraki, when transferred to Azpeytia, must be changed; the nomen novum Azpeytia shirakii is hereby proposed for this taxon. ACKNOWLEDGEMENTS I would like to thank Mr Volkert van der Goot for stimulating my work on the genus Merodon, his con- tinued interest in my studies, and the translation of the Zimina paper. This work would have been hardly possible without the generous co-operation of Mr Jan A. W. Lucas who kindly placed his extensive collec- tion of Merodon specimens at my disposal. My warm thanks go to Dr. Pjotr Oosterbroek who posed questions to increase my curiosity, was always willing to discuss ideas, stimulated the general progress of my studies, and critically read the various drafts of the manuscript. The facilities and advice provided by Dr. H. W. J. van Dijk were of great value in producing the final manuscript. I wish to thank Prof. Dr. J. Stock, Dr. P. Oosterbroek, Dr. A. C. Pont, London, Dra. M. A. Marcos Garcia, Salamanca, Mr C. Claussen, Flensburg, Mr. J. A. W. Lucas, Rotterdam, Dr. H. W. J. van Dijk, Zwolle, Mr. G. den Hollander, IJsselstein, Mr. P. L. H. Hanegraaf and Mrs. R. L. G. J. van der Voort, Houten, and Mr. V. S. van der Goot, Amsterdam for their valuable com- ments at the various stages of the work on Merodon, or discussion of my research in general. I would like to thank the persons already mentioned, Mrss. Marcos Garcia and Petanidou, and Messrs. Barke- meyer, Bausenwein, Ellis, Hauser, Lyneborg, Toth, Treiber, Van Ooijen, Vujic and Wyatt for the inte- resting material provided for examination. Financial support for museum and field work was gratefully received from the Uyttenboogaart-Eliasen Stichting, Amsterdam. My sincere gratitude is due to the following per- sons for permission to study preserved specimens from the collections in their care, and assistance in tracing the type-specimens of various species: Dr. Luca Bartolozzi, Firenze; Dr. Vlad. Bradescu, Bucuresti; Mr. Claus Claussen, Flensburg; Dr. Ruth Contreras-Lichtenberg, NHMW, Wien; Dr. M. Datta, Calcutta; Dr. A. Dély-Draskovits, TMA, Budapest; Dr. H. Feustel, HLMD, Darmstadt; Dr. R. Gaedike, DEIC, Eberswalde -Finow; Dr. D. Grimaldi, AMNH, New York; Dr. Patrick Grootaert, KBIN, Brussel; Dr. P. J. van Helsdingen, RMNH, Leiden; Mr. Gijs Den 208 Hollander, IJsselstein; Prior P. Bruno Hubl, Bsa, Admont; Dr. I. Izquierdo, IEE, Madrid; Dr. F. Kaplan, Tel Aviv; Dr. C. Leonardi, MCSN, Milano; Dr. Leif Lyneborg, ZMUC, Kobenhavn; Dr. Maria- Angeles Marcos Garcia, SAL, Salamanca; Dr. Carolina Martin, IEE, Madrid; Mrs. Sarah Mascherini, LSF, Firenze; Dr. L. Matile, MNHN, Paris; Dr. E. Mingo Pérez, IEE, Madrid; Mr. Peter van Ooyen, Utrecht; Dr. O. C. Ovchinnikova, St.Petersburg; Ir. Dirk Prins, Rhenen; Dr. Wolfgang Schacht, ZSBM, München; Dr. H. Schumann, ZMHB, Berlin; Dr. P. Tschorschnig, MENS, Stuttgart; Mr. Nigel P. Wyatt, BMNH, London. Also I wish to thank Dr. W. Tobias, Frankfurt/ Main and Dr. ©. G. Ovchinnikova, St. Petersburg for their help in tracing the Sack and Paramonov type-specimens, and Dr. Dely- Draskovits, Budapest for information on the TMA specimens. Last but not least I wish to express my gratitude to- wards my wife Marijke whose unstinting interest in my work greatly contributed to the result. REFERENCES Aubert, J., J-J. Aubert & P. Goeldlin, 1976. 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Über einige Merodon Arten. — Bulletin de l Académie des Sciences de |’ Ukraine IV (4) vol. 2: 317-323 [Also in Travaux du Musée Zoologique de Kiev 2: 73-79]. Paramonoy, S. J., 1927. Uber einige Arten und Varietäten von Dipteren (Fam. Stratiomyidae et Syrphidae). — Travaux du Musée Zoologique de Kiev 2: 87-93. Paramonoy, S. J., 1929. Dipterologische Fragmente 19. Uber die Gattung Merodon. — Travaux du Musée Zoologique de Kiev 7: 181-195. Paramonoy, S. J., 1935. Dipterologische Fragmente 31, 32. Wass ist ein echtes Weibchen von Lampetia armipes Rond.?; Die Unterschiedsmerkmale zwischen L. montico- la Villeneuve und L. alexeji Paramonov; Lampetia monti- cola Villeneuve 6. — Travaux du Musée Zoologique de Kiev 15: 163-166. Paramonoy, S. J., 1937a. Beiträge zur Monographie der Gattung Lampetia (Merodon olim.), Syrphidae, Diptera, 1. Teil. — Travaux du Musée Zoologique de Kiev 17: 3- 13. Paramonoy, S. J., 1937b. Dipterologische Fragmente 36. Uber einige Lampetia (Merodon) Typen. — Travaux du Musée Zoologique de Kiev 17: 69-77. Peck, L. V., 1988. Syrphidae. — Jn: A. Sods (ed.), Catalogue of Palaearctic Diptera vol. 8 Syrphidae, Conopidae pp. 11-230. Amsterdam, etc. Portschinsky, I., 1877. Materiali dlye Istorie Fauny Rossiye e Kavkaza. — Trudy Rosskago Entomologicheskago Obshtshestva 10: 102-198. Réder, H. von, 1887. Insekten der Insel Kreta. — Berliner Entomologische Zeitschrift 31: 73-74. Rondani, C., 1843. Dipteres nouveaux d’ Italie. — Revue Zoologique, Fevrier 1843: 43-44. Rondani, C., 1845. Sulle specie Italiane del Genere Merodon; Memoria decimaquarta, per servire alla Ditterologia Italiana. — Nuovi Annali di Società delle Scienze Naturale di Bologna, (serie II) 4: 254-267. Rondani, C., 1857. Dipterologiae Italicae Prodromus II. Species Italicae ordinis dipterorum in genera character- ibus definitae, ordinatim collectae, methodo analitica dis- tinctae, et novis vel minus cognitis descriptis, pars I: Oestridae, Syrphidae, Conopidae. — Parma. 264 pp. Rondani, C., 1865. Diptera Italica non vel minus descripta vel annotata observationibus nonnulis additis, fsaciculo 1: Oestridae, Syrphidae, Conopidae. — Atti della Societa Italiana delle Scienze Naturale di Milano 8: 127-146. Rondani, C., 1868a. Diptera Italica non vel minus descrip- ta vel annotata observationibus nonnulis additis, specie- bus Italicis adde sequentes: Familia II Syrphidae, genere Merodon Latr. — Atti della Societa Italiana delle Scienze Naturale di Milano 11: 22-23. Rondani, C., 1868b. Specierum Italicarum Ordinis Dipterorum Catalogus Geographicis auctus, fasciculo 1. oestridae, Syrphidae, Conopidae. — Atti della Societa Ita- liana delle Scienze Naturale di Milano 11: 559-575. Rondani, C., 1873. Species aliquae in Oriente lecta a Marchese J. Doria, anno 1862 63 Fragmentum II. — Annali di Museo Civico di Genova 1873: 295. Rossi, F., 1790. Fauna Etrusca, sistens Insecta quae in pro- vincias Florentina et Pisana praesertim collegit. — Pisa 1- 451 (2 vols.). Rossi, F., 1794. Mantissa Insectorum, exhibens species nup- er in Etruria collectas adjectis faunae Etruscae illustratio- nibus ac emendationibus (2). — Pisa. 154 pp. Sack, P., 1913a. Die Gattung Merodon Meigen (Lampetia Meigen olim.). — Abhandlungen des Senckenbergische Gesellschafts der Naturforscher 31: 427-462. Sack, 1913b. Zwei neue Paläarktische Merodon-Arten. — Annales Musei Nationalis Hungarici 11: 620-622. Sack, P., 1930. Zweifliigler (Diptera) IV: Schwebfliegen (Syrphidae). — Tierwelt Deutschlands 20: 1-118. Sack, P., 1931. Syrphidae. — /n: Lindner, die Fliegen der paläarktischen Region IV (6): 1-451. Sack, P., 1934. Lampetia crymensis Paramonov ®. — Konowia 13: 273-274. Sack, P., 1938. Zwei neue Syrphiden vom Balkan. — Konowia 17: 19-23. Schiner, J. R., 1857. Diptera Austriaca, Aufzahlung aller im Kaisertüme Oesterreichs bisher aufgefundene Zwei- flügler, III: die Oesterreichischen Syrphiden. — Verhand- lungen der Kaiserlichen Königlichen Zoologisch- Botanischen Gesellschaft in Wien 7: 279-506. Schiner, J. R., 1862. Fauna Austriaca, die Fliegen (Diptera) 1. Theil vi + xlii + 672 pp. Seguy, E., 1941. Dipteres accueillis par M. L. Berland dans le Sud Marocain, Syrphidae. — Annales du Société Entomologique de la France 110: 13-14. Séguy, E., 1961. Dipteres Syrphides de |’ Europe occiden- tale. — Mémoires du Musée d’ Histoire Naturelle, Nouvelle Série, Série A Zoologie 23: 1-248. Shiraki, T., 1930. Die Syrphiden des Japanischen Kaiserreiches, mit Beriicksichtigung der benachbarten Gebiete. — Memoirs of the Faculty of Agriculture, Taihoku Imperial University 1: i-xx; 1-446. Shiraki, T., 1968. Fauna Japonica II: Insecta Diptera, vol III Syrphidae, subfamily Merodontinae (Lampetiinae): 194- 252. Speight, M. C. D., 1987. External morphology of adult Syrphidae (Diptera). — Tijdschrift voor Entomologie 130: 141-175. Stackelberg, A. A. & Richter, V., 1968. Hower-Flies HURKMANS: Monograph of Merodon (Diptera: Syrphidae) of the Caucasus. — Trudy Vseso- yuznogo Entomologicheskogo Obschestva 52: 224-274. Strobl, G., 1893. Beitrage zur Dipterenfauna der Oesterreichischen Littorale. — Wiener Entomologische Zeitung 12 (2): 74-408. Strobl, G., 1900. Die Dipterenfauna von Bosnien, Hercegovina und Dalmatien. — Wissenschaftliche Mittei- lungen aus Bosnien und der Hercegovina (III. Natur- wissenschaft) 7: 352-670. Stuckenberg, B. R., 1956. The immature stages of Merodon bombiformis Hull, a potrential pest of bulbs in South Africa (Diptera: Syrphidae). — Journal of the Entomologi- cal Society of South Africa 19: 219-224. Suster, P., 1959. Diptera, Syrphidae. — Fauna Republicis Populare Romine, Insecta II pars III. Bucuresti. 286 pp. Szilady, L., 1940. Uber Paläarktische Syrphiden. — Annales Musei Nationalis Hungarici 33: 34-70. Thompson, F. C., 1988. Syrphidae (Diptera) described from unknown localities. — Journal of the New York Entomological Society 96 (2); 200-226. Treiber, R., 1987. Beobachtungen zur Okologie von Merodon rufus (Meigen, 1846). — Naturkundliche Beitrage des DJN 18: 64. Verrall, G. H., 1901. British Flies, vol. 3: Platypezidae, Pipunculidae, Syrphidae of Great Britain: 17-121 (Cata- logue); 127-691. — [reprint E. W. Massey, Faringdon 1969] Villeneuve, J., 1903. Contributions au Catalogue des Diptéres de France. — Feuille de la Jeunesse Naturaliste 33: 113-119. Villeneuve, J., 1909. Description d’ un nouveau Syrphide. — Wiener Entomologische Zeitung 28: 338-339. Villeneuve, J., 1910. Notes synonymiques. — Wiener Entomologische Zeitung 29: 304-305. Villeneuve, J., 1912. Notes synonymiques. — Wiener Entomologische Zeitung 31: 96-97. Villeneuve, J., 1924. Dipteres nouveaux. — Encyclopédie Entomologique (B) 2: Diptera 1: 5-8. Villeneuve de Janti, J., 1934. Notes diptérologiques. — Revue Francaise d’Entomologie 1: 180-181. Villers, R., 1789. Linnaeus, Entomologia, fauna Suecicae descriptionibus aucta, tomus tertius IV Insecta Diptera: Musca 1-657. — Lugduni [Lyon]. Violovitsh, N. A., 1983. Sirfydi Sibiri, Siberian Syrphidae (Diptera). — Verslagen en technische gegevens van het Instituut voor Taxonomische Zoologie (Zoologisch Museum), Universiteit van Amsterdam; [Translated by V. S. van der Goot and L. Verlinden; Foreword and keys only: 2-228. — published in Amsterdam, 1986]. Walker, F., 1849.List of the specimens of dipterous insects in the collection of the British Museum vol. 3: 485-687. Walker, F., 1852. Insecta Saundersiana, or characters of un- described insects in the collection of William Wilson Saunders, esq. — London. 474 pp. Walker, F., 1857a. Characters of undescribed Diptera in the collection of W. W. Sunders, esq. — Transactions of the Entomological Society of London, New Series 4: 119- 1572 Walker, F., 1857b. Catalogue of the dipterous insects col- lected at Sarawak, Borneo by Mr A. R. Wallace, with de- scriptions of new species. — Journal of the Proceedings of the Linnaean Society, Zoology vol. 1: 105-136. Walker, F., 1860. On Diptera collected at Makassar: Merodon. — Proceedings of the Linnaean Society of London 4: 120. 211 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Westwood, J. O., 1840. An introduction to the modern classification of insects: Synopsis of the genera of British Insects, Order 13: Diptera Aristotle (Antliata Fabricius, Halteriptera Clairv.). — pp. 125-158. Zimina, L. V., 1989. Novie Sirfidy Poda Merodon (Diptera, Syrphidae) iz Vostozhogo Kryma [New species of the ge- nus Merodon (Diptera, Syrphidae) from the east Crimea] — Vestnik Zoologia 1989 no. 1: 24-29. (in Russian). Received: January 1992. Revised version accepted: June 1993. Figs. 1-10. General morphology of Merodon. — Fig. 1. Lateral view of head. a, face; b, fions; c, oral margin; d, vertex; e, occi- put. — Fig. 2. Lateral view of antenna. a, antenna with 3rd article showing convex upper margin, apex rounded; b, 3rd article with upper margin basally convex, apically concave, apex acute; c, 3rd article straight margined, apex subacute; d, 3rd article strongly widened, upper margin convex, incised subapically. Lines above and below mark measured lengths of articles 2 and 3 respectively; the antennal ratio (length 3 : length 2) is often used in descriptions. — Fig. 3. Dorsal view of vertex and frons in d. Line portion a marks the length of tl, portion b the vertex height; angle alpha is vertex angle, beta is ocellar angle. — Fig. 4. Dorsal view of vertex and frons. (left), tl complete; (middle), tl incomplete: a linear strip separates the eyes, this condition might be termed subholoptic; (right), eyes separate (this occurs in some species in the d, and in all 9). The ocellar region is hatched. — Fig. 5. Lateral view of thorax (wings and legs omitted). 1, humeral knob; 2, dorsum with suture; 3, scutellum; 4, mesopleuron (katepisternum); 5, pteropleuron (anepisternum); 6, sternopleuron (anepimeron); 7, coxa I; 8, coxa II; 9, coxa III; W, wing insertion area; R, respiratory orifice (trachaea). — Fig. 6. Dorsal view of right wing. 1, anal cell (a); 2, discal me- dial cell (dm); 3, combined 4th+5th radial ce | (r4+5); 4, basal medial cell (bm); 5, basal radial cell (br); 6, 2nd+3rd radial cell (r2+3); 7, 1st radial cell (rl); 8, subcostal cell (sc); 9, costal cell (c); A, Costa (C); B, Subcosta (Sc); C, Ist branch of Radius (R1); D, main branch of Radius (R); E, 2nd & 3rd fused branches of Radius (R2+3); F, 4th & Sth fused branches of Radius (R4+5); G, Media (M); H, anterior Cubital (Cual); I, discal-medial Cubital (dm-cu); J, apical part of Media (M); K, analis (A); L, humeral (h); M, radial-medial (r-m); N, 2nd anterior Cubital (Cua2); O, basal part of Cual; P, basal-medial Cubital (bm-cu). — Fig. 7. Lateral view of leg III. a, trochanter (spine shown is present in d of some species only); b, femur, with c, triangular processus; d, tibia, with e, apical spur; f, metatarsus; g, tarsus. — Fig. 8. Ventral view of d genitalia (alagoezicus group). A, cercus; B, posterior surstyle lobe; C, apical margin of surstyle; D, ridge on inner face (medial face) of surstyle; E, epandrium; F, membraneous part of surstyle; G, apical extension of anterior surstyle lobe; H, anterior surstyle lobe; J, acces- sory lobe. — Fig. 9. Lateral view of & genitalia (alagoezicus group). A through J as in fig. 8; L, aedeagus; M, genital cap. — Fig. 10. Ventral view of aedeagus. A, apical shaft part; B, apex aedeagi; C, fringed plates; D, chitinous plates; E, subapical cavity; F, thecal ridge basally bordering same; G, basal humps. 212 Hurkmans: Monograph of Merodon Va A Gus À / 8 CE 4 : Po AIN TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Zz 7 18d 18e 18a 18b 18c Figs. 11-13. Aedeagus in Merodon. — 11, lateral view, lettering as in fig. 10, H, lateral ridge; 12, slender aedeagus with length- ened apical shaft part A; S, paired spines; 13, stout aedeagus with recumbent fringed plates. — Figs. 14-17. Lateral view of 3 genitalia of Merodon, lettering as in Fig. 9. — 14, M. apimimus; 15, M. distinctus, 16, M. nigritarsis, 17, M. longicornis. — Fig. 18. Lateral view of head in d of different Syrphidae. — a, Azpeytia scutellaris Walker; b, Merodon avidus, c, Eumerus strigatus Fallen; d, Mallota takasagoensis (Matsumura); e, Platynochaetus armipes Bezzi. 214 Hurkmans: Monograph of Merodon 33 20k 201 20m! 20n 200 ] Fig. 19. Dorsal view of right wing in d of different Syrphidae. — a, Eumerus strigatus Fallen; b, Azpeytia scutellaris Walker; c, Mallota takasagoensis (Matsumura); d, Merodon avidus e, Platynochaetus armipes Bezzi. — Fig. 20. Lateral view of leg III in d of different Syrphidae. — a, Merodon equestris, b, M. armipes, c, M. minutus, d, M. aberrans, e, M. ruficornis, f, Eumerus sulci- tibius Rondani; g, E. tuberculatus Rondani; h, E. olivaceus Loew and E. nudus Loew; j, Eumerus strigatus Fallen; k, Azpeytza scutellaris Walker; |, Azpeytia shirakii nomen novum; m, Mallota cimbiciformis Fallen; n, Mallota takasagoensis (Matsumura); o, Platynochaetus armipes Bezzi; p, Platynochaetus rufus (Macquart). 215 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Be % < = Pe iy Fig. 21. Merodon biarcuatus. a, b, variation in antennal shape; c, distal end of f3. — Fig. 22. M. clunipes. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 23. M. distinctus. a, lateral view of 3 genitalia; b, c, ventral and lat- eral view of aedeagus; d, habitus of 3. — Fig. 24. M. femoratus. a, b, ventral and lateral view of aedeagus; for lateral view of genitalia compare fig. 22a. — Fig. 25. M. mariae. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. 216 Hurkmans: Monograph of Merodon EE lil y == | VM NI i IM Fig. 26. Merodon ottomanus. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 27. M. testaceoides. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 28. M. alexeji. a, lateral view of d genitalia; b, ventral view of aedeagus. — Fig. 29. Posterior end of surstyle and cercus in M. alexeji (a) and M. altinosus (b). Pubescence de- leted to show difference in outlines. — Fig. 30. M. hirsutus. a, lateral view of & genitalia; b, c, ventral and lateral view of ae- deagus. — Fig. 31. M. hypochrysos. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. 277 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 = 4 303 32b Je SA | / ii) B Wp h lee NE My IA Fig. 32. Merodon kawamurae. a, lateral view of d genitalia; b, ventral view of overlapping posterior surstyle lobes; c, d, ven- tral and lateral view of aedeagus; e, size correlated shape variation in antennae. — Fig. 33. M. marginicornis. a, lateral view of g genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 34. M. rufitarsis. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 35. M. sophron. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus; over- lap of surstyle lobes as in Fig. 32b. 218 Hurkmans: Monograph of Merodon Tiii , Tiv Lan hr 39c Fig. 36. Merodon ankylogaster. a, lateral view of d genitalia; b, lateral view of posterior abdomen: T III, IV = tergites III and IV, S III, IV = sternites III and IV, A = appendage to S IV, G = genitalia. Pubescence on genital cap deleted. — Fig. 37. M. auronitens. a, lateral view of & genitalia; b, apex aedeagi; c, anterior view of apical end of tibia 3 and basitarsus 3. — Fig. 38. M. caudatus. a, lateral view of ¢ genitalia; b, c, ventral and lateral view of aedeagus; d, posterior view of distal tibiae 3 and metatarsi 3. — Fig. 39. M. oidipous. a, lateral view of d' genitalia; b, ventral view of aedeagus; c, lateral view of posterior abdo- men (lettering as fig. 36b); d, posterior view of metatarsi 3 and other tarsi 3. 219 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 INN SS SSSR N ESS Sì = = vy ys SO 2959 L En Pra a G x «6% GTA FIA Per SD HAE ARN [of VE 72s fe NS A I SS In DIES NS a 3 Fig. 40. Merodon persicus. a, lateral view of 3 genitalia; b, lateral view of posterior abdomen; c, posterior view of distal tibiae 3 and metatarsi 3, showing specialized hair-tuft on hump near base of metatarsi 3; d, lateral view of same hump, surrounding pubescence deleted. — Fig. 41. M. tangerensis. a, lateral view of d genitalia; b, ventral view of aedeagus; c, see 40c; d, lateral view of posterior abdomen. Pubescence on genital cap deleted. — Fig. 42. M. tarsatus. a, lateral view of & genitalia; b, c, ven- tral and lateral view of aedeagus; d, posterior view of distal tibiae 3 and metatarsi 3 showing specialized pubescent area; e, lat- eral view of posterior abdomen. 220 Hurkmans: Monograph of Merodon We. DE a Ns. Wd ande Fig. 43. Merodon turkestanicus. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus; d, posterior view of dis- tal tibiae 3 and metatarsi 3 (note extremely long floccus); e, lateral view of posterior abdomen. — Fig. 44. M. xanthipous. a, lat- eral view of & genitalia; b, lateral view of posterior abdomen; c, posterior view of distal tibiae 3 and metatarsi 3 (note long floccus). — Fig. 45. M. aberrans. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus; d, outline of abdo- men, d of aberrans aberrans (specimen of average size); e, same in aberrans isperensis. — Fig. 46. M. brevis. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. 22, TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 LS ) 4 IR \ TERESA A ESS ERIS ES Oe GORIS OOI RIDER OKRA PALI RSS 0% > % XS <2 RR es Y SER» COA KOSKY SR SKA Fig. 47. Merodon clavipes, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus; d, M. splendens, lateral view of d genitalia. — Fig. 48. M. cupreus. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 49. M. ha- mifer. a, lateral view of distal end of femur 3 and base of tibiae 3 in d ; b, same in 2. Fig. 50. M. lusitanicus (2). a, outline of femur 3; b, same in ® M. aberrans. — Fig. 51. M. quadrinotatus (® ). Dorsal view of abdomen, hatched area represents black pubescence; dotted, grey pubescence; blank, white pubescence. — Fig. 52. M. warnckei. a, lateral view of d genitalia; d, detail of same at different angle showing projection bearing pubescence on margin of posterior surstyle lobe; c, b, ventral and later- al view of aedeagus. — Fig. 53. M. pruni. a, lateral view of 8 genitalia; b, ventral view of aedeagus; c, d, lateral view of same in small and large specimen (note size correlated morphology). 222 Hurxmans: Monograph of Merodon ap FIR 7/4 iff il i Hi Su 179 N > Fig. 54. Colour patterns on abdomina of females: a, b, Merodon erivanicus (showing extremes of variation); c, M. longicornis, d, M. kaloceros. — Fig. 55. Outlines of antennae of females: a, M. erivanicus, b, M. longicornis, c, M. kaloceros. — Fig. 56. Merodon kaloceros. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus. - Fig. 57. M. longicornis. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 58. M. vandergooti. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 59. M. femoratoides. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus; d, outline of trochanter 3 in lateral view. 223 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 po 4 Se SE NINA U À N DIR, Ah EE TÉLÉ An ES Pa | 60 = mm A a Sz peser it Tn x < wat AU Mi fs Ta AN, seo = PS = = LS Di TAN £ Party | ; ES 60b n È LEER SS Fig. 60. Merodon nigritarsis. a, lateral view of d genitalia (lectotype); b, same from specimen from Handere, Turkey; c, d, ven- tral and lateral view of aedeagus (lectotype); e, f, same (Handere); g, variation of shape in outlines of trochanter 3. — Fig. 61. M. avidus. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus; d, ventral view of d genitalia; e, abdomi- nal colour pattern in pale 3; f, same in dark d; g, same in intermediate 2. — Fig. 62. M. crassifemoris. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 63. M. bequaerti. a, lateral view of & genitalia; b, c, ventral and lat- eral view of aedeagus. 224 Hurkmans: Monograph of Merodon 22 dl D a = = ZA = BET Fe Pf Fig. 64. Merodon elegans. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 65. M. manicatus. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 66. M. testaceus. a, lateral view of 3 genitalia; b, ©, ventral and lateral view of aedeagus. Pubescence on genital cap deleted. — Fig. 67. Distal ends of tibia 3 in: a, M. alagoezi- cus, b, M. schachti, c, M. satdagensis, d, M. lucasi. — Fig. 68. M. alagoezicus a, lateral view of 3 genitalia; b, c, ventral and lat- eral view of aedeagus; d, lateral view of triangular processus of femur 3. 225 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 he vi | tr, SN + ET TIZIA N Sy € I (y 2 S N RS aL TER Dy i} Fig. 69. Merodon lucasi. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus d, lateral view of triangular pro- cessus on femur 3.. — Fig. 70. M. nitidifrons. a, lateral view of & genitalia; b, c, ventral and lateral view of aedeagus; — Fig. 71. 8 M. satdagensis. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus. 226 HURKMANs: Monograph of Merodon at ig, == i @ | Sat — Al N ZB il Ps x N) \ FM net Im es 73a u N N N € 7 RES a 75 fo + Fig. 72. Merodon schachti. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 73. M. taniniensis. a, lateral view of d genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 74. M. toscanus. a, lateral view of 3 genitalia; b, c, ventral and lateral view of aedeagus. — Fig. 75. M. aureotibia. a, outline of antenna. 227 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Vi we ANN Fig. 76-79. Distribution of Merodon in biogeographic regions, hatched area: known conjunct range, dot or triangle: known localities. — 76, Ethiopian region; 77, western Palaearctic; 78, south-western Asia. — Fig. 79. Distribution of Merodon kaw- amurae, the only species with a certain natural occurrence in eastern Asia. — Fig. 80. Localities of Merodon in Turkey men- tioned in this paper: 1, Izmir Smyrna’). 2, Bursa [’Brussa’]; Karacabey lies slightly to the W, on the coast. 3, Ankara. 4, South of Ankara, near Ankara airport. 5, Aksehir [Ak-chehir]. 6, Konya. 7, Irmesan gedigi N of Akseki. 8, Sarihaci, 45 km W of Malatya. 9, Malatya [Melatia]. 10, Altı Haral Gölü NE of Celikhan. 11, Nemrut Dag near Adiyaman. 12, Buglan gecidi (pass) near Bingöl. 13, Palandöken Dagi (Kayak) SW of Erzurum. 14. Ovit pass, near Ispir. 15, near Karaurgan, 8 km W of Sarıkamıg. 16, Soganli railroad stop. 17, Handere, 20 km W of Sarikamis. 18, 11 km E of Karakurt. 19, Kars Deresi valley 5 km E of Sarıkamıs. 20, 20 km W of Uludere. 21, Akdamar, 40 km SW of Van. 22, Suvarihalil pass (on main east-west road through Hakkari province). 23, Mengene Dagi, N of Baskale. 24, Tanin-Tanin pass (on main east-west road through Hakkari province), W side. 25, Habero deresi valley, north of same road. 26. Tanin-Tanin pass near top. 27, Sat Dagları N of Mount Gavaruk. 28, Chilo Daglari. 29, Sat Dagları S of Sat Gölü. 30, Varegös, SW of Yüksekova. 31, Locality S of Yüksekova. 228 HURKMANs: Monograph of Merodon Fig. 81-88. Distribution of Merodon. — 81, M. alexeji (MM), M. sophron (©) and M. tener (4); 82, M. hirsutus (©), M. hypo- chrysos (A) and M. marginicornis (EB); 83, M. rufitarsis (8) and M. tarsatus (A); 84, M. ankylogaster (@), M. auronitens (W), M. caudatus (©) and M. oidipous (à); 85, M. persicus (©), M. quadrinotatus (4) and M. xanthipous (li); 86, M. brevis (©) and M. turkestanicus (MB); 87, M. kaloceros (WI) and M. tangerensis (A); 88, M. cupreus (©), M. hamifer (W), M. vandergooti (W) and M. warnckei (A). 229 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 89-93. Distribution of Merodon. — 89, M. aberrans (A) and M. clavipes (B); 90, M. aberrans isperensis (®); 91. M. velox (©) and M. splendens (A); 92, M. erivanicus (A); 93. M. longicornis (©) and M. lusitanicus (A). 230 HurkMANS: Monograph of Merodon Fig. 94-98. Distribution of Merodon. — 94, M. pruni (M); 95. M. crassifemoris (A) and M. femoratoides (©); 96, M. avidus (M); 97, M. nigritarsis (A); 98, M. bequaerti (©), M. manicatus (W) and M. testaceus (MM). 231 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 99-106. Distribution of Merodon. — 99, M. alagoezicus (©), M. dzhalitae (A), M. elegans (B) and M. toscanus (W); 100, M. schachti (©) and M. taniniensis (4); 101, M. nitidifrons (©) and M. satdagensis (A); 102, M. altinosus (A) and M. lucasi (©); 103. M. biarcuatus (A) and M. distinctus (©); 104, M. clunipes (MM) and M. femoratus (©); 105, M. mariae (@) and M. testaceoides (À); 106, M. aureotibia (A) and M. ottomanus (B). 292 INDEX References to the main treatment are in bold type. aberrans 147, 155, 156, 176-178, 181-183, 203-205, 215, 221, 222, 230 aberrans aberrans 176, 204, 221 aberrans flavitibius 176 aberrans isperensis 147, 177, 230 affinis 157, 203, 205 alagoezicus 156, 157, 185, 191, 193, 198, 200, 205, 206, 225,232 alagoezicus group 147, 150, 153, 157, 159, 192, 194, 196, 197-203, 205, 212 alexeji 154, 157, 163-169, 182, 204, 206, 217, 229 alexeji group 147, 153, 162-165, 167-169, 204 altinosus 147, 154, 163, 164-165, 204, 217, 232 amaryllidis 152 Amphoterus 148 angustiventris 207 ankylogaster 147, 154, 156, 169, 204, 219, 229 apimimus 214 armipes 214, 215 aureotibia 147, 157, 189, 203-205, 227, 232 aurifer 191, 205 auronitens 147, 154, 169, 170, 204, 219, 229 avidus 152, 153, 157, 180, 182, 185, 190, 191-194, 196, 203-206, 214, 215, 224, 231 avidus group 147, 150, 153, 191, 203, 205 Azpeytia 147, 150, 207, 208, 214, 215 balanus 207 bardus 207 bautias 207 bequaerti 147, 155, 194-197, 205, 224, 231 bessarabicus 153 biarcuatus 156, 157, 159, 204, 216, 232 bicolor 207 bipartitus 207 bombiformis 152 brevis 155, 176, 177, 178, 183, 205, 206, 221, 229 canipilus 178, 179, 205 x caudatus 151,154, 157, 169, 170, 171, 204, 205, 219, 229 chiragra 207 cimbiciformis 215 clauda 178, 179, 205, 206 clavipes 149, 155, 157, 178-186, 204-206, 222, 230 clavipes group 147, 153, 175, 181, 182, 204 clavipes var. albus 178, 180, 205 clavipes var. ater 178, 205 clavipes var. niger 178, 205 clunipes 154, 156, 158-160, 204, 216, 232 coerulea 207 contrarius 207 Copestylum 150 crassifemoris 153, 193, 194, 205, 206, 224, 231 crassifemoris group 147, 150, 153, 193, 205 cupreus 147, 155, 156, 179-181, 183, 205, 222, 229 curvipes 149, 178, 179, 205-207 dimorphus 159, 204 distinctus 150, 154, 156-159, 182, 204, 205, 214, 216, 232 distinctus assemblage 147, 153, 157, 204 dobrogensis 153 dzhalitae 155, 180, 181, 183, 205, 232 HURKMANS: Monograph of Merodon edentulus 207 elegans 147, 155, 157, 195-197, 204-206, 225, 232 elegans group 147, 153, 194, 195, 202, 205 equestris 149, 150, 152, 166, 215 Eristalis 149 erivanicus 157, 186-188, 205, 206, 223, 230 Eumerus 148, 150, 207, 214, 215 Exmerodon 149, 152, 167 femoratoides 156, 157, 189-191, 194, 205, 206, 223, 231 femoratus 154, 156, 158, 160, 161, 180, 204, 205, 216, 232 flavitibius 155, 156, 176, 178, 205 fulcratus 149, 152, 153, 167, 168 fulvus 185, 186, 205, 206 furcata 207 fuscinervis 185, 186, 205, 206 graecus 191, 205 gravipes 178, 179, 205, 206 grisea 207 hamifer 155, 156, 176, 181, 205, 222, 229 Helophilus 207 hirsutus 154, 156, 164, 165, 169, 204, 217, 229 hypochrysos 147, 154, 162, 165, 204, 217, 229 interveniens 207 isperensis 147, 155, 156, 177, 205, 230 italicus 191, 192, 205 kaloceros 147, 155, 157, 187, 188, 205, 223, 229 karadaghensis 155, 156, 181, 205 kawamurae 154, 157, 165, 166, 186, 204, 206, 218, 228 kiritshenkoi 153 knerii 176, 205 loewi 152 ‘ longicornis 155, 157, 186-188, 205, 214, 223, 230 longicornis group 147, 153, 162, 186, 188, 197, 203-205 lucasi 147, 156, 157, 198, 199, 205, 225, 226, 232 lusitanicus 147, 157, 181, 182, 205, 222, 230 Mallota 150, 207, 214, 215 manicatus 155, 157, 194-196, 205, 225, 231 marginicornis 147, 154, 162, 166, 167, 169, 204, 218, 229 mariae 147, 154, 157, 160, 161, 204, 216, 232 Megatrigon 148 melota 207 micromegas 165, 166, 204 minutus 215 morosus 207 nigritarsis 156, 157, 182, 190, 193, 194, 205, 214, 224, 25) nigritarsis group 147, 150, 153, 189, 203, 205 nitidifrons 147, 155, 157, 199, 200, 205, 226, 232 nudus 215 obscuripennis 176, 205 oidipous 147, 155, 171, 204, 219, 229 olivaceus 215 ornatus 207 Orthoprosopa 207 ottomanus 147, 154, 161, 162, 204, 217, 232 pallida 185, 205 Palpoda 207 persicus 147, 154, 171, 172, 204, 220, 229 Platynochaetus 150, 214, 215 Polydontomyia 207 pruni 153, 157, 183, 185, 186, 205, 206, 222, 231 pruni group 147, 153, 185, 205 pruni var. obscurus 185, 186, 205 quadrilineatus 191, 205 233 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 quadrinotatus 156, 182, 205, 222, 229 Quichuana 207 ruficornis 171, 195, 215 rufitarsis 154, 162, 167, 168, 204, 218, 229 rufitibius 191, 192, 205 rufus 152, 215 sacki 178, 179, 205 satdagensis 147, 156, 157, 198, 200, 201, 205, 225, 226, 232 schachti 147, 156, 157, 201, 205, 225, 227, 232 scutellaris 147, 207, 208, 214, 215 senilis 178, 179, 205 serrulatus 191, 205 shirakii 147, 208, 215 sicanus 185, 205 smirnovi 171-174, 204, 206 sophron 147, 154, 162-164, 168, 204, 218, 229 Sphegina 207 spinipes 189, 191, 193-195, 204-206 splendens 147, 155, 176, 182, 183, 205, 222, 230 strigatus 214, 215 sulcitibius 215 takasagoensis 214, 215 234 tangerensis 147, 154, 172, 173, 204, 220, 229 taniniensis 147, 155, 157, 201, 202, 205, 227, 232 tarsatus 154, 156, 167, 173, 174, 204, 206, 220, 229 tarsatus group 147, 153, 169, 171, 204 tenebricus 207 tener 153, 157, 168, 204, 229 testaceoides 147, 154, 162, 204, 217, 232 testaceus 155, 157, 162, 195-197, 205, 225, 231 torpidus 207 toscanus 147, 155, 157, 192, 202, 205, 227, 232 trizonus 169, 204 tuberculatus 207, 215 turkestanicus 155, 156, 169, 174, 204, 206, 221, 229 umbrifer 207 vandergooti 147, 153, 188, 189, 203, 205, 223, 229 vandergooti group 147, 150, 153, 188, 205 varicolor 207 velox 155, 157, 183-186, 205, 230 velox var. anathemus 183, 184, 205 velox var. armeniacus 183, 184, 205 viaticus 203, 204-206 warnckei 147, 155, 156, 176, 184, 185, 205, 222, 229 xanthipous 147, 154, 175, 204, 221, 229 HERMAN DE JONG Institute for Taxonomic Zoology, Amsterdam THE PHYLOGENY OF THE NEPHROTOMA FLAVESCENS SPECIES GROUP (DIPTERA: TIPULIDAE) De Jong, H., 1993. The phylogeny of the Nephrotoma flavescens species group (Diptera: Tipulidae). — Tijdschrift voor Entomologie 136: 235-256, figs. 1-114, tabs. 1-2, appendix. [ISSN 0040-7496]. Published 10 December 1993. The phylogeny of the currently recognized 22 species and subspecies of the west Palaearctic Nephrotoma flavescens species group (Diptera, Tipulidae) is analysed on the basis of 50 morpho- logical characters. The results of the present study are compared with the phylogeny of the fla- vescens group published by Oosterbroek (1980). A synopsis of the distribution of the species and subspecies is given. H. de Jong, Department of Entomology, Institute for Taxonomic Zoology (Zoological Museum), Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. Key words. — Diptera, Tipulidae, Nephrotoma, phylogeny, west Palaearctic. In this paper I will discuss the phylogenetic rela- tionships of the 22 species and subspecies of the Nephrotoma flavescens species group sensu Ooster- broek (1980). The concept of the flavescens group adopted here is in accordance with the views of Oosterbroek (1980, 1982). A list of the 22 species and subspecies of the flavescens group as recognized in the present paper is given in table 1. A systematic revision of the twenty species and subspecies of the flavescens group known at that time was incorporated in Oosterbroek (1978). Subsequently, Oosterbroek (1982) introduced N. cretensis and N. guestfalica hartigiana as new members of the group. Oosterbroek (1985a) furthermore in- cluded the Japanese species N. subpallida Alexander in the flavescens group. For reasons explained below (section “Discussion of adopted phylogeny’), I do not accept this combination. An as yet unnamed species, known of female specimens originating from north- ern Algeria and northern Tunisia, awaits description until the male sex becomes available. It is not includ- ed in the phylogenetic analysis of the present paper. The phylogeny of the flavescens group is analysed as part of a research project on the historical biogeogra- phy of the western Mediterranean (see De Jong in press). Previously, Oosterbroek (1980) presented a phylogenetic analysis of this species group, but addi- tional information on the distribution of certain char- acters within Nephrotoma prompted a reconsideration of Oosterbroek’s conclusions. In the section ‘Discussion of adopted phylogeny’, I will compare the results of the present study with Oosterbroek’s phylo- genetic analysis of the flavescens group. The flavescens group has an essentially west Palaearctic distribution, with most of its species and subspecies being confined to the Mediterranean sub- Table 1. The species and subspecies of the Nephrotoma fla- vescens species group as recognized in this paper. + = exam- ined; - = not examined. Females of an as yet undescribed species are known from north Algeria and north Tunisia. 3} 2 appendiculata appendiculata Pierre, 1919 + + appendiculata pertenua Oosterbroek, 1978 + + astigma Pierre, 1925 + + beckeri Mannheims, 1951 + + cretensis Oosterbroek, 1982 + + exastigma Oosterbroek, 1978 + + flavescens Linnaeus, 1758 + + fontana Oosterbroek, 1978 + + guestfalica guestfalica Westhoff, 1879 + + guestfalica hartigiana Oosterbroek, 1982 + + guestfalica surcoufi Pierre, 1925 + 4 lempkei Oosterbroek, 1978 + + minuscula Mannheims, 1951 + + nasuta Oosterbroek, 1975 + + quadrifaria quadrifaria Meigen, 1804 + + quadrifaria farsidica Savchenko, 1957 - + saccai Mannheims, 1951 + + schaeuffele Mannheims, 1964 + + spatha Oosterbroek, 1975 + + submaculosa Edwards, 1928 + + sullingtonensis Edwards, 1938 + + theowaldi Oosterbroek, 1978 + + 235 TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 1. Distribution of the Nephrotoma flavescens species group. region (fig. 1). The distribution range of six represen- tatives of the flavescens group extends towards north- west Europe. A single male specimen of the species flavescens Linnaeus has been reported from western Greenland (cf. Mannheims & Theowald 1971). Detailed information on the distribution of the species and subspecies of the flavescens group can be found in Oosterbroek (1978: distribution maps of the species and subspecies discussed therein) and Oosterbroek (1982, 1985b: additional distributional data). Oosterbroek & Theowald (1992) summarized the ranges of the Palaearctic Tipulidae, including the members of the flavescens group. Distribution maps of each of the species and subspecies of the flavescens group will be included in a forthcoming paper deal- ing with the historical biogeography of the western Mediterranean. A synopsis of the distribution of the members of the flavescens group is given at the end of the present paper. A key to the identification of the species and sub- species of the flavescens group (as well as all other western Palaearctic members of Nephrotoma known at the time) can be found in Oosterbroek (1979c). The more recently described N. cretensis and N. guest- falica hartigiana will key out under couplet 13 and 236 couplet 3 respectively. Oosterbroek (1982) listed dis- tinguishing characters for both taxa. MATERIAL, METHODS AND TERMINOLOGY The majority of specimens used for the present stu- dy are preserved in the collection of the Department of Entomology of the Institute for Taxonomic Zoology (Zoological Museum), Amsterdam. Addi- tional material was kindly lent by Dr. Eulalia Eiroa (Department of Animal Biology, University of Santiago de Compostela, Spain). Specimens were usually dry pinned, occasionally material was preser- ved in alcohol. As most characters used for the phylogenetic analy- sis pertain to structures of the terminalia of both sex- es, preparations of these parts were made. To this end the specimen was relaxed for a few hours in a humifi- er, after which the (end of the) abdomen was de- tached. The removed parts were macerated for about three to five minutes in a nearly boiling 10% KOH solution in a bain-marie. The macerated structures were rinsed with water (several times) and 70% alco- hol, upon which they were transferred to a watch- glass filled with glycerol. Examination of the termi- nalia was carried out using a Wild stereomicroscope with a magnification up to 100 X. The drawings were made with the aid of a drawing tube on the micro- scope. For lasting preservation the terminalia were stored in glycerol in a microvial pinned under the pertaining specimen. The programs HENNIG86, version 1.5 (Farris 1988) and PAUP, version 3.1.1 (Swofford 1993) were used to analyse the phylogeny. For details on the computa- tional methods employed, see the section ‘Discussion of adopted phylogeny’. The terminology used is largely in accordance with McAlpine (1981). I employ some additional terms to denote details of the male and female terminalia. Tangelder (1985) published a review of the morphol- ogy of the terminalia of the Tipulidae, and for further details I refer the reader to her paper. The terms used in the present text are explained in figures 3-10, 13, 14, 30 and 50. BIOLOGY The species of the flavescens group occur in a varie- ty of habitats, ranging from grassland, gardens, scrubs, deciduous as well as coniferous forests to dry sandy places. Adults are usually on the wing in spring and early summer, although minuscula Mannheims DE Jone: Phylogeny Nephrotoma flavescens group (Cyprus and the Levant) flies as early as the middle of January and records of flavescens Linnaeus (central and western Europe) are known from as late as September. The species of the flavescens group can be found from sea-level up to 2400 m, the highest alti- tude recorded for flavescens in the Alps. Several authors described immature stages of mem- bers of the group, viz. of appendiculata appendiculata Pierre, flavescens, guestfalica guestfalica Westhoff, quadrifaria quadrifaria Meigen, and submaculosa Edwards (cf. Brindle 1960, Chiswell 1956, Hemmingsen & Jensen 1972, Theowald 1957, 1967). Oosterbroek (1978) presented details on the biology of each of the then recognized members of the flavescens group. SYSTEMATIC POSITION Mannheims (1951) separated the west Palaearctic species of Nephrotoma with acutely pointed cerci in the female from the remaining species of the genus. Savchenko (1973), largely adopting this major divi- sion, included most of the species involved in his cor- nicina species group. The cornicina group sensu Savchenko contains the species of Nephrotoma that in the male carry an extension on the posterior margin Table 2. Character state matrix of the species and subspecies of the Nephrotoma flavescens species group. (Sub)species Character 5 10 15 20 25 30 35 40 45 50 appendiculata 00010 11000 11011 00100 01000 11011 10111 00000 00011 00000 a. pertenua 00010 11000 11011 00100 01000 11011 10111 00000 00011 00000 astigma 10010 00010 10111 10100 01110 01111 11000 10000 00011 01000 beckeri 00110 00000 10001 01110 01000 01010 00010 00000 00011 00010 cretensis 00110 00000 10001 01110 01000 01010 00000 00000 00011 00010 exastigma 10010 00000 10000 00100 01000 01111 00000 00000 00011 01000 flavescens 10010 00010 10111 10100 01110 01011 11000 10000 00011 01000 fontana 00010 00010 11011 00100 01100 01011 10000 : 00000 00011 00000 guestfalica 00011 00000 10001 10101 00001 01111 00000 01010 00011 10000 £ hartigiana 00011 00000 10001 10101 10001 01111 00000 01010 00011 10000 £ surcouf? 00011 00000 10001 10101 10001 01111 00000 01010 00011 10000 lempkei 10010 00011 10111 10100 01110 01111 11000 10000 00011 01100 minuscula 10010 10100 10011 00100 01000 01011 10111 00000 11011 00001 nasuta 00000 00000 10101 00100 01000 01010 00000 00000 00000 00000 quadrifaria 01000 00000 11011 01100 01000 01111 10010 00101 00101 00000 q. farsidica 01000 00000 11011 01100 01000 01111 10010 00101 00101 00000 SACCAL 00000 00000 1-001 00100 01000 01010 00010 00000 00010 00000 schaeuffelei 00000 00000 10000 00000 01000 00010 00000 00000 00100 00000 spatha -0010 00000 10001 10000 01000 01011 00000 00000 00100 00000 submaculosa 10010 00011 10111 10100 01110 01011 11000 10000 00011 01100 sullingtonensis 00010 00010 11011 00100 01000 01011 10000 00000 00011 00000 theowaldi 00010 10100 10011 10100 01000 01011 10111 00000 11011 00001 257 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 a LA = a 2 DE — © 0 2 © 5 o © ee EEE Sie NE Ze 2 = a * - E = © 3 o © o £ 5 2 S © E) 25 ee een: o © Ce] [= — — + © pesi wo ac) pe) £ E MCD ea CT OO OA 2 2 E ere SIE MO NNS SALE Tees He o 0 2 3 o U 2 0 a © = D © Dio. Si ES a € ee oee De CU. Cd un oo 0 £ 0 D © © DDD oo Gases D AS EU [134 (11016 [128 21 8,41, [128 10, 48 HAN De a 24, 32, 36 37, 39, 460017, 28, 43 2 [1 1, 13, 16, 47 O1, 4701 16 4, 44 6, 33,35 12 15 22 23 [134 9 3, 19, 49 [1 28 12, 14, 31 [117 [134 [113 44 114, 16, 18 30, 43 [143 15, 27 11,22, 29 Fig. 2. Consensus tree of two equally parsimonious trees with length 73, consistency index 68 and retention index 84 for the species and subspecies of the Nephrotoma flavescens species group. — synapomorphy; LJ: homoplasy; X : reversal. of sternite eight and that are devoid of bristles on the alar squama. Thus Savchenko excluded the species guestfalica Westhoff from his cornicina group, mean- while adding several central and east Palaearctic as well as Oriental species to the group. Based on three presumed synapomorphies, the fla- vescens species group was first recognized by Ooster- broek (1980) as a subgroup of the cornicina group. The monophyly of the flavescens group will be further discussed below. The exact position of the flavescens group within the cornicina group is not yet established. Ooster- broek (1980) presented a phylogeny of the cornicina group, but restricted his analysis to the western Palaearctic species. This approach, for obvious rea- sons, led to the conclusion that the flavescens group is closely related to western Palaearctic species of the cornicina group. I have made a preliminary analysis of the phylogeny of the cornicina group with the inclu- sion of some of its east Palaearctic and Oriental mem- bers. Results of this investigation show that the flaves- cens group is probably more closely related with east Palaearctic species of the cornicina group. An exhaus- tive analysis of the cornicina group, however, was be- 238 yond the scope of the present paper. In the phylogenetic analysis of the flavescens group I employed the non-flavescens group members of the cornicina group as the first level outgroup, and the re- maining species of Nephrotoma as the second level outgroup. PHYLOGENETIC ANALYSIS In this section I will discuss the characters used in the phylogenetic analysis of the flavescens group. Each character dealt with contains a brief description of its presumed plesiomorphous and apomorphous states, accompanied by their respective codings (0: plesio- morphous, 1: apomorphous). The character state ma- trix is given in table 2. The resulting cladogram for the flavescens group is shown in figure 2 and is discus- sed in the next section. Autapomorphies of the species and subspecies of the flavescens group are given in the appendix to this paper. Legs 1. — Tarsal claws of male: (0) medial tooth present (fig. 11, arrowed); (1) medial tooth absent (fig. 12) (cf. Oosterbroek 1980: 337, character 24). The male claws lack the medial tooth in the species exastigma, minuscula and the members of the clade astigma to submaculosa. In the species spatha the male claws usually bear this tooth, but there are specimens with some or all of the claws toothless. Absence of the medial tooth on the male claw is due to reduction. Females of Nephrotoma always have the tarsal claws devoid of teeth. Species with toothless claws in the male occur fre- quently in other groups of Nephrotoma. According to Oosterbroek (1980: 337) and Tangelder (1985: 160), this character state is usually restricted to smaller monophyletic groups. Referring, among other things, to this character state, Oosterbroek (1980) united the species exastigma and spatha with the members of the clade astigma to submaculosa, an interpretation not followed in the present paper. Wing 2. — Wing-membrane: (0) unmarked; (1) dark crossband running from pterostigma over base of discal cell, crossvein m-cu and apical portion of vein Cu; wing-tip darkened (fig. 13). The subspecies quadrifaria quadrifaria and quadri- faria farsidica are distinguished among the members of the flavescens group by dark shades below the pte- rostigma and on the wing-tip. Other species of Nephrotoma with similarly marked wings, e.g. atrosty- la Alexander, bifusca Alexander, gaganboi Tangelder, and neopratensis Alexander, are not closely related with quadrifaria. Male terminalia 3. — Aedeagal guide, apex: (0) gradually tapering towards tip (figs. 5, 14-17, 19-26, 28); (1) with knob-like enlarge- ment at tip (fig. 18). The species beckeri and cretensis are sister species that, among other things, are characterized by the shape of the aedeagal guide that terminates in an en- larged tip. Oosterbroek (1982) distinguished this shared derived character state in his discussion on cre- tensis (as ‘shape of central part of adminiculum’). 4. — Aedeagal guide, gonapophysis: (0) dorsal extension elongate (figs. 14, 16, 17); (1) reduced in length (figs. 5, 15, 18-20, 22-26, 28). The species spatha and the majority of species and subspecies of the clade beckeri to submaculosa either have a relatively short or completely reduced dorsal extension of the gonapophysis. Contrary to this, both subspecies of quadrifaria possess a relatively long dor- sal extension (fig. 21). I interpret the character state in quadrifaria as a reversal of character state 4(1). The majority of species of the cornicina group have DE Jone: Phylogeny Nephrotoma flavescens group relatively long dorsal extensions of the gonapophyses. Outside the cornicina group long dorsal extensions are known in species of the analis, brevipennis, dorsal- is, and pedunculata species groups of Nephrotoma (cf. Oosterbroek 1979b, 1984, 1985a; Tangelder 1983, 1984). 5. — Aedeagal guide, gonapophysis: (0) posterior exten- sion apically rounded (figs. 14-19, 21, 23-26, 28); (1) dor- soventrally flattened and with upcurved acute tip (fig. 20). The three subspecies of guestfalica show the apo- morphous state of this character. Within the flaves- cens group, the only other taxa with an acute dorsal tip of the posterior extension of the gonapophysis are both subspecies of appendiculata (fig. 22). In the lat- ter two subspecies the posterior extension is mediolat- erally flattened (see character 7). 6. — Aedeagal guide, gonapophysis: (0) posterior exten- sion parallel sided or tapering towards tip (figs. 5, 14-17, 19- 21, 24-26, 28); (1) spatulate at tip (figs. 22, 23). The members of the clade appendiculata to theo- waldi have the tip of the posterior extension of the gonapophysis dilated and laterally flattened, the pos- terior extension thus appearing as a spatulate struc- ture. In both subspecies of appendiculata the spatulate part has an acute dorsal tip (fig. 22; see next charac- ter), in minuscula and theowaldi it is rounded off and somewhat downcurved (fig. 23; see character 8). 7. — Aedeagal guide, gonapophysis: (0) tip of posterior ex- tension rounded off when seen in profile (figs. 14-17, 19, 21, 24-26, 28); (1) dorsally with acute tip (fig. 22). The two subspecies of appendiculata show the apo- morphous state of this character. The only other members of the flavescens group with a dorsally acute posterior extension of the gonapophysis are the three subspecies of guestfalica (fig. 20). Here the extension is dorsoventrally flattened and thus of an overall dif- ferent structure (see character 5). 8. — Aedeagal guide, gonapophysis: (0) ventral margin of posterior extension almost straight when seen in profile (figs. 5, 14-22, 24-26, 28); (1) downcurved at tip (fig. 23) (cf. Oosterbroek 1980: 337, character 25). The tip of the posterior extension of the gonapo- physis is somewhat downcurved in the species minus- culaand theowaldi. 9. — Aedeagal guide, gonapophysis: (0) variously shaped (figs. 14-23); (1) consisting of a mediolaterally flattened and relatively high anterodorsal part and an elongate posterior projection which gradually tapers towards its tip (figs. 5, 24- 26, 28). The species of the clade sullingtonensis to submacu- losa all have a similarly built aedeagal guide plus ap- 239 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 tg7 tg8 tg9 tg10 ogonst igonst lat shell centr rdg N NIT ANDER gon ventr app comp apod ant pt _ post apod ant lat pt = È aed ant apod 6 7 tg8 tg9 tg10 cerc A spmth genfk st9 240 pendages, which I interpret as a synapomorphy. 10. — Aedeagal guide, gonapophysis: (0) anterodorsal margin rounded (figs. 26, 27); (1) anterodorsal margin in- curved and acutely tipped (fig. 28, 29, arrowed) (cf. Oosterbroek 1980: 337, character 28). The apomorphous character state separates the spe- cies lempkei and submaculosa from the remainder of the flavescens group. 11. — Aedeagal guide, rod: (0) rod present ventral of sperm pump (fig. 30); (1) absent (fig. 31) (cf. Oosterbroek 1980: 332, character 12, as ‘adminicular rods’). The species of the flavescens group are devoid of rods of the aedeagal guide, although slight vestiges of these structures can be found in individual specimens of the species beckeri and saccai. Rods of the aedeagal guide are usually present in other species of Nephrotoma, including the species of the cornicina group outside the flavescens group. Reduction of the rods occurs throughout Nephrotoma (cf. Oosterbroek 1980: 349, character 5). 12. — Aedeagus: (0) basal, tubular section about 0.50 of total length or longer (figs. 32-35, 37, 39, junction of tubu- lar and branched sections of aedeagus arrowed); (1) basal, tubular section about 0.40 to 0.20 of total length (figs. 36, 38, junction of tubular and branched sections of aedeagus arrowed). Half of the species and subspecies of the clade quadrifaria to submaculosa have the branching point of the aedeagus situated before its midlength. Reversal of this state seems to have occurred twice within this clade, once in the ancestor of both minus- cula and theowaldi (cf. fig. 37) and once in the ances- tor of the clade astigma to submaculosa (cf. fig. 39). In the latter clade reversal of character 12(1) coincided with shortening of the aedeagus (see next character). The presumed plesiomorphous situation of charac- ter 12 can be found in non-flavescens group members of Nephrotoma with a partly tripartite aedeagus, e.g. alticrista Alexander, integra Alexander, medioproducta Alexander, nigrohalterata Edwards, and subpallida Alexander. DE Jone: Phylogeny Nephrotoma flavescens group 13. — Aedeagus: (0) relatively long and slender (figs. 32, 33, 35-38); (1) short and thick (figs. 34, 39). The species nasuta (fig. 34) and the members of the clade astigma to submaculosa (cf. fig. 39) have a rela- tively short aedeagus. Oosterbroek (1980: 336, char- acter 23) also mentioned the species exastigma and spatha as having a short aedeagus (as ‘intromittent organ’). Oosterbroek (1980: 350-351, diagram 1; 391-393, appendix 2B) related the length of the ae- deagus with the length of the second abdominal ter- gite. Contrary to this, the relative proportions of the aedeagus are perhaps better expressed by comparing the length of the aedeagus with its diameter. When the latter procedure is followed and the - arbitrary - limit between relatively long and short is appointed to a quotient of 100 for the total length of the aedeagus and its diameter just after the sperm pump, the aedea- gus of both exastigma (fig. 35) and spatha (fig. 33) turns out to be relatively long (cf. measurements in Oosterbroek 1980, appendix 2A). In the species of the cornicina group outside the fla- vescens group, the aedeagus is a relatively long structure. 14. — Gonocoxite: (0) dorsomedially fused with base of aedeagal guide (fig. 40); (1) separate (fig. 41, gap arrowed). The species of the clade quadrifaria to submaculosa show a membranous gap between the base of the ae- deagal guide and the adjacent dorsomedial corner of the gonocoxite. In the remaining species of the flaves- cens group both structures are fused, as they are in the other members of the cornicina group. Oosterbroek (1980: 337, character 26, as separation of ventral ap- pendages of adminiculum and sternite nine) distin- guished the same character state but recognized it in the species of the clade astigma to submaculosa only. Oosterbroek noted that the gap between both struc- tures can be narrow in other species of Nephrotoma. Several species of the flavescens group show this latter state, e.g. fontana. 15. — Gonocoxite, medisternal sclerotization: (0) fused with ventral appendage of aedeagal guide (fig. 42, connexion arrowed); (1) structures separate from each other (fig. 43, gap arrowed; figs. 8, 47-49). Figs. 3-10. Nephrotoma flavescens, 3-8, male; 9, 10, female. — 3, terminalia, lateral view; 4, tergite 10, ventral view; 5, aedea- gal guide, lateral view; 6, sperm pump, lateral view; 7, inner gonostylus, lateral view; 8, sternite 9 and gonocoxites, ventral view; 9, terminalia, lateral view; 10, hypogynial valve, medial view. Abbreviations: aed: aedeagus; aed gd: aedeagal guide; ant apod: anterior apodeme of sperm pump; ant lat pt: anterolateral part of inner gonostylus; ant pt: anterior part of inner gonostylus; centr rdg: central ridge of tergite 10; cerc: cercus; comp apod: compressor apodeme of sperm pump; crst: crest of inner gonostylus; ej duct: ejaculatory duct; gen fk: genital fork; gon: gon- apophysis of aedeagal guide; goncx: gonocoxite; hyp vlv: hypogynial valve; i gonst: inner gonostylus; lat shell: lateral shell of tergite 10; ma rdg: major ridge of hypogynial valve; med scl: medisternal sclerotization; mi rdg: minor ridge of hypogynial valve; o gonst: outer gonostylus; post apod: posterior apodeme of sperm pump; rdg: ridge of inner gonostylus; sp2: lateral remnant of genital bridge; spmth: spermatheca; st7 etc.: sternite 7 etc.; st8 ext: extension of sternite 8; tg7 etc.: tergite 7 etc.; ventr app: ventral appendage of aedeagal guide. 241 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Within the flavescens group, schaeuffelei and exas- tigma are the only species which have the ventral ap- pendages of the aedeagal guide fused with the medis- ternal sclerotizations. In all other members of the group these structures are separated by a membranous gap. When the medisternal sclerotizations are well de- veloped, they are united with the appendages of the aedeagal guide in the non-flavescens group members of the cornicina group. 16. — Gonocoxite, medisternal sclerotization: (0) well de- veloped, large (fig. 42, 44, 46-48); (1) reduced in size, small or absent (figs. 8, 43, 45, 49). It is assumed that the presence of medisternal scle- rotizations is a plesiomorphy within the genus Nephrotoma (Oosterbroek 1980: 321; Tangelder 1985: 144). Reduced medisternal sclerotizations oc- cur within the flavescens group in the species spatha (fig. 43), the three subspecies of guestfalica (cf. fig. 45), the species theowaldi, and the members of the clade astigma to submaculosa (figs. 8, 49). The shape of the reduced medisternal sclerotizations varies among these taxa. 17. — Gonocoxite, medisternal sclerotization: (0) not in contact with ventromedial margin of gonocoxite or only so along a relatively short zone (figs. 8, 42, 43, 45, 47-49); (1) fu- sed for more than half its length with gonocoxite (figs. 44, 46). When in contact with the gonocoxites, the medi- sternal sclerotizations are usually only fused with these structures near their anterior margins (cf. fig. 47). The sister species beckeri (fig. 44) and cretensis, and both subspecies of quadrifaria (cf. fig. 46) are characterized by a more complete fusion of both structures. I assume that fusion evolved independent- ly in both pairs of taxa. 18. — Inner gonostylus: (0) a projecting posterolateral part present (figs. 50-52, structure arrowed in figs. 51 and 52); (1) posterolateral part reduced (figs. 7, 53-66). Within the flavescens group, the species schaeuffelei and spatha have a projecting structure placed postero- laterally on the inner gonostylus. I interpret this structure as the homologue of the posterolateral part of the inner gonostylus of other genera and subgene- ra of Tipulidae. As such it is a plesiomorphy in Nephrotoma. In several species of the flavescens group its remnant is readily identifiable in the low protuber- ance covered with long hairs on the posterolateral margin of the inner gonostylus (e.g. nasuta, fig. 53; exastigma, fig. 57; appendiculata, fig. 62; minuscula, fig. 63; sullingtonensis, fig. 64). Several members of the cornicina group outside the flavescens group have a distinct projecting posterolat- eral extension on the inner gonostylus (fig. 50; cf. Oosterbroek 1980: 330, character 5). Oosterbroek interpreted this character state as an apomorphy of 242 his cornicina subgroup, but considering its occurrence outside and within Nephrotoma, it is probably better regarded as a plesiomorphy on this level of analysis. 19. — Inner gonostylus: (0) general shape various, seen in lateral view with relatively short and high anterior part (figs. 7, 50-53, 57-66); (1) general shape similar, in lateral view anterior part and crest appearing in line, elongate and slen- der (fig. 54). The inner gonostyli of the species beckeri and cre- tensis show a number of features that can be interpret- ed as apomorphies: the anterior part is acutely bent over laterally (fig. 55; less so in the other members of the flavescens group), the anterolateral part is smooth- ly curved when seen in dorsal aspect (fig. 56; acute in the other species of the flavescens group) and the crest is posteriorly produced into an acute tip (fig. 54; the only other representative of the flavescens group with an acute crest is the subspecies guestfalica guestfalica, fig. 58). Oosterbroek (1982) mentioned the synapo- morphous resemblance of the shape of the inner go- nostyli of beckeri and cretensis in his discussion on the latter species. 20. — Inner gonostylus: (0) general shape various, lateral edge toothless (figs. 7, 50-57, 61-66); (1) general shape si- milar, laterally with edge toothed (figs. 58-60). The three subspecies of guestfalica share some apo- morphous peculiarities in the structure of the inner gonostylus: the anterior part is large and produced, while the lateral margin is present as a rim carrying blackish sclerotized points. The inner gonostyli of the subspecies differ in degree of reduction of the dorsal crest (see next character). 21. — Inner gonostylus: (0) crest on anterior part well de- veloped, posteriorly extending (fig. 58); (1) crest reduced in length (figs. 59, 60). Oosterbroek (1980: 326, character 12) considered a posteriorly produced crest a synapomorphy of the species of the cornicina group. A posteriorly produced crest can therefore be interpreted as a plesiomorphy within the flavescens group. Reduction of the posteri- or extension of the crest is found in the subspecies guestfalica hartigiana and guestfalica surcoufi. In guest- falica hartigiana the crest is shortened (fig. 59), in guestfalica surcouf? it is completely absent (fig. 60). The members of the clade fontana to submaculosa also show reduction of the posterior extension of the crest. Reduction in this clade resulted in a distinct sit- uation and is treated under character 23. 22. — Inner gonostylus: (0) without ridge (fig. 50); (1) with ridge on its lateral side (figs. 7, 51-57, 61-66) (cf. Oosterbroek 1980: 332, character 11). All members of the flavescens group, with the ex- DE JONG: Phylogeny Nephrotoma flavescens group Figs. 11-29.—11, 12, Nephrotoma spatha, male last tarsal segment; 13, N. quadrifaria, wing; 14, N. schaeuffelei, aedeagal guide and gonapophysis, lateral view; 15, N. spatha, idem; 16, N. nasuta, idem; 17, N. saccai, idem; 18, N. beckeri, idem; 19, N. ex- astigma, idem; 20, N. guestfalica guestfalica, idem; 21, N. quadrifaria quadrifaria, idem; 22, N. appendiculata appendiculata, idem; 23, N. minuscula, idem; 24, N. sullingtonensis, idem; 25, N. fontana, idem; 26, 27, N. astigma, 26, idem; 27, aedeagal guide and gonapophysis, dorsal view; 28, 29, N. lempkei, 28, aedeagal guide and gonapophysis, lateral view; 29, aedeagal guide and gonapophysis, dorsal view. Abbreviations: aed gd: aedeagal guide; Cu: cubital vein; d: discal cell; dors ext: dorsal extension of gonapophysis; m-cu: me- dio-cubital crossvein; post ext: posterior extension of gonapophysis; sq: squama. 243 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 ception of the three subspecies of guestfalica, are char- acterized by the presence of a ridge on the posterior part of the inner gonostylus. (Contrary to Oosterbroek’s assertion, nasuta does have a ridge underneath the crest; cf. fig. 53, arrowed). Most other species of the cornicina group, and all other Nephrotoma examined, lack this feature. The only representatives of the cornicina group with a ridge-like structure are cornicina cornicina Linnaeus, cornicina sardiniensis Oosterbroek, and moravica Figs. 30-39. — 30, Nephrotoma ligulata, sperm pump and part of aedeagus, lateral view; 31, 32, N. schaeuffelei, 31, idem; 32, sperm pump and aedeagus, lateral view; 33, N. spatha, idem; 34, N. nasuta, idem; 35, N. exastigma, idem; 36, N. quadrifar- ia quadrifaria, idem; 37, N. minuscula, idem; 38, N. fontana, idem; 39, N. astigma, idem. Abbreviation: rod: rod of aedeagal guide. 244 Martinovsky. These three taxa together form a monophyletic group within the cornicina group (cf. Oosterbroek 1980). Here the ridge-like structure probably evolved independently of that of the flaves- cens group. 23. — Inner gonostylus: (0) crest on anterior part well de- veloped, posteriorly extending (figs. 52-58, 61-64); (1) crest reduced in length, posteriorly laterally downcurved (figs. 7, 65, 66). In the clade fontana to submaculosa, the apomor- phous state of this character can be found. Oosterbroek (1980: 337, character 27) recognized this apomorphy for the species of the clade astigma to submaculosa. I claim that it also applies to the species fontana (fig. 65). 24. — Inner gonostylus: (0) ridge without pubescence (figs. 51-57, 61-65); (1) ridge covered with dense pubescen- ce (fig. 7, 66) (cf. Oosterbroek 1980: 332, character 11X). The species of the clade astigma to submaculosa are distinguished from the remainder of species and sub- species of the flavescens group, by the presence of pu- bescence on the ridge of the inner gonostylus. 25. — Outer gonostylus: (0) at base with anteromedially directed extension (figs. 67, 69); (1) with elongate ventrola- teral extension (fig. 68). The outer gonostylus of the majority of species of Nephrotoma is provided with a more or less well de- veloped anteromedially directed extension at its base. In the three subspecies of guestfalica the outer gono- stylus basally bears a long ventrolaterally directed ex- tension. 26. — Outer gonostylus: (0) elongate, without strikingly long hairs (figs. 67, 68); (1) broad, long haired along poste- rior margin, with longest hairs about as long as width of ou- ter gonostylus (fig. 69). The outer gonostylus of the majority of species of Nephrotoma is a rather slender structure when seen in lateral view, covered with moderately long hairs. In the two subspecies of appendiculata, the outer gono- stylus is relatively broad and bears long hairs along its posterior margin. 27. — Sperm pump: (0) sclerotized part of ejaculatory duct attached to sperm pump (figs. 30, 31, 70, ejaculatory duct arrowed); (1) membranous gap between sperm pump and sclerotized part of ejaculatory duct (figs. 6, 71-73, 75) (cf. Oosterbroek 1980: 332, character 14). In the flavescens group the sclerotized part of the ejaculatory duct is attached to the sperm pump in the species schaeuffelei only (figs. 31, 70). In the other species and subspecies of the group there either is a membranous gap between the body of the sperm DE Jone: Phylogeny Nephrotoma flavescens group pump and the sclerotized part of the ejaculatory duct or the ejaculatory duct is not sclerotized at all (see next character). The gap between the sperm pump and the sclerotized part of the duct is narrow in the species beckeri (fig. 73), in the other relevant species and subspecies of the flavescens group it is wider (figs. 1972245) Oosterbroek (1980: 326, character 11) recognized the presence of a midposterior appendage to the sperm pump (as ‘aedeagus’) as a synapomorphy for the cornicina group. I interpret the midposterior ap- pendage as a sclerotization of the wall of the ejacula- tory duct (seminal duct of Byers 1961). The species of the cornicina group outside the flavescens group have this sclerotized structure well developed, being at- tached to the sperm pump (fig. 30). 28. — Sperm pump: (0) vestige of sclerotized portion of ejaculatory duct present (figs. 6, 71-73, 75); (1) absent (fig. 74). As mentioned under the preceding character, some members of the flavescens group do not show a trace of sclerotization of the ejaculatory duct. The species and subspecies involved are exastigma (fig. 74), the subspecies of guestfalica and quadrifaria, and astigma and lempkei. It seems that reduction of the sclerotiza- tion of the ejaculatory duct evolved along several lin- eages and to various degrees within the flavescens group. 29. — Sperm pump, anterior apodemes: (0) without long membranous extensions at tip (fig. 30); (1) with long mem- branous extensions at tip (fig. 6, 31, 71). All members of the flavescens group are character- ized by the possession of long translucent extensions at the tips of the anterior apodemes of the sperm pump. I did not find such extensions in species of Nephrotoma outside the flavescens group. 30. — Sperm pump, anterior apodeme: (0) broad, bulging in dorsal view (figs. 70, 73); (1) reduced to narrow strip (figs. 72, 74, 75). The anterior apodemes of the sperm pump are usu- ally well developed in Nephrotoma. Within the flaves- cens group, reduced anterior apodemes are present in the species spatha (fig. 72) and the members of the clade exastigma to submaculosa (figs. 74, 75). Oosterbroek (1980: 333, character 15) also recog- nized a reduction of the anterior apodemes (as ‘later- al appendages of the aedeagus’) as an apomorphy in the flavescens group. The largest anterior apodemes in the flavescens group are found in the species nasuta and saccai. Oosterbroek distinguished a monophylet- ic group composed of the remainder of species and subspecies of the flavescens group on account of this feature. As relatively large anterior apodemes are also 245 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 7 post lat pt Figs. 40-56. — 40, Nephrotoma guestfalica guestfalica, base of aedeagal guide in part plus right gonocoxite, posterolateral view; 41, N. quadrifaria quadrifaria, idem; 42, N. schaeuffelei, male sternite 9, gonocoxites and medisternal sclerotizations, ventral view; 43, N. spatha, idem; 44, N. beckeri, idem; 45, N. guestfalica guestfalica, idem; 46, N. quadrifaria quadrifaria, idem; 47, N. appendiculata appendiculata, idem; 48, N. minuscula, idem; 49, N. astigma, idem; 50, N. ligulata, inner gonostylus, later- al view; 51, N. schaeuffelei, idem; 52, N. spatha, idem; 53, N. nasuta, idem; 54-56, N. beckeri, inner gonostylus, 54, lateral view; 55, anterior view; 56, dorsal view. Abbreviation: post lat pt: posterolateral part of inner gonostylus. 246 present in schaeuffelei, beckeri, and cretensis, I do not adopt this interpretation. 31. — Sperm pump, compressor apodeme: (0) deeply in- cised (fig. 76); (1) almost entire, dorsal margin either slight- ly concave or entirely convex (fig. 77). Within the flavescens group, the species and sub- species of the clade quadrifaria to submaculosa have a compressor apodeme which consists of an undivided structure. 32. — Sperm pump, compressor apodeme: (0) entire, with midposterior keel; (1) without midposterior keel. The more basal species and subspecies of the clade quadrifaria to submaculosa have the compressor apo- deme provided with a well developed midposterior keel. The keel is completely reduced in the species of the clade astigma to submaculosa. 33. — Sperm pump, posterior apodeme: (0) yellowish co- loured, elongate (figs. 6, 30, 31, 70-72, 74); (1) dark colou- red and short (fig. 75). Dark sclerotized, short posterior apodemes occur in the species of the clade appendiculata to theowaldi of the flavescens group. The remainder of species of the group have the posterior apodemes less heavily sclerotized. In the clade appendiculata to theowaldi sclero- tization of the posterior apodemes coincides with re- duction of these structures. Oosterbroek (1980: 336, character 22) recognized short posterior apodemes (as ‘posterior appendages of the aedeagus’) as an apo- morphy of the clade appendiculata to theowaldi. However, reduced posterior apodemes are also known in the species nasuta and beckeri (fig. 73). The apodomes in the two latter species are not blackish sclerotized. 34. — Sp2: (0) haired (figs. 78, 83, arrowed); (1) bare (figs. 79-82). As in the majority of other species of the cornicina group, most members of the flavescens group have the posteromedial margin of sp2 near midlength provid- ed with a concentration of hairs. Within the flavescens group, these hairs are absent in the species saccai (fig. 79), beckeri (fig. 80) and in the clade quadrifaria to theowaldi (cf. figs. 81, 82). 35. — Sp2: (0) variously shaped (figs. 78-81, 83); (1) si- milar (fig. 82). The species of the clade appendiculata to theowaldi possess a similarly built sp2. The bare structure has the dorsal part well developed and anteriorly protrud- ing, while the ventral part is present as a straight ven- tromedially directed appendix. DE Jone: Phylogeny Nephrotoma flavescens group 36. — Sp2: (0) variously shaped (figs. 78-82); (1) similar (fig. 83). In the clade astigma to submaculosa the sp2 struc- ture has a small and protruding dorsal part, while it is slightly produced posteriorly near midlength at the point of insertion of the hairs. The ventral part is straight and gradually tapers towards its tip. 37. — Sternite 8: (0) with posterior extension (figs. 3, 84, 86); (1) without (fig. 85). The only members of the flavescens group without an extension on the posterior margin of sternite 8 are the three subspecies of guestfalica. All other species of the cornicina group have a well-developed posterior extension of sternite 8. 38. — Sternite 8: (0) posterior extension dorsoventrally compressed (fig. 84); (1) laterally compressed (fig. 86). The extension of sternite 8 is short and laterally compressed in both subspecies of quadrifaria. The majority of species of the cornicina group have the ex- tension either dorsoventrally flattened or rounded in cross-section. Only cornicina cornicina, cornicina sar- diniensis and moravica have the extreme apex of the extension laterally compressed. However, in these taxa the posterior extension of sternite 8 is dorsoven- trally flattened at its base and elongate. 39. — Tergite 10: (0) variously shaped (figs. 3, 4, 89-94); (1) consisting of two cup-shaped structures provided with black spines along their anteromedial and posterior margins, each carrying an acute lateral extension (figs. 87, 88). The three subspecies of guestfalica are characterized by their uniquely shaped tergite 10. Oosterbroek (1980: 336, character 17P) considered guestfalica and quadrifaria sister taxa, a relationship presumed, among other things, to be supported by similarities in the structure of tergite 10 (as ‘extension of tergite 9’). In my opinion the shape of tergite 10 in quadrifaria (figs. 89, 90) has no apomorphy in common with that of guestfalica (figs. 87, 88). 40. — Tergite 10: (0) variously shaped, with relatively nar- row medial incision, caudally directed (figs. 3, 4, 87, 88, 91- 94); (1) posteriorly widely emarginate, caudodorsally direc- ted (figs. 89, 90). The two subspecies of quadrifaria are characterized by their peculiarly shaped tergite 10. 41. — Tergite 10: (0) central ridges posteriorly not or only slightly produced (fig. 91); (1) central ridges posteriorly pro- duced (fig. 93) (cf. Oosterbroek 1980: 336, character 17S; tergite 10 as ‘extension of tergite 9’). The central part of tergite 10 protrudes only mod- erately in most species of the flavescens group. Only 247 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 minuscula and theowaldi show a posterior elonga- tion of this part of tergite 10. 42. — Tergite 10: (0) lateral shells rounded along ventral margin when seen in lateral view (fig. 92); (1) ventral mar- gin truncate (fig. 94). The species minuscula and theowaldi are the only members of the flavescens group which have the ven- tral margin of the lateral shells of tergite 10 straight. In the other members of the group, as well as in most other species of Nephrotoma, this margin is rounded. Female terminalia 43. — Cercus: (0) long, well-developed, gradually tapering towards tip (fig. 99); (1) reduced in length, tip rounded off (figs. 95, 97, 106). Within the flavescens group a shortened, blunt- tipped cercus is known in the species schaeuffelei (fig. 95), spatha (fig. 97), and the two subspecies of quad- rifaria (fig. 106). Besides being reduced in length, the cercus of both subspecies of quadrifaria is also re- duced in height. In the remainder of species and sub- species of the flavescens group the cercus is long and either gradually tapers towards the tip (fig. 99), or is produced into a nipple-like extension (see next char- acter). Oosterbroek (1980: 326, character 14 and page 328, character 1) listed the presence of pointed cerci in the female as one of the synapomorphies of the cor- nicina group. The cerci in this group are also more ro- bust than is usual in Nephrotoma (cf. Savchenko 1973: 35, 119). The plesiomorphous form of the cer- cus in the cornicina group can be found in supposed- ly basal species like forcipata Pierre and ligulata Alexander. Here the well-developed cercus gradually tapers towards its tip. In the flavescens group a similar- ly shaped cercus is found in the species nasuta (fig. 99) and in the as yet undescribed species from Algeria and Tunisia. The cercus of nasuta and the north-west African species probably represents the plesiomor- phous form in the flavescens group. 44, — Cercus: (0) gradually tapering towards tip (fig. 99); (1) tip produced into a nipple-like extension (figs. 9, 101). With the exception of both subspecies of quadri- faria, the species and subspecies of the clade saccai to submaculosa are characterized by the possession of a nipple-like projection of the tip of the cerci. (The nipple-like projection is frequently worn off in older specimens). As stated under the preceding character, the plesio- morphous condition of the female cercus in the fla- vescens group is probably represented by a moderately pointed structure as found in nasuta (fig. 99). 45. — Hypogynial valve: (0) tip entire, with ventral mar- 248 gin evenly convex (figs. 96, 98, 100, 102); (1) tip irregular, appearing to be broken off (figs. 9, 10, 103-105, 108-110) (cf. Oosterbroek 1980: 333, character 16). The species of the clade beckeri to submaculosa show the apomorphous condition of this character. Within the cornicina group a similar situation is known in the non-flavescens group species saghaliensis Alexander, where it apparently evolved independent- ly (cf. fig. 6 of Oosterbroek 1985a. Fig. 13 of Oosterbroek 1985a, depicts a damaged specimen of esakii Alexander in which the tip of the hypogynial valve is actually broken off; in intact specimens of this species the hypogynial valve terminates in an acute tip). The basal species of the flavescens group, as well as the other members of the cornicina group, have slen- der, reduced hypogynial valves of which the ventral margin of the tip is entire (cf. figs. 96, 98, 100, 102). 46. — Hypogynial valve: (0) dorsal margin continuous (figs. 9, 10, 96, 98, 100, 102-104, 107-110); (1) dorsal mar- gin with posteriorly directed point at about two-thirds of its length (fig. 105). The three subspecies of guestfalica are, among oth- er things, characterized by their apomorphously shaped hypogynial valves. 47. — Hypogynial valve: (0) major and minor ridges ba- sally in contact with each other (figs. 102, 103, 108); (1) se- parate (figs. 10, 109, 110). In the Tipulidae there are usually two ridges on the medial side of the hypogynial valve, a major (dorsal) and a minor (ventral) one (cf. fig. 10). As this situa- tion is found also in the majority of species of Nephrotoma, I agree with Oosterbroek (1980: 326), who interpreted the presence of two ridges as a plesio- morphy for the genus. Tangelder (1985: 151), how- ever, considered the same character state an apomor- phy within Nephrotoma. When present, both ridges are usually separate, like they are in the species schaeuffelei of the flavescens group (fig. 96). A partly or entirely reduced minor ridge is frequently found in the flavescens group, e.g. in schaeuffelei (fig. 96), spatha (fig. 98), nasuta (fig. 100) and the subspecies of guestfalica (fig. 105). In other species of the group the minor ridge is well de- veloped and both major and minor ridges are fused at their base, e.g. in saccai (fig. 102), beckeri (fig. 103), cretensis, the subspecies of appendiculata and in fonta- na (fig. 108). In the sistergroup of fontana, viz. the clade astigma to submaculosa, both ridges are separate. Judged by the situation in the nearest outgroups of this clade, this trait can be interpreted as an apomorphy (cf. figs. 109, 110). The species exastigma (fig. 104) also has the ridges separate, probably as the result of a secon- DE Jone: Phylogeny Nephrotoma flavescens group Figs. 57-77. — 57, Nephrotoma exastigma, inner gonostylus, lateral view; 58, N. guestfalica guestfalica, idem; 59, N. guestfalica hartigiana, idem; 60, N. guestfalica surcoufi, idem; 61, N. quadrifaria quadrifaria, idem; 62, N. appendiculata appendiculata, idem; 63, N. minuscula, idem; 64, N. sullingtonensis, idem; 65, N. fontana, idem; 66, N. astigma, idem; 67, N. exastigma, out- er gonostylus, lateral view; 68, N. guestfalica guestfalica, idem; 69, N. appendiculata appendiculata, idem; 70, N. schaeuffelei, sperm pump, dorsal view, compressor apodeme omitted; 71, 72, N. spatha, 71, sperm pump, lateral view; 72, sperm pump, dorsal view, compressor apodeme omitted; 73, N. beckeri, idem; 74, N. exastigma, idem; 75, N. appendiculata appendiculata, idem; 76, N. guestfalica guestfalica, compressor apodeme, anterior view; 77, N. quadrifaria quadrifaria, idem. 249 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 78-100. — 78, Nephrotoma nasuta, sp2, medial view; 79, N. saccai, idem; 80, N. beckeri, idem; 81, N. quadrifaria quad- rifaria, idem; 82, N. appendiculata appendiculata, idem; 83, N. astigma, idem; 84, N. exastigma, posterior margin of male ster- nite 8, posterolateral view; 85, N. guestfalica guestfalica, idem; 86, N. quadrifaria quadrifaria, idem; 87, 88, N. guestfalica guestfalica. 87, male tergite 10, ventral view; 88, male tergite 10, lateral view; 89, 90, N. quadrifaria quadrifaria, 89, male ter- gite 10, ventral view; 90, male tergite 10, lateral view; 91, 92, N. appendiculata appendiculata, 91, male tergite 10, dorsal view; 92, male tergite 10, lateral view; 93, 94, N. minuscula, 93, male tergite 10, dorsal view; 94, male tergite 10, lateral view; 95, 96, N. schaeuffelei, 95, cercus, lateral view; 96, hypogynial valve, medial view; 97, 98, N. spatha, 97, cercus, lateral view; 98, hypogynial valve, medial view; 99, 100, N. nasuta, 99, cercus, lateral view; 100, hypogynial valve, medial view. dary development. (There is in my opinion no com- pelling reason to consider the dorsal ridge in exastig- ma and in the clade astigma to submaculosa a new structure, non-homologous with the major ridge of the other members of the flavescens group; cf. Oosterbroek 1980: 336, character 19A and B). 250 48. — Hypogynial valve: (0) minor ridge short (fig. 109); (1) posteriorly extended (fig. 110). The minor ridge is elongate in the species lempkei and submaculosa, reaching towards the tip of the hy- pogynial valve. 49. — Sternite 9: (0) entire (fig. 112); (1) with break be- tween its midposterior and anterolateral parts (fig. 111, ar- rowed). The species beckeri and cretensis are the only repre- sentatives of the flavescens group that have a membra- nous interruption in between the broad anterolateral portion of sternite 9 and its slender posterior projec- tion. 50. — Sternite 9: (0) yellowish coloured (fig. 111); (1) an- teriorly blackish coloured, i.e. heavily sclerotized (fig. 112, arrowed). Contrary to the other members of the group, the anterior margin of sternite 9 is blackish coloured in the species minuscula and theowaldi. DISCUSSION OF ADOPTED PHYLOGENY Phylogenetic analysis of the character state matrix given in table 2, with a zero-vector as outgroup, resul- ted in two most parsimonious trees with length 73, consistency index 68, and retention index 84. I used exact tree-calculating methods with both computer programs employed, viz. the ‘ie’ (implicit enumera- tion) command of HENNIG86 and the branch-and- bound algorithm of PAUP. The binary coded charac- ters had the default weight 1. No a priori or a posteriori adaptations of the data were introduced to adjust the results of the analysis. Both programs pro- duced the same pair of equally parsimonious trees. The two trees obtained differ in the position of the species flavescens. It either is the sister species of a clade (astigma (lempkei, submaculosa)) or constitutes a trichotomy with astigma and the clade (lempkei, sub- maculosa). In the first tree character state 28(1) is con- sidered an apomorphy for the clade (astigma (lempkei, submaculosa)), with subsequent reversal to the plesio- morphous state in submaculosa. The second tree de- picts character state 28(1) as a homoplasy in the spe- cies astigma and lempkei. The second tree is topologically identical with the strict consensus tree which is shown in fig. 2. The four most basal species in the cladogram split off consecutively. Next in the transition series follows the pair of sister species beckeri and cretensis. The remainder of the cladogram contains several lineages of which the monophyly is only weakly cor- roborated. The clade exastigma to submaculosa is substantiated by a single homoplasy, viz. the narrow anterior apo- demes of the sperm pump, which are also found in the species spatha (character 30). The clade contain- ing the species exastigma and the three subspecies of guestfalica is also relatively weakly supported by one homoplasy only (character 28). The character state involved, viz. the complete reduction of the sclero- tized part of the ejaculatory duct, also occurs in the DE Jone: Phylogeny Nephrotoma flavescens group two subspecies of quadrifaria, as well as in astigma and lempkei. The clade quadrifaria to theowaldi is the third line- age within the cladogram of which the monophyly is demonstrated by a single homoplasy, in this case the bare sp2 (character 34). Within the flavescens group the same character state is known in the species saccai and beckeri. In certain aspects the results of the phylogenetic analysis of the present paper differ considerably from those presented by Oosterbroek (1980: 333, clado- gram 2B; see also Oosterbroek & Arntzen 1992). Fig. 113 shows the topology of Oosterbroek’s cladogram of the flavescens group. For ease of comparison, the topology of the consensus tree of the present paper is given in fig. 114. Oosterbroek (1980: 332) listed three synapomor- phies in support of the monophyly of the flavescens group, two of which he considered weak and one strong. According to his views, the members of the flavescens group should be distinguished by a straight- ened dorsal margin of the gonocoxite (‘dorsal margin of the male sternite nine is flattened’; Oosterbroek 1980: 332, character 10; considered weak). Ooster- broek noted that this character state is absent in the species nasuta and spatha. My observations demon- strate that the posterodorsal edge of the gonocoxite is curved also in exastigma, the three subspecies of guest- falica (cf. fig. 38) and in schaeuffelei. I therefore do not accept the straightened dorsal margin of the gon- ocoxite as a synapomorphy of the flavescens group. The other two synapomorphies of the flavescens group distinguished by Oosterbroek are also used in the present paper. They are the presence of the mem- branous ridge underneath the free part of the crest (Oosterbroek 1980: 332, character 11; considered weak because of its presumed absence in nasuta and guestfalica, but see character 22 of the present paper) and the absence of the rods of the aedeagal guide (Oosterbroek 1980: 332, character 12; considered strong; cf. character 11 of the present paper). Oosterbroek (1980: 332, characters 13 and 14; 346, character 2) distinguished saccai as the sister spe- cies of a clade containing the remainder of species and subspecies of the flavescens group. The monophyly of this latter clade was substantiated by the presence of a partly trifid aedeagus and the reduction of the sclero- tized part of the ejaculatory duct. However, contrary to Oosterbroek’s (1980, 1985a) assertion, non-flavescens group species with a partly trifid aedeagus are known within Nephrotoma. East Palaearctic and Oriental Nephrotoma species showing this feature include alticrista Alexander, dodabettae Alexander, inorata Alexander, integra Alexander, me- dioproducta Alexander, nigrohalterata Edwards, rajah Alexander, and subpallida Alexander. These species 251 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 101-112. — 101, 102, N. saccai, 101, cercus, lateral view; 102, hypogynial valve, medial view; 103, N. beckeri, hypogy- nial valve, medial view; 104, N. exastigma, idem; 105, N. guestfalica guestfalica, idem; 106, 107, N. quadrifaria quadrifaria, 106, cercus, lateral view; 107, hypogynial valve, medial view; 108, N. fontana, idem; 109, N. astigma, idem; 110, N. lempkei, idem; 111, N. beckeri, female sternite 9, dorsal view; 112, N. minuscula, idem. do not belong to the cornicina group in its current conception. It should be noted, moreover, that the majority of the Afrotropical species of Nephrotoma have the apical section of the aedeagus modified. The apical portion in these species looks like a ribbon or a flattened tube with sclerotized lateral margins, much as in the east Palaearctic and Oriental species with a trifid aedeagus, but then without the medial filament. A partly trifid aedeagus, which is also known in spe- cies of the tipulid subgenera Tipula (Acutipula Alexander), 7. (Lunatipula Edwards), T. (Odonatisca Savchenko), 7. (Papuatipula Alexander), and 7. (Pterelachisus Rondani) could well represent a plesio- morphy in Nephrotoma. I therefore assume that the structure of the aedeagus in saccai, which is tubular throughout, represents an autapomorphy of the spe- cies (cf. appendix). Likewise, the reduction of the sclerotized part of the ejaculatory apodeme is evaluat- 252 ed differently in the present study (cf. characters 27 and 28). In this context it should be noted that Oosterbroek (1985a: 247) assigned the Japanese species subpallida Alexander to the flavescens group on account of its partly trifid aedeagus. When a trifid aedeagus is re- garded a plesiomorphy on this level of analysis, the feature - of course - advances no argument to incor- porate any species in the flavescens group. Moreover, the male of subpallida has well developed rods of the aedeagal guide, a plesiomorphous character state not found in the flavescens group (cf. character 11 of the present paper). The ovipositor of subpallida has blunt-tipped, well-developed cerci and large hypogy- nial valves, both structures thus being of a different, presumably plesiomorphous, type compared with the cerci and hypogynial valves of the members of the fla- vescens group (cf. characters 43 and 45 of the present text). For these reasons I do not include subpallida in the flavescens group. Next in Oosterbroek’s transition series is the spe- cies nasuta, which was considered the sister species of the remainder of the flavescens group on account of the plesiomorphously large anterior apodemes of the sperm pump (cf. Oosterbroek 1980: 333, character 15). Reduction of the size of the anterior apodemes is employed in a slightly different manner and at an- other branching point in the present paper (cf. char- acter 30). The sister group of schaeuffelei in Oosterbroek’s phylogeny is characterized by the serrate hypogynial valves (cf. Oosterbroek 1980: 333, character 16). However, incorporated in this clade is the species spa- tha, a species of which the female sex was unknown to Oosterbroek in 1980. The female of spatha was de- scribed by Eiroa & Novoa (1987) based on material collected in Galicia, Spain. It has the ventral margin of the tip of the hypogynial valve smoothly curved (fig. 98) and can therefore not be placed in the clade distinguished by the serrate hypogynial valve (cf. character 45 of the present paper). [Oosterbroek & Artzen (1992) excluded spatha from their cladogram of the flavescens group]. Figures 113 and 114 both depict the species pair beckeri and cretensis as the sister group of the remain- der of species and subspecies of the flavescens group. According to Oosterbroek’s views, guestfalica and quadrifaria should be considered sister taxa. This re- lationship was postulated on account of the reduction of the hypogynial valves (Oosterbroek 1980: 334, character 16B), the shape of tergite 10 in the male (loc. cit.: 336, character 17P) and the presence of bristles on the alar squama (as ‘neala’; loc. cit.: 336, character 18). The shape of the hypogynial valves of both species is very different (cf. figs. 105, 107). The same holds for the shape of male tergite 10 (cf. figs. 87-90, and characters 37 and 38 of the present paper). Although probably a plesiomorphy in the family Tipulidae, the presence of bristles on the squa- ma is quite unique in the genus Nephrotoma. I have to assume that these bristles developed independently in both guestfalica and quadrifaria. The position of the species fontana in the clado- gram of Oosterbroek differs noticably from its allot- ted place in the consensus tree of the present text. The sister group of fontana is, according to Oosterbroek, characterized by the absence of the downcurved part of the minor ridge of the hypogynial valve (Oosterbroek 1980: 336, character 19). However, not all members of this ‘sister group’ lack the down- curved part of the minor ridge, viz. both subspecies of appendiculata and flavescens (fig. 10). My interpreta- tion of the position and state of development of the ridges of the hypogynial valve, which deviates from DE Jong: Phylogeny Nephrotoma flavescens group Oosterbroek’s views, is explained under characters 47 and 48 of the present text. The clade appendiculata to theowaldi of Oosterbroek fully concords with my ideas on the re- lationships of the species involved. The exact phylogenetic relationship of the species sullingtonensis was left unresolved by Oosterbroek. On account of the shape of the gonapophysis of the aedeagal guide it can be considered to be more close- ly related with the species of the clade astigma to sub- maculosa (cf. character 9 of the present paper). According to Oosterbroek, exastigma is closely re- lated with the species of the clade astigma to submac- ulosa. He corroborated this postulate by the presence of both minor and major ridges of the hypogynial valves (Oosterbroek 1980: 336, character 19A, with the major ridge assumed to represent a newly devel- oped structure), the short aedeagus (loc. cit.: 336, character 23), and the toothless claws in the male (loc. cit.: 337, character 24). The three characters are dis- cussed under the respective characters 47, 13 and 1 of the present text. My argumentation results in a more basal position of exastigma in the flavescens group. Oosterbroek’s clade astigma to submaculosa is iden- tical to the clade astigma to submaculosa presented here, inclusive of the trichotomy I was unable to re- solve. In summary, both our cladograms (figs. 113, 114) recognize the same basal species in the flavescens group, with the exception of the species spatha. In Oosterbroek’s solution spatha occupies a more de- rived position. The same applies to exastigma, which is placed next to spatha in Oosterbroek’s phylogeny. A sister group relationship of guestfalica and quadri- faria, as suggested by Oosterbroek, was not substan- tiated in the present study. The species fontana has a more derived position in my consensus tree. The tri- chotomy of which sullingtonensis forms part in fig. 113 is resolved in fig. 114. Here sw/ingtonensis is de- picted as the sister species of the clade fontana.to sub- maculosa. Both Oosterbroek’s and my solutions agree on the composition of the clades appendiculata to theowaldi and astigma to submaculosa. Differences between figs. 113 and 114 are the result of selection and interpretation of characters. They do not repre- sent artefacts of the analytical methods employed, viz. the ‘manual’ approach of Oosterbroek versus the computational procedure of the present study. DISTRIBUTION In this section I will give a synopsis of the distribu- tion of the species and subspecies of the flavescens group. For the time being, more detailed information can be found in Oosterbroek (1978, 1982, 1985b) and in Oosterbroek & Theowald (1992). 253 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 saccai nasuta schaeuffelei becker cretensis guestfalica g. hartigiana g. surcoufi quadrifaria q. farsidica fontana appendiculata a. pertenua minuscula theowaldi sullingtonensis exastigma spatha astigma flavescens lempkei submaculosa Fig. 113. (Left) Topology of the phylogeny of the Nephrotoma flavescens species group according to Oosterbroek (1980) and Oosterbroek & Arntzen (1992). The most basal species in the phylogeny of the fla- vescens group, schaeuffelei, is known from the Elburz mountains (Iran) and the Talysch mountains (Azerbaydzhan) to the south-west of the Caspian Sea. The species spatha is endemic to north-west Spain. The species nasuta is known from a few localites in western, central and eastern Turkey, while saccai oc- curs in south-west Italy and northern Sicily. The species pair beckeri and cretensis has an eastern Mediterranean distribution, with beckeri being dis- tributed in Cyprus and the adjacent mainland of Turkey, as well as Lebanon and Israel. Its sister spe- cies is endemic to Crete. The species exastigma is confined to northern Algeria and Tunesia. The three subspecies of guestfalica are essentially west Mediterranean, with the nominate subspecies having the largest range of the three, extending from England and south Sweden in the northwest, Spain in the south-west to western Turkey in the east. The subspecies guestfalica hartigiana is endemic to 254 schaeuffelei spatha nasuta saccai beckeri cretensis exastigma guestfalica g. hartigiana g. Surcoufi quadrifaria q. farsidica appendiculata a. pertenua minuscula theowaldi sullingtonensis fontana astigma flavescens lempkei submaculosa Fig. 114. (Right) Topology of the consensus tree of the Nephrotoma flavescens species group of the present paper. Sardinia, while the subspecies guestfalica surcoufi is distributed in Morocco, Algeria and Tunesia. The nominate subspecies of quadrifaria is distrib- uted over west and central Europe with its range ex- tending towards the east over the Crimea peninsula and the Caucasus mountain range. The subspecies quadrifaria farsidica is known from north-west Iran in the area adjacent to the Caspian Sea. The nominate subspecies of appendiculata is large- ly distributed over west and central Europe, eastwards reaching towards the westcoast of the Caspian Sea and southwards to Anatolia. It is absent from south- west Europe (Spain, Italy) where it is replaced by the subspecies appendiculata pertenua. In this context the occurrence of appendiculata sensu stricto on Sardinia is remarkable. The subspecies appendiculata pertenua is known from north-west Morocco and south-west Europe, including Corsica, Sicily and Malta. In its turn, this subspecies is absent from Sardinia. The spe- cies minuscula is known from Cyprus and the Levant, theowaldi occurs in south and south-west Turkey and in the Greek island Rhodos. The species sullingtonensis is distributed from northern Morocco through Spain and France to southernmost England. The species fontana is en- demic to Algeria, while astigma is confined to the north of Algeria and Tunesia. Both flavescens and sub- maculosa are widespread throughout western and cen- tral Europe, with submaculosa occurring in Corsica, Sardinia, Sicily and the mainland of Italy. The species flavescens is absent in the four last mentioned areas. The species /empkei is at present known only from the islands Mallorca and Menorca. ACKNOWLEDGEMENTS For the loan of material of both sexes of the species Nephrotoma spatha\ would like to thank Eulalia Eiroa (Santiago de Compostella, Spain). Pjotr Oosterbroek and Frederick Schram, and an anonymous referee, read and commented upon an earlier draft of this paper. Although I did not follow all of their suggestions for adjustment of the text, I greatly appreciate their willingness to review the paper. The investigations were supported by the Foundation for Biological Research (BION), which is subsidized by the Netherlands Organization for Scientific Research (NWO). REFERENCES Brindle, A., 1960. The larvae and pupae of the British Tipulinae (Diptera: Tipulidae). — Transactions of the Society for British Entomology 14: 64-114. Byers, G. W., 1961. The crane fly genus Dolichopeza in North America. — University of Kansas Science Bulletin 42: 665-924. Chiswell, J. R., 1956. 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Tipulidae. — Fliegen der palaearktis- chen Region 15, Lieferung 167: 1-64, pls. I-VII. DE JONG: Phylogeny Nephrotoma flavescens group Mannheims, B. & B. Theowald, 1971. Die Tipuliden von Grönland (Diptera, Tipulidae). — Bonner zoologische Beiträge 22: 332-342. Oosterbroek, P., 1978. The western palaearctic species of Nephrotoma Meigen, 1803, (Diptera, Tipulidae) Part 1. — Beaufortia 27: 1-137. Oosterbroek, P., 1979a. The western palaearctic species of Nephrotoma Meigen, 1803, (Diptera, Tipulidae) Part 2. — Beaufortia 28: 57-111. Oosterbroek, P., 1979b. The western palaearctic species of Nephrotoma Meigen, 1803 (Diptera, Tipulidae) Part 3. — Beaufortia 28: 157-203. Oosterbroek, P., 1979c. The western palaearctic species of Nephrotoma Meigen, 1803 (Diptera, Tipulidae) Part 4, including a key to the species. — Beaufortia 29: 129-197. Oosterbroek, P., 1980. The western palaearctic species of Nephrotoma Meigen, 1803 (Diptera, Tipulidae), Part 5, Phylogeny and Biogeography. — Beaufortia 29: 311-393. Oosterbroek, P., 1982. New taxa and data of western Palaearctic Nephrotoma (Diptera: Tipulidae). - Entomologische Berichten, Amsterdam 42: 41-44. Oosterbroek, 1984. A revision of the crane-fly genus Nephrotoma Meigen, 1803, in North America (Diptera, Tipulidae). Part Il: non-dorsalis species-groups. — Beaufortia 34: 117-180. Oosterbroek, P., 1985a. The Nephrotoma species of Japan (Diptera, Tipulidae). — Tijdschrift voor Entomologie 127: 235-278. Oosterbroek, P., 1985b. Some Tipulidae new for Corsica and Sardinia (Diptera). — Entomologische Berichten, Amsterdam 45: 121-122. Oosterbroek, P. & J. W. Arntzen, 1992. Area-cladograms of Circum-Mediterranean taxa in relation to Mediterranean palaeogeography. — Journal of Biogeography 19: 3-20. Oosterbroek, P. & B. Theowald, 1992. Family Tipulidae. — Catalogue of Palaearctic Diptera 1: 56-178. Savchenko, E. N., 1973. Crane flies (fam. Tipulidae), sub- fam. Tipulinae (conclusion) and Flabelliferinae. — Fauna SSSR (N.S. 105) Two-winged insects II (5): 1-281. (In Russian). Swofford, D. L., 1993. PAUP, Phylogenetic Analysis Using Parsimony, version 3.1. — Illinois Natural History Survey, Champaign, Illinois. Tangelder, I. R. M., 1983. A revision of the crane fly genus Nephrotoma Meigen, 1803, in North America (Diptera, Tipulidae). Part I: the dorsalis species-group. — Beaufortia 33: 111-205. _ Tangelder, I. R. M., 1984. The species of the Nephrotoma dorsalis-group in the Palaearctic (Diptera, Tipulidae). — Beaufortia 34: 15-92. Tangelder, I. R. M., 1985. Phylogeny of the Nephrotoma dorsalis species-group (Diptera, Tipulidae), mainly based on genital characters. — Beaufortia 35: 135-174. Theowald, B., 1957. Die Entwicklungsstadien der Tipuliden (Diptera, Nematocera), insbesondere der west- palaearktischen Arten. — Tijdschrift voor Entomologie 100: 195-308. Theowald, B., 1967. Familie Tipulidae. (Diptera, Nematocera). Larven und Puppen. = Bestimmungsbücher zur Bodenfauna Europas 7: 1-100. Received: 12 May 1993 Accepted: 27 August 1993 295) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 APPENDIX In this section the recognized autapomorphies of the species and subspecies of the Nephrotoma flaves- cens group are listed. appendiculata appendiculata: no autapomorphies recog- nized. appendiculata pertenua: differing from nominal subspecies in details of male tergite 10 only. astigma: shape of lateral shells of male tergite 10 in lateral view. beckeri: shape of male tergite 10; shape of inner gonostylus. cretensis. shape of compressor apodeme of sperm pump. exastigma: lateral prescutal stripes straight; inner gonostylus with sclerotized ridge from anterior to lateral part. flavescens medial prescutal stripes with dull lateral margins, downcurved part of lateral prescutal stripes dull. fontana: shape of extension of male sternite eight; shape of gonapophysis of aedeagal guide. guestfalica guestfalica: position and shape of crest of inner go- nostylus. guestfalica hartigiana: crest of inner gonostylus partly re- duced. 256 guestfalica surcoufi: crest of inner gonostylus entirely re- duced. lempkei: medial projection on male sternite eight slender, somewhat tapering towards tip. minuscula: aedeagal guide with dorsolaterally extending ridges; form and length of central ridges of male tergite 10. nasuta: nasus long; shape of male tergite 10; shape of aedea- gus, particularly at branching point. quadrifaria quadrifaria: shape of inner gonostylus; shape of male tergite 19. quadrifaria farsidica: medial projection of male sternite eight directed dorsally. saccat: shape of male tergite 10; shape of gonapophysis of ae- deagal guide; sperm pump posteriorly with upcurved plate; aedeagus single throughout; genital fork reduced. schaeuffelei: shape of inner gonostylus; outer gonostylus an- teriorly with pubescent incurved swelling; shape of sp2. spatha: hairs on vertex short; shape of male tergite 10. submaculosa: no autapomorphies recognized. sullingtonensis. prescutal stripes partly confluent. theowaldi: aedeagus near branching point with pair of later- al extensions; crest of inner gonostylus twisted along its longitudinal axis. A. NEBOISS Museum of Victoria, Melbourne, Australia NEW SPECIES OF THE GENUS MOLANNA CURTIS FROM SULAWESI (TRICHOPTERA: MOLANNIDAE) Neboiss, A., 1993. New species of the genus Molanna Curtis from Sulawesi (Trichoptera: Molannidae). — Tijdschrift voor Entomologie 136: 257-258, figs. 1-4 [ISSN 0040-7496]. Published 10 December 1993. The description and figures of Molanna jolandae sp. n. are given. This is the second species re- corded from Sulawesi, and extends the range of generic distribution to southern Sulawesi Tenggara. A. Neboiss, Department of Entomology, Museum of Victoria, 71 Victoria Crescent, Abbotsford 3067, Victoria, Australia. Key words. — Trichoptera; Molannidae; Molanna; Sulawesi; taxonomy. The present paper is based on material recently collected in Sulawesi. Among the numerous speci- mens from a number of localities, members of the family Molannidae are scarce and have been collected at only one location. There is only one previous Sulawesi record, a single specimen on which the de- scription of Molanna cupripennis Ulmer (1906) is based. The type locality ‘Bonthain Peak’ is in south- western Sulawesi (Sulawesi Selatan). The present ma- terial is from the southeastern part of the island (Sulawesi Tenggara) near Moramo (4°09’S 122°38’E). It may be noted, that the extensive collec- tion from Dumoga Bone National Park in the north of the island (Sulawesi Utara), accumulated by the Wallace Expedition 1985, did not contain any speci- mens of Molanna. MATERIAL AND METHODS Specimens were prepared by clearing the abdomen in cold KOH solution, then transferring to glycerol for drawing and later storage in microvial. Drawings of genitalia were made with the assistance of a camera lucida on a Wild M20 compound microscope. The wings, prepared as temporary microscope mounts in glycerol, were examined and drawn using a Wild M8 dissecting microscope also equipped with camera lu- cida. All dissected and figured specimens are identi- fied by the author’s notebook number with prefix ‘PT-.... All specimens have been deposited in the National Museum of Natural History, Leiden, The Netherlands (RMNH). ‘TAXONOMIC PART Molanna jolandae sp.n. (figs. 1-3) Type material. — Holotype d: Sulawesi Tenggara, Moramo (4°09’S 122°38’E), Sungai Sena, 50 m asl, 15 Nov 1989, R. de Jong and J. Huisman (site S 8946: mixed forest, dense undergrowth) in RMNH [Genitalia prep. PT-1968 fig- ured, specimen dry mounted]. — Paratypes: Sulawesi Tenggara, Moramo, Sungai Moramo, 200 m asl, 16 Nov. 1989, at light, R. de Jong and J. Huisman, 26 (site S 8947: mixed forest, dense undergrowth) in RMNH [wing prep. PT- 1969 figured; both specimens in alcohol]. Description Wings slender, rounded apically (fig. 3), length d 8.6-9.0 mm, densely covered with short, dark brown pubescence, hind wing anal margin with long, dark brown fringe, venation similar to that of Molanna cupripennis Ulmer. Male. — Dark, blackish brown, antennae, maxillary and labial palpi as well as fore legs densely covered with coarse blackish brown hairs, mid- and hind-tarsi also covered with dark hairs, whereas femora and tib- iae are pale yellow with scattered small, dark blackish semi-erect spines, apical spurs covered with dark hairs, preapical spurs with paler hairs. Genitalia (figs. 1, 2) with abdominal segment IX in lateral view broadly rounded ventrally, mid-dorsal margin broad with slight mesal depression, anterolat- eral angles may be more or less produced. Superior appendages elongate ovoid, flattened, lower basal margin produced into an incurved lobe. Segment X large, extended and strongly curved ventrad, dorsom- esally separated almost to the base, usually with three strong spines on dorsal margin on either side. Inferior 257 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-4. Molanna species. — 1, M. jolandae sp. n., male genitalia (holotype), lateral; 2, ventral; 3, wing venation (paratype); 4, M. cupripennis Ulmer, male genitalia lateral (from Ulmer 1951). appendages short, two branched, lower branch with terminal half curved dorsad, a distinct spine apically, upper branch directed posteriorly, slightly concave, apex rounded. Phallus short, rather robust, strongly curved, membranous, small chitinous spine internal- ly. Female unknown. Remarks This species is very similar to Molanna cupripennis Ulmer, but, besides being smaller (length of forewing 9 mm instead of 14 mm), differs by the presence of spines on segment X, and the shape of the inferior ap- pendages, the branches of which are not incised api- cally. The figures by Ulmer (1906, 1951) (here repro- duced as fig. 4) also show notable differences in shape and position of segment IX and the superior appen- dages. Etymology. — The species is named after Jolanda Huisman in recognition of her efforts gathering ex- 258 tensive material of Trichoptera from many localities in Sulawesi. ACKNOWLEDGEMENT I wish to express my sincere thanks to Jan van Tol of RMNH Leiden for access to this most interesting material. REFERENCES Ulmer, G., 1906. Neuer Beitrag zur Kenntnis aussereu- ropaische Trichopteren. — Notes from the Leyden Museum 28: 1-116. Ulmer, G., 1951. Kôcherfliegen (Trichopteren) von den Sunda-Inseln (Teil 1). — Archiv fiir Hydrobiologie (Supplement) 19: 1-528 + plate 1-28. Received: 16 March 1993 Accepted: 15 April 1993 NICO NIESER' & PING PING CHEN’ 1 Tiel, The Netherlands 2 Beijing Academy of Agriculture & Forestry Sciences, Beijing, P. R. China THE RHAGOVELIA (HETEROPTERA: VELIIDAE) OF SULAWESI (INDONESIA) Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), IV. Nieser, N. & P. P. Chen, 1993. The Rhagovelia (Heteroptera: Veliidae) of Sulawesi (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), IV. — Tijdschrift voor Entomologie 136:259-281, figs. 1-84 [issn 0040-7496]. Published 10 December 1993. Apart from additional records for several species for Sulawesi, ten new species are described: Rhagovelia blagiokommena sp. n., Rhagovelia chrysomalla sp. n., R. daktylophora sp. n., R. horaia sp. n., À. pseudocelebensis sp. n., R. robina sp. n., R. trichota sp. n., R. tropidata sp. n. from Sulawesi, À. kastanoparuphe sp. n. from Sulawesi and Buton and R. kalami sp. n. from Buton. A revised key to species from Sulawesi and Buton is given. Additional records for Rhagovelia from Sabah are given in an appendix. Correspondence: Dr. N. Nieser, Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands. Key words. — Sulawesi (= Celebes); Buton; Sabah; Veliidae; Rhagovelia, key; new species. As our contribution to the Fauna Malesiana pro- gram, we are working on the Nepomorpha and Gerromorpha mainly collected by staff members of the Zoological Museum Amsterdam and the National Museum of Natural History Leiden. The present one is the fourth in the series, the first dealt with Naucoridae, Nepidae and Notonectidae (Nieser & Chen 1991), while the second and third treated the Gerridae (Nieser & Chen 1992, Chen & Nieser 1992) Rhagovelia is a large tropicopolitan genus of small, stream inhabiting, Veliidae. About 150 species have been described so far, but the number of extant spe- cies is probably twice as high. The subfamily Rhagoveliinae is characterized by a large, deeply cleft, apical segment of the middle tarsus with a fan-like structure (fig. 1). This is considered an adaption to li- ving on waters with a strong current. Two Oriental genera are included in the subfamily, viz. Tetraripis, with both middle and hind tarsi with cleft apical seg- ment provided with a fan-like structure, and Rhagovelia, with only middle tarsi modified in this way. Tetraripis is only known from Sri Lanka and western Malaysia. Formerly, it was thought that tropical America would be much richer in species of Rhagovelia than Asia (Bacon 1956, Hungerford & Matsuda 1961, Lundblad 1936, 1937). In a recent revision, however, Polhemus & Polhemus (1988) described 26 new spe- cies from Borneo, Sulawesi and Maluku (the Moluc- cas). Only two species were previously known from this area. They stated, moreover, that they had many more undescribed Indo-Australian species in their collection. Thus the apparent paucity of species in tropical Asia is at least partly due to insufficient knowledge of the fauna. Although a complete survey of the Rhagovelia of Sulawesi was included in the revision by Polhemus & Polhemus (1988), we recognised again ten new spe- cies in the collections we studied, thus doubling the number of known species. As very probably more spe- cies are still to be discovered, the key to species from Sulawesi and Buton presented here is to be used with care. New records of this genus for Sabah are presen- ted in appendix 1. Measurements are in millimetres, length has been measured in dorsal view from the anterior margin of the head along the central axis to the posterior margin of the last visible abdominal tergite in apterous, and to the apex of the hemielytra in macropterous speci- mens. ‘Width’ has been measured at the base of the connexiva in dorsal view. The first two segments of the fore tarsi have not been measured, as they are small (0.02-0.03), usually partly hidden in an excava- tion of the fore tibia and thus difficult to measure pre- cisely. Moreover, they do not provide specific charac- ters. For the same reason the measurements of the first two segments of the hind tarsi have been omit- ted. The seventh abdominal segment, on which the connexiva end, is the last before the modified genital segments. In females the eighth tergite is sometimes 259) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 1-16. — 1, diagrammatical drawing of the fan like structure in the cleft apical segment of intermediate tarsi of Rhagoveliinae. — 2-6. R. blogiokommena paratypes, 2-4 male, 2 apex of abdomen, lateral view, 3 paramere, 4 hind femur; 5-6 female, 5 apex of abdomen, lateral view, 6 right half of thorax and abdomen in dorsal view. Scales 2,4-6 1 mm, 3 0.25 mm. —7-11. R. chrysomalla paratypes; 7-9 male, 7 apex of abdomen, lateral view, 8 paramere, 9 hind femur; 10-11 female, 10 ap- ex of abdomen, lateral view, ‘7’ and ‘8’ indicate 7th and 8th (gonocoxa) segment, 11 right half of thorax and abdomen in dor- sal view, ‘7’ and ‘8’ indicate tergites 7 and 8. Scales 7,9-11 1 mm, 8 0.25 mm. — 12-16. R. daktylophora paratypes; 12-14 male, 12 apex of abdomen, lateral view, 13 paramere, 14 hind femur; 15-16 female, 15 thorax and abdomen, lateral view, 16 right half of thorax and abdomen in dorsal view. Scales 12,14-16 1 mm, 13 0.25 mm. 260 more or less horizontal (fig. 11) in which cases the eighth tergite is the last visible in dorsal view. In other species the eighth tergite is pointing downward (fig. 34), in which case the seventh tergite is the last one fully visible in dorsal view. Another character that may give some trouble is the presence of minute den- ticles on the body, especially in greyish black species. Good optics with a magnification of at least 30 X and a good light source is needed to study this character. These denticles are more difficult to distinguish in wet than in dry specimens. They are often most dis- tinct on the proepisterna, which are the sclerites ante- rodorsally of the anterior coxae, on both sides of the rostrum. The species dealt with in this paper were mainly collected by J. P. Duffels (ZMA), N. Nieser (NC), J. van Tol (RMNH) and G. Zimmermann (NC/ZC) on several expeditions to Sulawesi, Buton and Sabah (Eastern Malaysia). Unless otherwise indicated speci- mens are deposited in the collections mentioned in brackets after the collectors. Some reference speci- mens have been sent to the additional collections mentioned in following list: Museum Zoologi Bogor, Bogor, Jawa (MBBJ); Bagian Pertanian, Universitas Haluoleo, Kendari, Sulawesi (BPUH); J. T. Polhemus collection (University of Colorado Museum), Englewood, Co. U. S. A. (PC); Nieser collection, Tiel, The Netherlands (NC); Chen collection, Beijing, P. R. China (PCHC); National Museum of Natural History, Leiden, The Netherlands (RMNH); G. Zimmerman collection, Marburg, B. R. D. (zc); Zoölogisch Museum, afd. Entomologie, Amsterdam, The Netherlands (ZMA) and Zoological Museum, Copenhagen, Denmark (ZMC). Some additional in- formation on localities can be found in Nieser & Chen (1991). ACKNOWLEDGEMENTS Thanks are due to Drs J. P. Duffels, J. van Tol and G. Zimmermann for putting specimens in their care at our disposal and to Dr. J. T. Polhemus for making paratypes of most of the species described by Polhemus & Polhemus (1988) available for study. SYSTEMATIC PART Key to Rhagovelia of Sulawesi (apterous specimens) 1. Pronotum short, its length much shorter than SMT DES ce ND aide os R. samarinda Polhemus & Polhemus [Described from E Kalimantan and Sarawak, one macropterous female has been reported from Lore Lindu N. P. If a species with short pro- notum is found, it may well represent a new spe- cies] 12, NIESER & CHEN, Rhagovelia of Sulawesi Pronotum long, covering, all or nearly all of mes- ONOMIME NE 2 Dorsum of pronotum and abdomen covered by striking appressed golden pubescence. Transverse yellowish-orange band anteriorly on pronotum separated from yellowish prosternum by variable and sometimes indistinct dark brown spots be- hind the eyes. [Large species, length 3.8-4.3 mm] CIR denke anne R. chrysomalla sp. n. Dorsum of pronotum and abdomen not covered by striking appressed golden pubescence .......... 3 . Transverse yellowish-orange band anteriorly on pronotum narrow, laterally at most reaching half- Waythelgyes®. onere na 4 Transverse yellowish-orange band anteriorly on pronotum wide, reaching laterally of eyes … … 20 Males wann cata Seber seh tue > Bemalessn cn PR na 12 Abdominal sternite 7 (sixth visible ventrally) not longitudinally carinate (fig. 56) nnen 6 Abdominal sternite 7 longitudinally carinate ....7 . Dorsum of abdomen hirsute; apex of paramere ENE (HE) een R. trichota Dorsum of abdomen with normal pubescence, not hirsute; apex of paramere pointed (fig. 75) ... Silicone POOR: MN RON en RER R. hamjadi [Only known by the type series from Lore Lindu N. P. Polhemus & Polhemus 1988: 194-195, figs. 134-140] Abdominal sternite 7 with a sharply pointed tuft of stiff setae, looking at first sight like a short stout peg, at basal third of longitudinal median carina (fig. 68). Hind femur inflated to a variable extend, at least twice as wide as middle femur ... 7 Don haa en Deer wiles rate tas antes à R. pruinosa Abdominal sternite 7 without peg like tuft of setae, hind femur 1.5 times or less as wide as mid- dlefemunka tee tn en 8 Abdominal sternites 4-6 with thick long golden yellowish pilosity (fig. 35)... 9 Abdominal sternites 4-6 with normal inconspi- cuous pubescence .......... R. kastanoparuphe sp. n. Abdominal tergite 7 about as long as its basal width Aten rh heh os een ite 10 Abdominal tergite 7 longer (about 1.4x) than its bald ee th eee 10 . Paramere apically broad, leaf-like (fig. 36, 43-44) ara ee aa R. pseudocelebensis sp. n. Paramere apically less broad (fig. 41-42) hte a NT, eretico mic. has agi R. celebensis . Sternites 5-7 with ventrolateral impressions resul- ting in a broad median keel longitudinally on the- se,sternites (tie PD) a R. daktylophora Ventrolateral impressions and keel restricted to sterne Ale R. sulawesiana Hind femur dorsoventrally flattened, lower mar- 261 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 17-29. — 17-20. R. horaia; 17-19 male, 17 apex of abdomen of holotype, lateral view, 18 paramere of paratype, 19 hind trochanter, femur and tibia of holotype; 20 female paratype, apex of abdomen, lateral view. Scales 17, 19, 20 1 mm, 18 0.25 mm. — 21-23. Right half of proctiger of males; 21 R. horaia paratype, scale 0.25 mm, 22 R. grayi, 23 R. lorelinduana, 22 & 23 drawn after J. & D. Polhemus (1988). — 24-29. R. kalami paratypes; 24-26 male, 24 apex of abdomen, lateral view, 25 pa- ramere, 26 hind trochanter, femur and tibia; 27-29 female, 27 hind trochanter, femur and tibia, 28 right half of thorax and abdomen, dorsal view, 29 apex of abdomen, lateral view. Scales 24, 26-29 1 mm, 25 0.25 mm. 262 13. 17e 18. 2D gin more or less straight (fig. 78) … R. hamjadi Hind femur not flattened dorsoventrally, with a convex lower margin (fig. 77) Connexiva margined with yellowish brown and set with long setae (longer than width of hind ti- bia) along entire length, converging caudally but Bon folded.over territe 7 .........e sen R. unica [Known only by the unique holotype female from Dumoga Bone N. P., Polhemus & Polhe- mus 1988: 195, figs. 43-45] Combination of characters not as above ......... 14 . Tergite 8 more or less horizontal, in the same pla- me as GEE nen ne R. pruinosa Abdominal tergite 8 distinctly directed ventrad... RE oe eh RF ont AIRE A A 15 . Caudal part of connexiva folded over abdominal tergite(s), at least covering most of tergite 7 ...16 Connexiva not folded over abdominal tergites . .. . Connexiva strongly constricted on segments 4 and 5, more or less parallel and nearly meeting on segments 6 and 7 (fig. 61). Tergite 7 with a me- diocaudal finger-like process ........ R. sulawesiana Connexiva constricted on segment 4, convergent caudally and meeting dorsally over posterior part of tergite 7 which lacks a mediocaudal finger-like PEO GCESSM LTE OO) Linn R. trichota Connexiva converging caudally in a straight or aid Sa NES. re 18 Connexiva distinctly constricted at the suture be- tween segment 6 and 7, tergite 7 with a distinct mediocaudal upwardly directed finger-like pro- CESS (asl OI) AES R. daktylophora Upper margin of connexiva brownish, their cau- dal apices long, tergite 7 with a more or less trian- gular hind margin (figs. 33, 34) o iena R. kastanoparuphe Upper margin of connexiva blackish, caudal api- ces short, hind margin of tergite 7 either truncate or with a mediocaudal finger-like projection (figs. EON des 19 . Hind margin of tergite 7 truncate, straight ......... PR e ne ee R. celebensis Hind margin of tergite 7 with a mediocaudal fin- ger-like projection . R. pseudocelebensis „Melles benede Ilio AT 21 Reese RSR er re 29 . Posterior tibia straight or softly curved, without larger subapical tooth in inner row (fig. 64) ....22 Posterior tibia curved, usually distinctly, with one or more larger teeth subapically in inner row (figs. 19, 26) Metanotum and first abdominal tergite orange-li- ke to light reddish brown. Posterior femur relati- vely slender, about 3.5 times as long as wide a ee I IRE SERRE à R. robina DI 24. 29: 26. 27. 28 29) 30. al. NIESER & CHEN, Rhagovelia of Sulawesi Metanotum and first abdominal tergite dark. Posterior femur more inflated, three times or slightly less as long as wide Sternite 7 laterally compressed, resulting in a strongly developed ventral keel, which is accentu- ated by a well-developed fringe of cilia (fig. 62) .. rradeina licia lett R. tropidata Sternite 7 ventrally flattened, with a low median keel Abdominal sternites 5-6 with postero-ventrally directed tufts of stiff setae (fig 71), paramere as in GINT GO RL EDER NOLI RIM DI R. wallacei Abdominal sternites 5-6 with normal pubescence only, without postero-ventrally directed tufts of stiff setae (fig 69), parameres as in figs. 3, 75 ...25 Minute black denticles present on pro- and meso- pleura, mesopleura without coarse punctation .... TOI Stia RE TEN STE R. minahasa Minute black denticles absent on pro- and meso- pleura, mesopleura with coarse punctation ......... MAL PTT AID ANNE 1 R. blogiokommena About 30 minute teeth in row proximal to large spine on posterior femur ........................ R. grayi [Only known by the type series from Sulawesi Utara, Lake Mala near Mokabang, Polhemus & Polhemus 1988: 207-208, figs. 208-215] About 20 minute teeth in row proximal to large spine on posterior femur Sternite 7 ventrally halfway with a tuft of caudal- lidirecedisemel(tip PA en ne R. kalami Sternite 7 ventrally halfway without a tuft of cau- dally directed setae Posterior femur nearly three times as long as wi- de, proctiger as in fig. 23 ............ R. lorelinduana [Only known by the type series from Lore Lindu N. P. in Sulawesi Tengah, Polhemus & Polhe- mus 1988: 208, figs. 141-148] Posterior femur about 2.5 times as long as wide, PIOGL PE HAS MAO E R. horaia Connexiva caudally folded over abdomen at least on tergite 7 Connexiva convergent posteriorly but not folded OE EHE 7 non e 2er 32 Minute black denticles distinctly and abundantly extending onto thoracic pleura. Connexiva strongly constricted on segment 5 and conver- ging in a more or less straight line caudally of constriction. Caudal apices of connexiva small, trlangulârd ster me Se RRs: ne R. minahasa Minute black denticles not distinctly extending onto thoracic pleura. Caudal apices of connexiva astenia on. 31 Connexiva only slightly constricted on segments 4/5, more or less straightly convergent caudally. Middle femur not dorsoventrally flattened ......... PR Re eee R. grayi 263 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 — Connexiva strongly constricted on segment 5, curved outward again on segment 6. Middle fe- mur dorsoventrally flattened ....... R. lorelinduana 32. Outer lateral margin of connexiva distinctly con- stricted on segment 4 (figs. 52, 67) ................ 33 — Outer lateral margin of connexiva more or less SUIS TE Ts. nen ee ke! 35 33. Meso- and meta-thoracic pleura yellow to pale orange-like, densely set with minute black denti- cles. Metanotum caudolaterally with small swel- lings (fiom EAR ae Mis Se he R. robina - Meso- and meta-thoracic pleura largely dark with at most a few minute black denticles. Metanotum without a pair of caudolateral swellings (fig. 67) 34. Length over 3 mm, connexiva swollen and slight- ly curved inward on segments 4-5 but not sinuate (fig. 67), mesopleuron without coarse punctation hace ee R. tropidata — Length just under 3 mm, connexiva slightly sinu- ate and hardly swollen on segments 4-5 (fig. 6), mesopleuron with coarse punctation .................. ee ern ae R. blogiokommena 35. Connexiva in dorsal view slightly thickened on segment 4. Abdominal tergite 7 only slightly lon- ger than! basall width... R. kalami — Connexiva not thickened on segment 4. Abdom- inal tergite 7 over 1.25 times as long as its basal Width a Me ns deo ea 36 36. Generally a dark grey brown species, yellow transverse band on pronotum on median line less than one fifth the median length of pronotum. Length of body about 3.2, ratio length to maxi- TALIA CIO ILO ee R. wallacei — Generally a reddish brown species, yellow trans- verse band on pronotum on median line more than one fifth the median length of pronotum. Length of body about 3.4, ratio length to maxi- muniwidthel74.. Re R. horaia Rhagovelia celebensis Polhemus & Polhemus (figs. 41-42, 81) Rhagovelia celebensis Polhemus & Polhemus, 1988: 199- 200, figs. 179-186, 229. Material. — SULAWESI UTARA: Dumoga Bone N.P., Base Camp, bridge, 16.x.1985 103d 729 (5d 19 macr.); same, flussabw. Brücke (downstream bridge), 22.x.1985, 2d 39; same, Base Camp, Stausee (‘barrage lake’), 22.x.1985, 6d 109 (12 macr.); same, Tumpah R.(type locality), Staustufe (barrage dam) 23.x.1985, 17¢ 189 (36 macr.); same, Tumpah R., quiet part, 28.x.1985, 34 39; Tumpah R. beach, 19.x.1985, 36d 369 (1d macr.); Tumpah, Water- fall oberh. Seitenbach (upstream tributary), 21.xi.1985, 1d 12; Südküste, Strasse, Bach, (South coast, road, stream), 18.xi.1985, 8d 139; all leg. G. Zimmermann; Dumoga Bone N. P. Tumpah river near confluence Toraut, UTM- 264 XL00631, 210 asl., 18.v.1985, 28 29, same, 23.v.1985, 106 49 (19 macr.); Dumoga Bone N. P., waterfall creek, tributary of Tumpah, UTM-XL0064, c. 225m asl., 24.v.1985, 29; same, hygropetric zone of Waterfall Creek, 22.iv.1985, 1d 19; Dumoga Bone N.P., Edward’s sub- camp, UTM-WL9365, 664m asl., 3.vi.1985, 59 (1 macr.); all leg. J. van Tol; 20 km E of Kotamobagu, Mt Ambang, alt. 1210 m, 8.xi.1985, multistr. evergreen forest, soil litter sifted, 19, leg. J. Krikken. — SULAWESI TENGAH, 50 km SE of Palu, Lore Lindu N. P., sur. Dongi Dong: shelter, 1°13’S 120°11’E, tributary of Sopu river, edge of multistr. ever- green forest, 950 m, 4.xii.1985-C, 16 19; Palu, Sopu river nr. Dongi Dongi, SJ 85, 950m, 5.xii.1985, 59; same, 8.xii.1985, 16, J. van Tol. — SULAWESI SELATAN, c. 10 km NW Palopo (km 15 along road Palopo-Rantepao), Salo Tandung, 2°58’S 120°07’E, 300-400 m asl., width 10m, boulders, torrents and seepage areas, open sec. forest, 27.iv.1991, 3d 109, leg. S. Kofman, 91JvT15; Mamasa, 2°56°S 119°22’E, 1050m, cultivated area, river flowing through village, rather fast flowing, bottom with boulders and some coarse sand, 8. IV. 1991, leg. J. van Tol, 26 29. Distribution. — This species is widespread in Sulawesi (fig. 81). Its type locality is Dumoga National Park, Tumpah River. Rhagovelia minahasa Polhemus & Polhemus (figs. 70, 73-74, 84) Rhagovelia minahasa Polhemus & Polhemus 1988: 205- 206, figs. 216-222, 228. Material. — SULAWESI UTARA: Lakes, Bach unter Sulphur spr., Z. Dorf, 21.xi.1985, 48d 459 (5d 69 macr.), many larvae; Dumoga Bone N. P., Tumpah River, 19. X. 1985, 19, leg. G. Zimmermann. Apterous, unless otherwise indi- cated. Distribution. — Only known from the eastern part of North Sulawesi (fig. 84). Rhagovelia pruinosa Polhemus & Polhemus (figs. 68, 77, 83) Rhagovelia pruinosa Polhemus & Polhemus, 1988: 195-196, figs. 149-157, 229. Material. - SULAWESI SELATAN: E of Maros, Bantimurung Area, Sg. Pattunuang Asue (type locality), 5°03’S 119°41’E, fast flowing stream, ponded areas, through disturbed forest on limestone, 23.iv.1991, J. van Tol, 68 82 (26 29 macr.). — SULAWESI TENGAH: Luwuk area, Sungai Batui, Singsing Camp, 19.x.1989, 18 macr.; Sungai Biak, 22.x. 1989, J. van Tol, 26. — Sulawesi Tenggara: Sungai Mowewe, 28.11.1989, N8922, 38; marsh S of Pomalaa, 2.iii.1989, N8930 36 119; 20 km E Kolaka, mountain stream, 3.iii.1989, N8934 1 6 , leg. N. Nieser; Konaweha R. near Sanggona, 7.xi.1989, J. van Tol, 48 109. — BUTON. Stream N of Baubau, 8.iii.1989, N8935 48 6%, leg. N. Nieser. Apterous unless otherwise indicated. Distribution. — Southern and eastern parts of Sulawesi, Buton (fig. 83). NIESER & CHEN, Rhagovelia of Sulawesi Table 1. Measurements of leg segments in Rhagovelia Tarsal segments 1 and 2 of fore and hind legs have not been measured for most species. When not given, they measure to- gether 0.05-0.07 and 0.12-0.15 respectively and do not seem to provide specific characters. femur tibia tars 1 tars2 tars3 R. blogiokommena 3 fore leg 0.80 0.82 = - 0.22 middle leg 1.30 1.05 0.06 0.40 0.62 hind leg 1.20 1.06 = = 0.23 R blogiokommena 2 fore leg 0.81 0.82 - = 0.22 middle leg 1.29 1.01 0.07 0.40 0.61 hind leg 1.11 1.10 = — 0.23 R. chrysomalla 3 fore leg 1.23 1.28 = = 0.30 middle leg 2.00 1.62 0.07 0.72 0.81 hind leg 1.44 1.72 = - 0.38 R. chrysomalla 2 fore leg 1.21 1.25 = = 0.29 middle leg 1.86 1.60 0.08 0.75 0.85 hind leg 1.41 1.73 — - 0.36 R. daktylophora 3 fore leg 0.93 1.01 = = 0.25 middle leg 1.58 1.26 0.07 0.57 0.71 hind leg 1.32 1.41 = = 0.30 R. daktylophora 9 fore leg 0.87 0.98 = = 0.27 middle leg 1.49 1.16 0.07 0.50 0.71 hind leg 122 1.39 = — 0.31 R. horaia 3 fore leg 1.04 1.09 = = 0.30 middle leg 1.60 1.37 0.08 0.49 0.69 hind leg 1.67 1.49 = — 0.40 R. horaia ® fore leg 1.00 1.03 = = 0.28 middle leg 1.55 1.26 0.007 0.47 0.70 hind leg 1.37 1.43 = = 0.40 R. kalami & fore leg 0.94 0.97 = = 0.22 middle leg 1.52 1.20 0.08 0.50 0.69 hind leg 1.39 1.20 = = 0.30 R. kalami ? fore leg 0.93 0.95 = = 0.22 middle leg 1.51 1.18 0.08 0.53 0.69 hind leg 1.31 1.23 = — 0.31 R. kastanoparuphe 3 fore leg 0.93 1.02 = = 0.26 middle leg 1.63 1.28 0.08 0.56 0.70 hind leg 1.47 1.39 — = 0.35 R. kastanoparuphe ? fore leg 0.95 1.03 = = 0.25 middle leg 1.58 1.23 0.08 0.54 0.70 hind leg 1.32 1.45 — = 0.32 265 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 e) À 32 30 37 39 2 Dean 45 46 Figs. 30-46. — 30-34. R. kastanoparuphe paratypes; 30-32 male, 30 apex of abdomen, lateral view, 31 paramere, 32 hind fe- mur; 33-34 female apex of abdomen, 33 dorsal view ‘7’ and ‘8’ abdominal tergites 7 and 8 respectively; 34 lateral view. Scales 30, 32-34 1 mm, 31 0.25 mm. — 35-40. R. pseudocelebensis paratypes; 35-37 male, 35 apex of abdomen, lateral view, 36 pa- ramere, 37 hind leg; 38-40 female, 38 hind femur, 39 apex of abdomen lateral view, 40 left half of abdomen, dorsal view. Scales 35, 37-40 1 mm, 36 0.25 mm. — 41-46, parameres of 41-42, R. celebensis (Dumoga Bone N. P.), 43-44, R. pseudoce- lebensis (Kolaka and Buton respectively), and 45-46, R. sulawesiana (paratype); 45, lateral view, 46, medial view. Scale 0.25 mm. 266 R pseudocelebensis 3 fore leg 0.96 0.99 middle leg 1.48 1.17 hind leg 1.26 1.35 R. pseudocelebensis 9 fore leg 1.00 0.95 middle leg 1.41 1.11 hind leg 1.23 1.35 R. robina 3 fore leg 0.91 0.93 middle leg 1.47 1.11 hind leg 1.37 1.14 R robina 9 fore leg 0.90 0.93 middle leg 1.44 1.08 hind leg 1.29 1.28 R. trichota & fore leg 0.89 0.91 middle leg 1.41 1.13 hind leg 1.20 1.24 R. trichota 2 fore leg 0.96 0.96 middle leg 1.43 1.1177 hind leg 1.22 1.31 R.tropidata 3 fore leg 0.92 0.96 middle leg 1.52 1.23 hind leg 1.40 1.22 R.tropidata 9 fore leg 0.90 0.93 middle leg 1.42 1.11 hind leg 1.23 1.23 NIESER & CHEN, Rhagovelia of Sulawesi = - 0.28 0.07 0.53 0.70 = — 0.31 = = 0.23 0.08 0.50 0.69 = — 0.29 En = 0.24 0.08 0.52 0.66 - — 0.30 = = 0.23 0.08 0.51 0.66 - — 0.31 — — 0.20 0.06 0.43 0.65 — — 0.30 = — 0.22 0.08 0.47 0.70 - — 0.32 — - 0.22 0.09 0.48 0.68 — — 0.32 — — 0.22 0.08 0.41 0.64 = — 0.36 Rhagovelia sulawesiana Polhemus & Polhemus (figs. 61, 69, 81) Rhagovelia sulawesiana Polhemus & Polhemus, 1988: 198- 199, figs. 158-164, 230. Material. — SULAWESI SELATAN: E of Maros, Bantimurung Area, Sg. Pattunuang Asue [type locality], 5°03’S 119°41’E, fast flowing stream, ponded areas, through disturbed forest on limestone, 23.iv.1991, J. van Tol, 38 59; SE of Malino, rivulet at N side of Lompobattang, cultivated area, 20.vi.1985-A, J. van Tol 26 . All apterous. Distribution. — S Sulawesi (fig. 81). Rhagovelia wallacei Polhemus & Polhemus (figs. 71, 76, 82) Rhagovelia wallacei Polhemus & Polhemus, 1988: 203-205, figs. 187-193, 230. Material. — SULAWESI UTARA: Kotamobagu, Caves, 26.x.1985, 116 72; Dumoga Bone N. P., Base Camp, Toraut R., maze, 7.xi.1985, 78 159; Tumpah R., beach, 19.x.1985, Id 42 (12 macr.); Tumpah R., Waterfall oberh. Seitenbach, 21.xi.1985, 2d ; Malibagu Z., 8.xi.1985, 12 macr.; Lakes, Kleine See, Abfluss des Flusses (small lake, drainage of stream), 16.xi.1985, 6d 119 (22 macr.); Lakes, kleiner See, Fischteich, 16.xi.1985, 26 49; Südküste, Strasse, Bach, 18.xi.1985, 28 99; all leg. G. Zimmermann; Dumoga Bone N. P., waterfall creek (type locality), 24.v.1985, 16; Tumpah river near confluence Toraut, UTM-XL00631, 210 asl., 21.v.1985, 1d, both J. van Tol; 20 km E of Kotamobagu, Mt Ambang, alt. 1210 m, 8.xi.1985, multistr. evergreen forest, soil litter sifted, 1d 19, leg. J. Krikken. — SULAWESI TENGAH, 50 km SE of: Lore Lindu N. P., sur. Dongi Dongi shelter, 1°13’S 120°11’E, tributary of Sopu river, edge of multistr. evergreen forest, 950 m, 4.xii.1985-C, 19. Apterous, unless otherwise indi- cated. Distribution. — Northern and eastern parts of Sulawesi (fig. 82) 267 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Rhagovelia blogiokommena sp. n. (figs. 2-6, 82) Type material. — Holotype d : Sulawesi Tenggara, moun- tain stream between Tamborasi and Wolo, 1 mar. 1989, leg. N. Nieser, N8926 (ZMA). — Paratypes, same data as holoty- pe, 7d 99 (19 allotype ZMA, 36 39 NC, 1d 19 PCHC, 1d 19 Jrrc, 1d 19 MBBJ, 1d 19 RMNH, 1% ZMC). All ap- terous (fig. 82). Description. — Apterous form. Dimensions. Length d 2.80-2.92, 9 2.85-3.00; width d 1.0-1.1, 2 1.1-1.2; width of head d 0.70-0.73, £ 0.72-0.74; width of pronotum d 0.90-0.95, 2 0.95-1.02. Colour, generally dark grey to blackish, anterior quarter of pronotum orange-yellow, connected with yellowish propleura, and prosternum. Dorsal half (males) to two thirds (females) of connexivum oran- ge. Venter dark, juga, prosternum and abdominal sternite 7 yellowish, genital segments of male yello- wish. Basal half of first antennal segment, basal part of rostrum, basal three quarters of fore femur, basal third of hind femur, acetabula, coxae and trochanters pale; hind femur in ventral view medium brown. Minute black denticles virtually restricted to poste- rior half of jugum of head and propleura on both sides of rostrum. Mesopleuron in front of acetabulum with a group of 11-16 coarse punctures. Dorsum rather bare, abdominal tergites pruinose laterally. Vertex, interoculus, sides of thorax and abdomen, an- tennae and legs with the usual pubescence and setae. Punctures on disk of pronotum indistinct except for a row at posterior margin of transverse orange-yellow part. Length of antennal segments I-IV d 0.70 : 0.37 : 0.47 : 0.41, 2 0.68 : 0.35 : 0.45 : 0.41. Pronotum long, covering mesonotum, length: width 0.75 : 0.92. Length of metanotum on midline, 0.10. Abdominal tergites 1-5 subequal in length (0.15), tergite 6 slight- ly longer (0.20), tergite 7 0.30 in male, 0.35 in fema- le. Male. Posterior trochanter with 1-3 small but dis- tinct blunt teeth. Posterior femur slightly less than four times as long as wide (1.20/0.32), on ventral margin proximally with about 10 minute and closely set teeth, followed about halfway by a large spine about as long as half the width of femur, with a row of about 11 spines of decreasing length distally (fig. 4). Anteroventrally a row of 5 smaller spines located in distal part. Larger spines black tipped, smaller spi- nes and teeth entirely black or nearly so. Posterior ti- bia virtually straight, armed beneath with a row of about 15 rather coarse pegs, a few proximally paired. Connexiva more or less horizontal, gradually conver- ging posteriad. Basal width of abdominal tergite 7 subequal to its median length. Abdominal sternite 7 with a pair of shallow impressions separated by a nar- 268 row and low keel, sternite 8 with an indistinct ventral keel (fig. 2). Parameres as in fig. 3. Female. Posterior trochanter without small teeth. Posterior femur slightly less than four times as long as wide (1.1/0.3), row of minute teeth absent, row of larger spines essentially as in male. Posterior tibia with small and less distinct pegs as in male. Connexiva mo- re or less vertical, on tergites 5 and 6 somewhat sinu- ate, otherwise gradually converging caudally, with the pointed apices of connexiva touching, or nearly so (fig. 6). One female has the connexiva nearly hori- zontal. Tergite 8 slanting ventrad at an angle of about 45°, proctiger pointing ventrad. Sternite 7 about half as long as remaining abdominal sternites (0.5/1.0). Gonocoxa hidden in segment 7. Etymology. — Blogiokommenos, greek adjective meaning pockmarked, referring to the coarse puncta- tion on mesopleura. Comparative notes. — Similar to R. tropidata, R. wallacei and, in males, R. minahasa, all of which have distinct minute black denticles on pro- and meso- pleura. À. minahasa lacks mesothoracic punctures, À. tropidata has fewer (4-9) and indistinct punctures in front of mesoacetabula, R. wallacei has slightly less (about 10) and less coarse punctures there. Male ster- nite 7 and female connexiva are also different (figs. 2, 62, 70-71, 6, 67) Rhagovelia chrysomalla sp. n. (figs. 7-11, 83) Type material. — Holotype d : Sulawesi Tengah, 60 km SE of Palu: Lore Lindu N. P., UTM-SJ95, Danau Tambing and brooklets, 1600 m asl., Pandanus, 7 Dec. 1985-A, leg. J. van Tol (RMNH). — Paratypes 48 89, same data as holotype (23 39, including allotype RMNH, 26 39 NC, 19 MBB}, 12 yrpc). All apterous (fig. 83). Description. — Apterous form. Dimensions. Length d 3.80-3.90, 9 4.10-4.33; width d 1.3-1.4, ? 1.4-1.6; width of head d 0.88-0.90, 9 0.90-0.92; width of pronotum d 1.20-1.28, 9 1.22-1.40. Colour, dark grey brown to blackish, eyes castane- ous. Anterior fifth of pronotum orange-yellow. Propleura, pro- and mesosterna, first antennal seg- ment except apex, acetabula, coxae and trochanters anterior femur except apex yellowish. Middle and hind femur variably yellowish and brown, connexiva entirely in female, distal two thirds in male orange yellow. Sternites 7 and 8 in male, tergite 8, proctiger and gonocoxa in female yellowish. The orange-yellow anterior transverse band of pronotum tends to be se- parated from the yellow area of the propleura by a dark spot behind the eyes which is quite variable and may be absent. Thoracic venter with minute black denticles, ex- Nieser & CHEN, Rhagovelia of Sulawesi Figs. 47-61. — 47-55. R. robina paratypes; 47 apex of abdomen, lateral view, 48 paramere, 49 hind trochanter and femur; 50- 52 female, 50 hind femur, 51 dorsal part of thorax and abdomen lateral view, 52 left half of thorax and abdomen, dorsal view; 53-55 hemielytra of males (54-55 of same specimen). Scales 47, 49-55 1mm, 48 0.25 mm. — 56-60. R. trichota paratypes; 56- 58 male, 56 apex of abdomen lateral view, 57 paramere, 58 hind trochanter, femur and tibia; 59-60 female, 59 apex of ab- domen, lateral view, 60 right half of thorax and abdomen, dorsal view. Scales 56, 58-60 1 mm, 57 0.25 mm. - 61, R sula- wesiana paratype female, right half of thorax and abdomen, dorsal view. Scale 1 mm. 269 TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 tending onto pleura, most distinctly on jugum and proepisternum. Interoculus, thoracic and abdominal tergites clothed with dense, appressed golden-yellow pilosity. (This golden pilosity is also present in larva V). In addition dorsum of thorax and abdomen with sparse long erect dark cilia. Vertex sides of thorax an- tennae and legs with the usual pubescence and setae. Length of antennal segments I-IV d 0.95 : 0.44 : 0.69 : 0.67, 2 0.99 : 0.47 : 0.68 : 0.59. Pronotum long, covering mesonotum. Disk of pronotum with some indistinct punctures, obscured by pilosity. Length: width of pronotum male 0.93 : 1.22, female 1.18 : 1.32. Length of metanotum on midline, 0.20. Abdominal tergites 1-5 subequal in length (0.20 in male, 0.25 in female), tergite 6 subequal in male, slightly longer (0.30) in female, tergite 7 distinctly longer (0.43 in male, 0.50 in female). Male. Posterior trochanter length: width 0.40 : 0.21, without teeth or warts. Posterior femur slender, slightly less than four and a half times as long as wide (1.44/0.33), on ventroposterior margin about half- way with a rather small brown spine, its length so- mewhat less than one thirds the width of femur, with a row of about 7 spines of decreasing length distally (fig. 9). Posterior tibia straight, armed beneath with a row of about 20 small teeth. Connexiva slanting up- ward about 45°, convex on segments 3-5 strongly converging posteriad on segment 7, no caudal points. Basal width of abdominal tergite 7 1.5 times its me- dian length (0.6/0.4). Abdominal sternite 7 compara- tively short, about 1.5 times as long as tergite 6 (0.3/0.2), somewhat flattened medially, without a ca- rina. Genital segments prominent (fig. 7), fusiform, tergite 8 slightly longer than tergite 7 (0.45/0.40). Sternite 8 laterally compressed at base, resulting in a low blunt carina. Median length of pygophore dis- tinctly longer than median length of sternite 8 (0.4/0.3). Parameres as in fig. 8. Female. Posterior femur about five times as long as wide (1.4/0.23). Posterior femur and tibia without teeth or spines. Connexiva vertical, sinuate, slightly curved inward on segments 4-5 and curved outward on segments 6-7, in some specimens nearly straight (fig. 11). In lateral view connexiva slightly convex dorsally on segments 4-6, caudal apex truncate with a tuft of pilosity at the tip (fig. 10). Basal width of ter- gite 7 slightly larger than its median length (0.55/0.50). Tergite 8 nearly horizontal, with caudo- lateral tufts of bristles. Sternite 7 large, about one third as long as the remaining abdominal sternites to- gether (0.6/1.8). Gonocoxa clearly visible, about half the length of sternite 7, ventro-laterally slightly com- pressed. Proctiger distinctly visible. Etymology. — Chrysomallos, greek adjective mean- ing ‘with a golden fleece’, refers to the golden pubes- 270 cence on dorsum of this species. Comparative notes. — The size combined with stri- king golden pubescence dorsally set this species apart from its Sulawesi congeners. Rhagovelia daktylophora sp. n. (figs. 12-16, 79, 81) Type material. — Holotype d: Sulawesi Tengah, SW Luwuk, Toptop Camp along Batui River, 1°09'S, 122°31’30”E, 120 m, lowland rainforest, 19-21 Oct 1989, leg. J. P. Duffels, sample Sul. 18 (ZMA). — Paratypes (inclu- ding allotype 9) same data as holotype 146 72 (zmA, 2d 29 NC, 1d 19 MBBJ); Batui River at Singsing Camp, SW of Luwuk, c. 1°09’S 122°31’E, 90m, middle sized stream in lowland rainforest, 14-17 Oct 1989, sample Sul.14, J. P.Duffels, 38 (16 RMNH, 26 ZMA); NE Luwuk, Sungai Bantayan, near road Kayutanyo to Siuna, 0°47’S 123°00’E, 50-100 m, medium sized river in slightly disturbed lowland rainforest, 7.x.1989, leg. J. P. Duffels, sample Sul. 2, 106 30/2 (ZA 37/2 NI reg SPC aloe) = Luwuk area, Sg. Batui and tributaries at Singsing Camp, 17.x.1989, J. van Tol 236 119 (48 22 macr., RMNH, 1d apt., 2ó 12 maer. Ne, 26 22 apt. jrrG Lon lie rapt SNOW); Luwuk area, Sg. Biak, 21.x.1989, leg. J. van Tol, 8d 92 (12 macr.). Apterous unless otherwise indicated (fig. 81). Description. — Apterous form. Dimensions. Length d 3.04-3.25, 2 3.05-3.28; width d 1.2-1.3, 2 1.3-1.4; width of head d 0.80-0.84, © 0.80-0.83; width of pronotum d 1.02-1.13, 9 1.10-1.20. Colour, black, pronotum with anterior transverse orange band usually slightly narrower than interocu- lar space. Basal third of first antennal segment, aceta- bula, fore and hind coxae and trochanters and basal part of anterior femur ventrally pale yellowish. In some specimens middle coxa medium to light brown, but in most blackish. Venter largely pruinose ash grey. Venter with minute black denticles, which are dif- ficult to see due to pruinose dark grey background, extending onto episterna. Dorsum with a few short golden-yellow setae, longer and denser on genital seg- ments of male. Vertex sides of thorax antennae and legs with the usual pubescence and setae. Length of antennal segments I-IV d 0.80 : 0.40: 0.44 : 0.43, 2 0.80 : 0.39 : 0.43 : 0.42. Pronotum long, covering mesonotum, usually laterocaudal ang- les of mesonotum narrowly visible. Disk of pronotum with comparatively fine punctures, about 30/0.25mm. Length: width of pronotum 0.85 : 1.10. Length of metanotum on midline, 0.15. Abdominal tergites 1-5 subequal in length (0.12 in male, 0.15 in female), tergite 6 slightly longer (0.20 in male, 0.30 in female), tergite 7 distinctly longer (0.40). Male. Posterior trochanter length: width 0.32: 0.18, pilose but without teeth or warts. Posterior fe- mur slender, nearly five times as long as wide (1.4/0.3), on ventroposterior margin about halfway with a large black spine its length two thirds the width of femur, with a row of 6 black spines of de- creasing length distally and three very small spines proximally of large spine (fig. 14). Posterior tibia straight, armed beneath with a row of about 16 small teeth, proximally three or four extra teeth forming pairs. Connexiva more or less horizontal gradually converging posteriad, no caudal points. Basal width of abdominal tergite 7 equal to its median length. Abdominal venter with long, quite dense, golden yel- low setae on sternites 3-6 and the base of sternite 7. Sternite 7 in lateral view distinctly compressed dorso- ventrally in posterior three fourths. Apical part of sternite 7 without pubescence, shining black, with a pair of sublateral impressions, separated by a low, nar- row median carina (fig. 12). Segment 8 prominent, elongate, in dorsal view, length 0.55, basal width 0.37, sternite 8 shining black. Parameres relatively large, as in fig. 13. Female. Generally with a more compact build as compared to male, due to compressed genital seg- ments. Posterior femur nearly five times as long as wi- de (1.18/0.27). Spines on posterior femur and tibia essentially as in male. Connexiva vertical, strongly curved inward on segment 5-6; ending in narrow cau- dal points with a tuft of pilosity at the tip (fig. 16). Another tuft of bristles just ventral of apices of con- nexiva and a smaller group of somewhat more scatte- red bristles on dorsal margin of connexiva at segment 6, best visible in lateral view (Fig. 15). Tergite 7 with a well developed mediocaudal projection on hind margin, with a short tuft of setae apically. Tergite 8 nearly vertical, genital segments retracted into seg- ment 7. Sternite 7 large, shining black under quite dense ash white pubescence, slightly over half as long as the remaining abdominal sternites together (0.67/1.2), sides of segment 7 pruinose grey. Macropterous form. — Mostly as apterous except for development of pronotum and hemielytra. Length d 3.5-3.7, 2 3.7-3.8; humeral width of pro- notum of both sexes 1.37-1.45. Submedial carinae at base of abdomen just not reaching the hind margin of first abdominal tergite. Female without the finger-li- ke projection on caudal margin of tergite 7 and the constriction of connexiva on segments 5-6 less pro- nounced. Etymology. — Daktylophora, Greek adjective mean- ing ‘bearing a finger’ refers to the finger-like projec- tion on tergite 7 in apterous female. Comparative notes. — Very similar to À. celebensis and À. pseudocelebensis, but the connexiva of apterous females curved inward on segments 5-6 in combina- NieseR & CHEN, Rhagovelia of Sulawesi tion with a well developed finger-like projection on the caudal margin of abdominal tergite 7 (figs. 15-16, 39-40) and the parameres of males, with the broad apical part longer (figs. 13, 36, 41-44), are distinctive. Rhagovelia horaia sp. n. (figs. 17-20, 84) Type material. — Holotype d: Sulawesi Tenggara, Centipede Camp, 3.xi.1989, J. van Tol (RMNH). — Paratypes 23 69 (including allotype), same data as holotype (RMNH, 16 22 NC, 12 MBBJ); S. Mokowu, 5.xi.1989, J. van Tol, 19, all apterous (fig. 84). Description. — Apterous form. Dimensions. Length d 3.4-3.7, 9 3.5-3.8, width of head d 0.78- 0.81, © 0.78-0.83; width of pronotum ¢ 1.07-1.12, ® 1.08-1.12; thoracic width d 1.12-1.23, 2 1.20- 125: Colour, reddish brown, eyes dark castaneous to blackish, anterior quarter of pronotum and entire propleura pale yellow. Basal two thirds of first anten- nal segment, basal two thirds of fore femur, coxae and trochanters, basal half of hind femur especially in fe- male, distal margin of connexiva more distinctly dor- sally than ventrally, pale. Venter with minute black denticles extending onto thoracic pleura, distinctly on propleura only a few on meso- and metapleura. Dorsum with normal pubes- cence and bristles, rather dense yellowish setae on ge- nital segments of male. Vertex, head, sides of thorax and apex of abdomen, antennae and legs with the usual pubescence and setae. Length of antennal segments I-IV d 0.81 : 0.40: 0.49 : 0.46, 2 0.80: 0.40: 0.47: 0.43. Pronotum long, covering mesonotum, length: width of prono- tum 0.85 : 1.1, length of metanotum on midline d 2 0.12. Posterior three quarters of disk with coarse punctures, about 50/0.25 mm’. Abdominal tergites 1-6 in male and 1-5 in female, subequal in length (0.20-0.22), tergite 7 in male longer (0.42), tergite 6 in female slightly longer (0.30), tergite 7 distinctly so (0.55). Male. Posterior trochanter length : width 0.40 : 0.25, with 7-8 well developed teeth. Posterior femur strongly inflated, about two and a half times as long as wide (1.73/0.78 to 1.57/0.60), on posteroventral margin proximally a row of about 15 minute teeth, about halfway a large spine about a quarter as long as the width of femur, with a row of 7 spines of decre- asing length distally. Anterior to the distal row a row of 4-5 smaller teeth. Posterior tibia curved (fig. 19), armed on inner side with a proximal row of about 13 teeth, then a larger tooth, followed by four distal teeth. In addition a more posterior row of about 12 smaller teeth. Connexiva horizontal slightly convex 271 TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 62-80. — 62-67, R. tropidata paratypes; 62-64 male, 62 apex of abdomen, lateral view, 63 paramere, 64 hind leg; 65-67 female, 65 hind leg; 66 abdomen in lateral view, 67 left half of thorax and abdomen dorsal view. Scales 62, 64-67 1 mm, 63 0.25 mm. — 68-73, Apex of abdomen, lateral view. 68, R. pruinosa male, 69 R. sulawesiana male, 70 R. minahasa male, 71 R. wallacei male, 72 R. grayi female, 73 R. minahasa female. Scale 1 mm. — 74-76. Paramere (75-76 redrawn after Polhemus & Polhemus 1988); 74 R. minahasa, 75 R. hamjadi, 76 R. wallacei. Scale 0.25 mm. — 77-78, hind femur of female;77 R. prui- nosa, 78 R. hamjadi (redrawn after Polhemus & Polhemus 1988); 79-80. Right hemielytron; 79 R. daktylophora, 80 R. pseu- docelebensis. Scale 1 mm. 272 on segment 4-6, otherwise gradually converging to- ward tergite 7, without apical points. Basal width of abdominal tergite 7 subequal to its median length (0.4), centrally more or less bare and somewhat shi- ning. Abdominal sternite 2 with a well defined me- dian keel, sternite 7 flattened medially, nearly bare, shining, without median keel, its median length one third of median length of remaining sternites together (fig. 17). Sternites 8 and 9 of subequal length, some- what shorter (0.25/0.35) than sternite 7. Paramere as in fig. 18. Female. Posterior trochanter with three or four small teeth. Posterior femur four and a half times as long as wide (1.35/0.31). Posterior femur on ventral margin proximally with 0-3 small minute teeth, large spine halfway three fifth the width of femur, followed by 4-6 smaller spines, hind tibia straight, with a, dis- tally double, row of about 18 teeth on inner surface. Connexiva slanting 45° upwards, slightly convex on segments 3-5, converging caudally with short, nar- rowly triangular, caudal points. Abdominal tergite 7 distinctly longer than its basal width (0.55/0.30). Tergite 8 slanting 45° downward, proctiger visible in dorsal view. Abdominal sternite 7 half as long as pre- ceding ones together, bare, shining, no minute black denticles, gonocoxa short but well visible (fig. 20). Macropterous form, unknown. Etymology. — Horaios, a greek adverb meaning e. g. youthfully beautiful. Comparative notes. — The male is nearly identical to À. grayi Polhemus & Polhemus, except for the proctiger which lacks a pair of proximal tubercles (figs. 21, 22). Females of R. grayi have connexiva up- turned, expanded and incurvate along tergites 4 and 5 and apically folded over abdomen, whereas in R. ho- raia the connexiva of female are simple. Another si- milar species is R. lorelinduana which has in male a quite different proctiger (figs. 21, 23) and in female curved and folded connexiva much as in R. grayi. Rhagovelia kalami sp. n. (figs. 24-29, 82) Type material. - Holotype d : Pulau Buton, first stream along road from Baubau to the North, 8 Mar. 1989, N8935, leg. N. Nieser (RMNH). — Paratypes, same data as holotype, 66 172 (29 including allotype RMNH, ZMA 16 22, zme lé 29, rest NC); Buton, small stream near sea, 9.iii.1989, N8941, 146 182 (46 42 PCHC, 16 1 BPUH, 18 19 MBB), 28 32 Jrrc, 1d 12 Zc); Buton, small stream 16 km E Baubau, 10.iii.1989, N8944, 66 99. All apterous (fig. 82). Description. — Apterous form. Dimensions. Length d 3.08-3.28, 2 3.20-3.50; width d 1.15- 1.19, £ 1.30-1.42; width of head d 0.72-0.80, 9 NIESER & CHEN, Rhagovelia of Sulawesi 0.78-0.80; width of pronotum d 0.98-1.08, 2 1.10- 1.18. Colour, dark greyish. Eyes dark castaneous to black, interoculus black, pronotum and dorsum of abdomen largely black with a greyish pruinose bloom. Anterior quarter of pronotum, propleura and prosternum yellowish orange. Connexiva light orange brown, proximally blackish. First antennal segment, acetabula, coxae trochanters, anterior femur except for distal apex, base of posterior surface and most of anterior surface of hind femur, variable part of anteri- or surface of middle femur, mesosternum and medio- caudal part of abdominal venter including genital seg- ments pale yellow orange. Venter with minute black denticles which extend onto jugum, proepisternum and dorsal sides of acetabula. Dorsum of abdomen of male with golden-yellow setae. Vertex sides of thorax antennae and legs with the usual pubescence and setae, except for hind tibia of male which is more hai- ry than in most other species. Length of antennal segments I-IV d 0.78 : 0.39 : 0.47 : 0.44, ® 0.77 : 0.38 : 0.47 : 0.42. Pronotum long, covering mesonotum, in some specimens latero- caudal angles of mesonotum narrowly visible. Disk of pronotum with coarse punctures, about 40/0.25mm’. Length : width of pronotum 0.80 : 1.05. Length of metanotum on midline, 0.12. Abdominal tergites 1-6 in male and 1-5 in female subequal in length (0.2) Male. Posterior trochanter length: width 0.3: 0.2, with 5-6 small teeth of which 2 twice as large as the others. Posterior femur strongly inflated, less than three times as long as wide (1.4/0.5), on ventral mar- gin distally with about 20 minute teeth, about half- way a large spine its length about one fourth the width of femur, with a double row of 6-7 spines of decreasing length distally, the larger spines pale, with blackish tip. Posterior tibia slightly curved armed beneath with a row of about 17 short blunt teeth of which the apical one and the fourth or fifth from apex are larger (fig. 26), anterior to row a few extra teeth. Connexiva more or less horizontal, distal margin straight to slightly convex on segments 4-6, conver- ging posteriad, no caudal points. Basal width of ab- dominal tergite 7 three quarters its median length (0.32/0.40). Sternite 7 in lateral view laterally com- pressed in posterior three fourths, with a low median carina marked at its base by a small but dense tuft of setae resembling a spine (fig. 24). Genital segments hirsute, tergite 8 as long as wide (0.32/0.33). Parameres as in fig. 25. Female. Posterior femur four times as long as wide (1.3/0.32). Spines on posterior femur essentially as in male. Posterior tibia straight, armed beneath with a row of about 17 short blunt teeth all of subequal size. Connexiva slanting about 45° upwards, distal margin slightly sinuate, thickened on segment 4, where there 275 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 is a patch of short dense pubescence. Apex of con- nexiva triangularly produced caudally, accentuated by a tuft of bristles (fig. 29). Tergite 7 slightly longer than its basal width (0.40/0.35). Tergite 8 nearly ho- rizontal, with laterocaudal tufts of pilosity, basal width equal to its median length (0.32), apical mar- gin truncate. Sternite 7 large, shining, half as long as the remaining abdominal sternites together (0.6/1.2). Gonocoxa not distinctly visible. Macropterous form, unknown. Etymology. — Kalami, greek noun, meaning shin, referring to the modified tibia of male. Comparative notes. — À. kalami is not very similar to other species in the region, those with a similar male tibia (R. grayi, R. horaia) are more slender and longer. R. tropidata and R. wallacei are of similar sha- pe, but have the male hind leg less modified. See also key to species. Rhagovelia kastanoparuphe sp. n. (figs. 30-34, 84) Type material. — Holotype & (ZMA): Sulawesi Tenggara, NE Kolaka, Mokowu Camp, along Mokowu River, nr. Gng. Watuwila, c. 3°49°S 121°40’E, 200m, 29-31 Oct. 1989, disturbed lowland rainforest, J. P. Duffels, sample Sulawesi 24 (ZMA). — Paratypes: same data as holotype, 14¢ 269 (78 189 including allotype 9 ZMA, 16 19 BPUH, 19 JTPC, 16 19 MBBJ, 36 32 NC, 16 19 PCHC, 16 19 RMNH); Sungai Mokowu, 30.x.1989, J. van Tol, 29; Rd. Kendari-Lalimboe, 21.ii.1989, N8903, 22; tributary Sg. Mowewe, 28.11.1989, N8923, 18 49. — P. Buton, stream N of Baubau, 8.iii.1989, N8935, 106 199 (68 109 NC, 16 19 BPUH, 16 39 yrec, 16 29 PCHC, 22 RMNH, 29 ZMA, 16 19 ZMC); P. Buton, Sg. Labuhan Tobelo at Jismil camp, 18.xi.1989, leg. J. van Tol 968 329 (19 macr., 5d 249, incl. 1 macr., RMNH, 48 89 NC). Apterous, unless otherwise indicated (fig. 84). Description. — Apterous form. Dimensions. Length d 3.0-3.3, 2 3.0-3.2; width d 1.0-1.1, © 1.1-1.2; width of head 6 0.79-0.82, 2 0.82-0.83; width of pronotum d 0.92-1.01, 9 1.0-1.08. Colour, black with a greyish tinge. Orange trans- verse band anteriorly on pronotum only slightly wi- der than interoculus. Propleurae orange, separated from pronotal transverse band by a narrow black mark behind eyes. Distal margin of connexiva orange brown, more distinctly so in females than in males, without obvious pilosity. Basal half of first antennal segment, acetabula, coxae, trochanters, basal half of fore femur, basal quarter of hind femur pale yellow. Rostrum castaneous except for darker apex. Abdo- minal sternites 7 and 8 in male and sternite 7 in fe- male, castaneous. Venter with minute black denticles, which do not extend onto pleura except for a few on metapleuron. 274 Dorsum mostly with a greyish pruinose cover, sparse pilosity except for some shiny yellowish setae on geni- tal segments of male and a tuft of long dark setae at caudal apex of connexiva. Vertex anteriorly with short dark setae, sparse long dark setae on head, sides of thorax and apex of abdomen. Antennae and legs with the usual pubescence and setae. Length of antennal segments I-IV d 0.81 : 0.40: 0.49 : 0.46, 2 0.80 : 0.40 : 0.47 : 0.43. Pronotum long, covering mesonotum. Length: width of prono- tum 0.85: 1.4. Disk of pronotum with about 60 punctures/mm’. Length of metanotum on midline, 0.12. Abdominal tergites 1-5 subequal in length (0.14), tergite 6 slightly longer (0.25), tergite 7 dis- tinctly longer (0.45). Male. Posterior trochanter length: width 0.30: 0.18, without teeth or warts. Posterior femur rather slender, four and a half times as long as wide (1.3/0.28), on ventral margin about halfway with a large spine about 0.6 as long as the width of femur, with a row of 5-7 spines of decreasing length distally (fig. 32). Posterior tibia straight, armed beneath with small blunt teeth proximally shorter and more dense- ly placed. Connexiva slanting up- and outward, about 45°, gradually tapering downward on tergite 7, with- out apical points. Basal width of abdominal tergite 7 0.30, its median length 0.45; sternites 6-8 with a well defined median keel. Segment 8 prominent (fig. 30), elongate, in dorsal view, length 0.4 width 0.30. Parameres as in fig. 31. Female. Generally with a more compact build as compared to male, due to compressed genital seg- ments. Posterior femur five times as long as wide (1.3/0.27). Length of large spine on posterior femur three quarters the width of femur, followed by 3-5 smaller spines, hind tibia essentially as in male. Connexiva variable more or less horizontal to vertical, dorsal margin straight to very slightly convex, apex of connexiva narrowly triangularly produced dorsocau- dally, accentuated by a tuft of setae (fig. 34). Posterior margin of tergite 7 bluntly triangular with a short me- diocaudal tuft of setae, often suggesting a spine (fig. 33). Tergite 8 nearly vertical, gonocoxa mostly hid- den within segment 7. Sternite 7 large, shining, half as long as the remaining abdominal sternites together. Macropterous form. The single macropterous fe- male, which has its wings torn off, is identical to the apterous form except for stronger development of pronotum, humeral width 1.40, and for stronger de- velopment of pronotum, humeral width 1.40, and absence of the mediocaudal tuft of setae on abdomi- nal tergite 7. Etymology. — Kastanoparuphe, greek noun mean- ing brown band or brown rim, referring to the upper margin of connexiva. Comparative notes. — At first sight similar to À. ce- lebensis and related forms, the lack of long pilosity on abdominal venter in male and the brownish upper rim of connexiva in female separate this species easily. The paramere (fig. 31) and the apex of abdomen of female (fig. 33, 34) are also characteristic. Rhagovelia pseudocelebensis sp. n. (figs. 35-40, 80-81) Type material. - Holotype d, SULAWESI TENGGARA: Tamborasi-Wolo, mountain stream, 1 Mar. 1989, N8926, leg. N. Nieser (RMNH). Paratypes: same data as holotype, 138 49 (incl. allotype) (19 macropterous) (allotype RMNH); Sungai Kolaka, upstream of Kolaka, 27 Feb. 1989, N8921A, 168 189 apt, 166 209 macr.(39 apt. PCHC); same, N8921B, 26 (macr.) 39 (29 apt, 19 macr. JTPC); Tamborasi, mouth of subterranean stream, 1.111.1989, N8925 816 219 (46 PCHC, 46 JTPC); 20 km E of Kolaka, mountain stream, 3 Mar. 1989, N8934 29 (all leg. N. Nieser). — PULAU BUTON, Ist stream road N of Bau-Bau, 8.i11.1989, N8935 4d 49, leg. N. Nieser. All apterous un- less otherwise indicated (fig. 81). The following museums received 1d 1@ paratypes each: BPUH, CHC, JTPC, MBBJ, ZMA; ZMC received a single 9, remaining paratypes in NC. Description. — Apterous form. Dimensions. Length d 2.98-3.80, 9 2.78-3.10; width d 1.1-1.2, 2 1.3-1.4; width of head d 0.76-0.82, 2 0.80-0.83; width of pronotum d 1.00-1.03, 9 1.02-1.13. Colour, black, pronotum with anterior transverse orange band only slightly broader than interocular space, laterally obscured by pale grey pruinose pilosi- ty. Basal third of first antennal segment, acetabula, fo- re and hind coxae and trochanters and basal third of anterior femur pale yellowish. Ventral pilosity of ab- domen in male yellowish. Venter with minute black denticles, which are dif- ficult to see due to pruinose dark grey background, extending onto episterna and head. Dorsum with a few short golden-yellow setae, longer and denser on genital segments of male. Vertex anteriorly with short dark setae, sparse long dark setae on head, sides of thorax and apex of abdomen. Antennae and legs with the usual pubescence and setae. Length of antennal segments I-IV d 0.83 : 0.42 : 0.46 : 0.48, 2 0.81 : 0.38 : 0.42 : 0.45. Pronotum long, covering mesonotum, in some specimens is the posterior margin of mesonotum narrowly visible. Length: width of pronotum 0.75 : 1.0. Length of me- tanotum on midline, 0.12. Abdominal tergites 1-5 subequal in length (0.12), tergite 6 slightly longer (0.17), tergite 7 distinctly longer (0.55). Male. Posterior trochanter length: width 0.30 : 0.17, without teeth or warts. Posterior femur rather slender, four times as long as wide (1.2/0.3), on ven- troposterior margin about halfway with a large spine about 0.6 as long as the width cf femur, with a row of NIESER & CHEN, Rhagovelia of Sulawesi 5-6 spines of decreasing length distally (fig. 37). Posterior tibia straight, armed beneath with five small blunt teeth proximally and three slightly larger shar- per teeth apically. Connexiva more or less horizontal or slightly slanting upward, gradually converging pos- teriad. Basal width of abdominal tergite 7 0.38, its median length 0.40. Abdominal venter with long, quite dense, golden yellow pilosity. Sternite 7 in la- teral view distinctly compressed dorsoventrally in posterior three fourths (fig. 35). Apical part of sterni- te 7 without pubescence, shining black, flattened with a low, narrow median carina. Segment 8 promi- nent (fig. 35), elongate, in dorsal view, length 0.50 width 0.30. Parameres as in fig. 36. Female. Generally with a more compact build as compared to male, due to compressed genital seg- ments. Posterior femur five times as long as wide (1.3/0.25). Spines on posterior femur and tibia essen- tially as in male. Connexiva more or less vertical, apex of connexiva triangularly produced caudally, with a long tuft of medially curved pilosity (figs. 39, 40). Tergite 7 with a short mediocaudal projection on hind margin, with a short tuft of setae apically, this character is, however, quite variable. Especially in some females from N8921A the projection is hardly recognizable. Tergite 8 distinctly (about 60°) slanting ventrad, proctiger pointing ventrad and somewhat re- tracted into segment 7 in most specimens. Sternite 7 large, shining, half as long as the remaining abdomi- nal sternites together (1.0/1.8). Gonocoxa not dis- tinctly visible. Macropterous form, as apterous except for full de- velopment of pronotum and wings. Dimensions, length d 3.5-3.8, 2 3.5-3.7, width of head 0.80- 0.82, humeral width of pronotum d 1.40-1.43, © 1.37-1.42. Pronotum broadly rounded posteriorly. Hemielytra dark brown with darker veins, three clo- sed cells, two basal, reaching about one third the length of hemielytron, the distal costal cell reaching well into distal half (fig. 80). Submedial carinae on base of abdomen reaching to hind margin of tergite 2. Etymology. — Pseudocelebensis, greek adjective mean- ing looking like celebensis, refers to the similarity be- tween R. celebensis and R. pseudocelebensis. Comparative notes. — Very similar to R. celebensis but the female has a small caudal projection on tergi- te 7, the male paramere is different (figs. 36, 41-44). In R. sulawesiana which is also very similar, the fema- le has the connexiva folded over the tergites apically, the paramere of male is different (fig. 45-46). See also comparative notes under R. daktylophora. Rhagovelia robina sp. n. (figs. 47-55, 82) Type material. — Holotype d: Sulawesi Tenggara: Road 275 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Rhagovelia A Rhagovelia | 7 R. celebensis ù @ R.minahasa R. pseudocelebensig @ A. lorelinduana R. daktylophora R. sulawesiana % R horaia Rhagovelia i Rhagovelia @ Rpruinosa i R.wallacei B A hamjadi i R. trichota % RB chrysomalia È R. tropidata R. blogiokommena R. robina R. kalami Figs. 81-84. Distribution of Rhagovelia species in Sulawesi. — 81, R. celebensis, R. daktylophora, R. pseudocelebensis, R. sulawe- siana; 82, R. wallacei, R. tropidata, R. blogiokommena, R. trichota, R. robina, R. kalami; 83, R. pruinosa, R. hamjadi, R. chryso- malla, 84 R. minahasa, R. unica, R. kastanoparuphe, R. lorelinduana, R. horaia. 276 Kolaka Kendari, 20 km E Kolaka, stream in hilly forest, 3 mar. 1989, N8934, leg. N. Nieser (RMNH). — Paratypes 66 49 apterous, 26 macropterous, same data as holotype (1? allotype RMNH, 58 29 NC, 1d 19 MBBJ, 1d PCHC, 1d JIPC, 16 19 ZMA) (fig. 82). Dimensions. Length d 3.38-3.50, 9 3.58-3.67; width of head d 0.80-0.81, © 0.82-0.87; width d 1.18-1.20, 2 1.27-1.32, width of pronotum d 1.07- 1.10, £ 1.10-1.12. Colour, yellow to orange reddish. Eyes dark casta- neous. Interoculus, posterior three quarters of prono- tum, abdominal tergites 2-6, proximal and lateral margins of tergite 7 in male, mesonotum and abdo- minal tergites except for central spot on tergite 6 in female, distal half of antennal segment 1 and remai- ning antennal segments, apical segment of rostrum, apex of fore femur, most of fore tibia and tarsus, mid- dle leg except for coxa and trochanter, dorsoapical streak on hind femur, hind tibia and tarsus dark brown to blackish. Body ventrally and dorsally densely covered with small black denticles. Dorsum without longer pilosity except on genital segments of male and posterior mar- gin of abdominal tergite 8 and connexiva in female. Head, sides of thorax, antennae and legs with the usu- al setae. Disk of pronotum without coarse punctures. Length of antennal segments I-IV d 0.70 : 0.40 : 0.49 : 0.41, 2 0.72 : 0.39 : 0.50 : 0.42. Pronotum long, covering mesonotum, length: width 1.1 : 0.85. Length of metanotum on midline, 0.12. Abdominal tergites 1-5 subequal in length (0.15-0.20), tergite 6 slightly (0.22) longer in male more distinctly so(0.30) in female, tergite 7 longer (0.40-0.45). Male. Posterior trochanter one and a half times as long as wide, with one small teeth and a about four minute teeth or warts. Posterior femur three and a half times as long as wide (1.37/0.40), on ventral margin about halfway a large spine with a length just over half the width of femur, with a row of 8-10 spi- nes of rapidly decreasing length distally and a row of 16 very small warts proximally (fig. 49). Anteriorly in distal half a row of 4-8 small warts. Larger spines black tipped, smaller spines and teeth entirely black or nearly so. Posterior tibia straight, armed beneath with a double row of about 20 pairs of warts of which the posterior ones are largest. Connexiva more or less horizontal, gradually converging to caudal margin of tergite 7, connexival points absent. Basal width of ter- gite 7 0.5, diverging caudally, caudal width 0.6. Segment 8 prominent in dorsal view, length and width about 0.6, sides slightly convex. Abdominal sternite 2 bluntly keeled in midline, length of sterni- tes 2-5 subequal (about 0.20), length of sternite 6 0.28, and of sternite 7 0.45. Abdominal sternite 7 la- terally compressed resulting in a broad median keel (fig. 47). This keel accentuated by a fringe of long pi- NieseR & CHEN, Rhagovelia of Sulawesi losity which runs over the midline of all abdominal sternites. Sternite 8 lateroproximally compressed. Parameres fig. 48. Female. Posterior trochanter without small teeth. Posterior femur four and a half times as long as wide (1.28/0.28), proximal row of 0-2 minute teeth, row of larger spines essentially as in male, posteroventral row of 0-2 smaller spines. Posterior tibia as in male. Metanotum with a pair of caudolateral broad blunt tooth-like swellings. Abdominal tergites 1 and 2 swol- len, in lateral view distinctly rising above the dorsal margin of connexiva. Tergites 4 and 5 with a broad blunt median keel. Connexiva (fig. 52) constricted on segments 4 and 5, caudally ending in blunt triangular apices fringed with black setae. Tergite 8 only hori- zontal in basal part, slightly curved ventrad apically, proctiger well visible. Gonocoxa 1 well visible (fig. 51). Macropterous form as apterous except for develop- ment of pronotum, hemielytra and dorsal abdominal carinae. Pronotum with coarse ill defined pits in pos- terior part, about 20/0.25 mm. Humeral width of pronotum 1.35, median length 1.40. Hemielytra with 4 cells and one or two variable adventitious cells (figs. 53-55). Dorsal abdominal carinae long, reach- ing to caudal margin of tergite 4. Etymology. — Robinos, a greek adjective meaning rose-coloured, refers to the reddish colour of the spe- cles. Comparative notes. — Similar to À. tropidata and À. wallacei but differing by the orange-like to pale red- dish metanotum and first abdominal tergite in À. ro- bina, which are dark in the other species. Rhagovelia trichota sp. n. (figs. 56-60, 82) Type material. — Holotype d, Sulawesi Tenggara, Tamborasi, creek of subterraneous stream, 1 Mar. 1989, N8925 (ZMA). — Paratypes: same data as holotype, 50d 329 (16 29 incl. allotype ZMA; 16 12 PCHC, 1d 19 BPUH; 16 19 MBBJ, 1d 19 JTPC; 264 22 RMNH, remaining specimens NC); Sg. Kolaka, 27.11.1989, 26 ; Sg. Konaweha at Sanggona, 7.xi.1989, J. van Tol, 38 32 (RMNH, 1d 19 NC). All apterous (fig. 82). Description. — Apterous form. Dimensions. Length d 3.10-3.17, 2 3.30-3.40; width 3 1.07- 1.14, 2 1.18-1.36; width of head d 0.78-0.80, 2 0.80-0.83; width of pronotum d 0.99-1.02, 2 1.10- 1.18. Colour, dull black, orange yellow transverse band on anterior margin of pronotum laterally reaching halfway the eyes. Basal half of first antennal segment, coxae, trochanters and basal half of anterior femur yellowish. Distal margin of connexiva, brownish ZII], TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 orange distinct in female, often indistinct in males. Abdominal venter with variable area along median line and on posterior sternites medium brown. Minute black denticles difficult to distinguish, on the propleura they are, however, usually quite dis- tinct. Dorsal side of male, except on head, with loose- ly set long erect setae. Dorsal side of female bare ex- cept on connexiva, abdominal dorsum especially on tergites 1-4 with a grey pruinose bloom. Thoracic pleura with grey pruinose bloom best developed in fe- males. Vertex, sides of thorax, antennae and legs with the usual pubescence and setae. Length of antennal segments I-IV & 0.80 : 0.41 : 0.48 : 0.49, 2 0.80 : 0.43 : 0.49 : 0.49. Pronotum long, covering mesonotum. Central part of pronotum smooth, lateral and hind margins with comparatively fine punctures, along hind margin of orange yellow transverse band a row of coarser punctures. Length: width of pronotum 0.8 : 1.0. Length of metanotum on midline, 0.12. Abdominal tergites 1-5 subequal in length (0.20-0.22), tergite 6 subequal (0.22) in male, slightly longer (0.30) in female, tergite 7 distinctly longer (0.40 in male, 0.50 in female). Male. Posterior trochanter length : width 0.30 : 0.15, with about 5 small teeth. Posterior femur rather slender, slightly less than four times as long as wide (1.20/0.33), on ventroposterior margin about halfway with a large, yellowish to blackish, usually medium brown, spine, its length (0.17) half the width of fe- mur, with a row of 12-13 very small black teeth proxi- mally and a row of 5-7 black spines of decreasing length distally (fig. 58). Posterior tibia straight, armed beneath with an irregular row of about 20 small teeth. Connexiva more or less horizontal gradually conver- ging posteriad, no caudal points. Basal width of abdo- minal tergite 7 about three quarters its median length (0.28/0.40). Abdominal sternites 2-5 with a low broad ridge, accentuated by a band of long pilosity (fig. 56). Sternite 7 slightly less than half as long as the remaining ones (0.4/1.0), without median keel. Genital segments relatively small, tergite 8 about three quarters the length of 7 (0.3/0.4). Sternite 8 flattened with a low broad carina. Parameres as in fig. 57. Female. Posterior femur nearly five times as long as wide (1.20/0.25). One or two small teeth proximally of large spine, which is about half as long as width of femur, on hind femur, distal decreasing row of about six spines. Teeth on posterior tibia essentially as in male. Connexiva more or less vertical, distal (‘dorsal’) rim pubescent except on segments 3 and 4 where it is bare and strongly shiny. Connexiva curved inward on segments 3 and 4, folded and dorsocaudally touching; ending in narrowly triangular points (fig. 60). Tergite 7 mostly hidden from view. Tergite 8 and proctiger nearly vertical, genital segments retracted into seg- ment 7 (fig. 59). Sternite 7 large, shining, half as long 278 as the remaining abdominal sternites together (0.6/1.2). Macropterous form, unknown. Etymology. — Trichotos, greek adjective meaning hairy, refers to the hairy aspect of dorsal surface of male. Comparative notes. — The long and relatively den- se dorsal pilosity distinguishes males from its Sulawesi congeners. Females are similar to those of R. sulawe- stana but a caudal finger like process is lacking and the connexiva are different (figs. 60, 61). Rhagovelia tropidata sp. n. (figs. 62-67, 82) Type material. — Holotype d : Sulawesi Tengah, between Desa Seseba and Singsing Camp, SW of Luwuk, c. 1°09’S 122°31’E, 80m, narrow stream in lowland rainforest, 14.Oct.1989, sample Sul.12, J. P.Duffels (ZMA). — Paratypes: same data as holotype 7d 69 (4d 49, including allotype 2 ZMA, 2d 29 NC, 16 PCHC); Batui River at Singsing Camp, SW of Luwuk, c. 1°09S 122°31’E, 90m, middle sized stream in lowland rainforest, 14-17 Oct. 1989, sample Sul. 14, J. P. Duffels, 38 22 (ZMA, 16 NC); SW Luwuk, Toptop Camp along Batui River, 1°09’S, 122°31’30”E, 120 m, lowland rainforest, 19-21 Oct 1989, leg. J. P. Duffels, sample Sul. 18, 16; Luwuk area, S. Biak, 21 Oct. 1989, J. van Tol, 38 22 (16 12 NC); Batui River at Singsing camp, 15 Oct. 1989, J. van Tol, 16 19 (RMNH). All apterous (fig. 82). Description. — Apterous form. Dimensions. Length d 3.20-3.32, 9 3.22-3.35; width d 1.2-1.3, 2 1.2-1.3; width of head d 0.76-0.80, 2 0.77-0.80; width of pronotum d 1.18-1.30, 2 1.26-1.32. Colour, dorsally dark brown, broad orange yellow band anteriorly on pronotum extending to propleura, covering anterior quarter of pronotum. Dorsal two thirds of connexivum orange yellow, in male caudal half of abdominal tergite 7 and posterior segments orange yellow. Venter brownish, laterally (including acetabula) and caudally orange yellow. First antennal segment pale yellow, other segments brown. Legs ba- sally (up to halfway femur) yellowish, apically casta- neous. ; Posterior half of jugum of head, anterior half of propleura, thoracic sternum, dorsal margin of aceta- bula and abdominal sternites 1-6 with small black denticles. On darker parts of venter difficult to see be- cause of lack of contrast. Dorsum rather sparsely clothed with golden pubescence, longer and thicker towards apex of abdomen, especially in male. Vertex anteriorly with short dark setae, sparse long dark setae on head, sides of thorax and apex of abdomen. Disk of pronotum with large impressed pits, about 25/0.25 mm. Antennae and legs with the usual pubescence and setae. Length of antennal segments I-IV d 0.74 : 0.40: 0.49 : 0.41, 2 0.70 : 0.39 : 0.49 : 0.41. Pronotum long, covering mesonotum, length : width 0.8 : 1.1. Length of metanotum on midline, 0.12. Abdominal tergites 1-6 subequal in length (0.15-0.20), tergite 7 longer (0.35-0.40). Male. Posterior trochanter with one larger, two small teeth and a group of about five small warts. Posterior femur three times as long as wide (1.35/0.45), on ventral margin proximally with about 20 minute and closely set teeth, followed about half- way by a large spine about as long as half the width of femur, with a row of 9-10 spines of decreasing length distally. Anteroventrally a row of 5 smaller spines lo- cated in distal part. Larger spines black tipped, smal- ler spines and teeth entirely black or nearly so (fig. 64). Posterior tibia virtually straight, armed beneath with a double row of about (2x) 15 pegs. Connexiva more or less horizontal, gradually converging poste- riad. Abdominal sternite 7 laterally compressed in posterior two thirds, resulting in a broad distinct me- dioventral keel in posterior part. Keel accentuated by a dense fringe of long pilosity (fig. 62). Genital seg- ments prominent, 1.5 times as long as sternite 7, ster- nite 8 with an indistinct ventral keel. Parameres long (fig. 63), apically curved mediad. Female. Posterior trochanter without small teeth. Posterior femur four times as long as wide (1.30/0.30), row of minute teeth absent, row of larger spines essentially as in male, anteroventral row of 2-4 smaller spines located in basal part. Posterior tibia as in male. Connexiva more or less vertical, on tergites 4 and 5 slightly curved inward, thickened and beset with dense pilosity. Apex of connexiva truncate, dor- soapically with a small tuft of pilosity (figs. 66-67). Tergite 8 only slightly slanting ventrad, proctiger pointing ventrad in most specimens. Gonocoxa | well visible, laterally somewhat compressed. Macropterous form, unknown. Etymology. — Tropidatos, greek adjective meaning ‘provided with a keel’, referring to the well developed keel on abdominal sternite 7 of male. Comparative notes. — Similar to À wallacei Polhemus & Polhemus which differs in the male by a less well developed keel on sternite 7 and shorter pa- rameres (fig. 76) and in the female by connexiva not thickened on segments 4 and 5 and apical segments (8 and 9) hardly visible, withdrawn into segment 7. Abdominal segment 8 and the long paramere remind somewhat of R. meikdelyi Polhemus & Polhemus from Ambon but in detail there are many differences of which the more strongly inflated posterior femur (only slightly over twice as long as wide in males, th- ree times as long as wide in females) is one of the most distinct. Finally À. lorelinduana Polhemus & Polhe- Nieser & CHEN, Rhagovelia of Sulawesi mus has a strongly developed keel on sternite 7 of male but this species has an aberrant paramere and proctiger (fig. 23) in male and a posteriorly prolonged abdomen with the connexiva bent over its apex in fe- male. Discussion Polhemus & Polhemus (1988) distinguished sever- al groups within the genus, of which three occur in Sulawesi. The sarawakensis-group, however, has been recorded by a single macropterous specimen of R. sa- marinda only. So we will restrict our discussion to the orientalis- and papuensis- group. The orientalis-group is a small group of about eight species distributed in the Philippines and Sulawesi. The main synapomorphy for this group is the hemi- elytral venation, three cells, of which the apical one reaches into the apical quarter of hemielytron (figs. 79-80). This seems a rather fickle character in view of the variability of hemielytral venation in some species and even between hemielytra of a specimen (figs. 53- 55). As the general appearance of at least the Sulawesi species of this group is homogeneous and, apart from males of R. sulawesiana, distinct from the species of the other group, it seems, nonetheless, justified. In Sulawesi this group is represented by three species: R. celebensis, R. daktylophora and R. pseudocelebensis. The remaining Sulawesi species belong to the pa- puensis-group, a large group with a wide distribution: India and Sri Lanka through Taiwan, the Philippines, Northern Borneo, Sulawesi and Maluku (the Moluccas) to New Guinea, Solomons and Australia. According to Polhemus & Polhemus (1988) this group is probably polyphyletic. A more restricted pa- puensis-group is characterized by a strongly inflated hind femur, a curved hind tibia with one or more lar- ger subapical teeth and a comparatively long third an- tennal segment. R. papuensis Lundblad (Papua New Guinea), R kawakamii (Matsumura) (Northern Borneo), R. grayi, R. horaia and R. lorelinduana (Sula- wesi) and R. kalami (Buton) are all representatives of this group. But as many species of this group (s. 1.) are still to be described and the Sulawesi fauna constitu- tes only a fraction of it an analysis of this group here would be premature. The distribution of most species seems to be rather restricted, although one should not forget, looking at these small maps, that the distance between the northern and southern points of the main island of Sulawesi is over 1000 km and the west-east length of the northern peninsula c. 600 km. Secondly, more or less extensive collecting has been done in a rather small number of restricted areas only. The maps are based on the data from Polhemus & Polhemus (1988) and our own. 279 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 The three species of the orientalis-group have, so far, neatly separated distributions (fig. 81). The most widely distributed species seem to be R. pruinosa and R wallacei (figs. 83, 82). R. kastanoparuphe and R. pseudocelebensis are interesting as they occur with R. pruinosa both on the main island and on Buton (figs. 81, 83-84), R. trichota has been found on both sides of Pegunungan Mengkoka. The remaining species are as yet only known from restricted areas. Nearly all specimens have been collected by experienced collec- tors. So we may assume that having been collected only on one or very few close localities of these local species indeed reflects restricted distribution rather than more cryptic habits. Available data suggest that the widely distributed species tend to be lowland spe- cies, and most, but not all, of the restricted species live on higher ground. A definite conclusion in this respect has to wait until more data become available. Likewise our data are not sufficient to establish altitu- dinal separation between species occurring geographi- cally at the same ‘points. Polhemus & Polhemus (1990) have postulated ‘centres of species endemism’ in Sulawesi. As average areas of distribution of species of various genera of Gerromorpha and Nepomorpha in Malesia are not yet known, we prefer to avoid the predicates endeme and endemism for the present. Although their centres look somewhat like areas in which more intensive col- lecting has been done compared to the remaining of Sulawesi, our results with Rhagovelia and Enithares (Nieser & Chen 1991) coincide with the regions with different faunal elements of water bugs in Sulawesi they postulated. Our present data also support their supposition that Sulawesi Tenggara (apparently in- cluding Buton) and the eastern peninsula of Sulawesi Tengah probably are such regions. It should, how- ever, been pointed out that our results with the gerri- ne genus Limnometra (Nieser & Chen 1992), which 280 also contains mostly stream inhabiting species, do not strongly support this pattern. In Zimnometra there are fewer species of which relatively more occur at least in two of these regions. REFERENCES Bacon, J. A., 1956. A taxonomic study of the genus Rhagovelia of the Western Hemisphere. — Kansas University Science Bulletin 38:695-913. Chen, P. P. & N. Nieser, 1992. Gerridae, mainly from Sulawesi and Pulau Buton (Indonesia). — Tijdschrift voor Entomologie 135: 145-162. Hungerford, H. B. & R. Matsuda, 1961. Some new species of Rhagovelia from the Philippines. — University of Kansas Scientific Bulletin 42: 257-279. Lundblad, ©., 1936. Die altweltlichen Arten der Veliidengattungen Rhagovelia und Tetraripis. — Arkiv för Zoologi 28A(21): 1-21, 13 pls. Lundblad, O., 1937. Einige neue oder wenig bekannte osta- siatische Rhagovelia-Arten. — Entomologisk Tidskrift 58: 1-9. Nieser, N., & P. Chen, 1991. Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau Buton (Indonesia). — Tijdschrift voor Entomologie 134: 47-67. Nieser, N., & P. Chen, 1992. Revision of Limnometra Mayr (Gerridae) in the Malay Archipelago. — Tijdschrift voor Entomologie 135: 11-26. Polhemus, J. T., & D. A. Polhemus, 1988. Zoogeography, ecology, and systematics of the genus Rhagovelia Mayr (Heteroptera: Veliidae) in Borneo, Celebes, and the Moluccas. — Insecta Mundi 2: 161-230. Polhemus, J. T., & D.A. Polhemus 1990. Zoogeography of the aquatic Heteroptera of Celebes: regional relationships versus insular endemism. — In: Knight, W. J. & J. D. Holloway (Eds.), Insects and the rain forests of South East Asia (Wallacea). Royal Entomological Society of London. Received: 7 October 1992 Revised manuscript accepted: 1 June 1993 Appendix 1 Rhagovelia species collected in Sabah Rhagovelia kawakamii (Matsumura) Kotovelia kawakamii Matsumura, 1913: 99. Rhagovelia kawakamit, Polhemus & Polhemus 1988: 206- 207, figs. 74-80, 227 (redescription, synonymy). Material. — EAST MALAYSIA: Sabah, Pulau Ranggi, logging camp, 7.10N 117.10E, 23. IV. 1987, leg. J. Huisman 2d 1 apterous (1d 19 RMNH, 1d NC). Distribution. — Taiwan, Philippines, N. Borneo. Rhagovelia samardaca Polhemus & Polhemus Rhagovelia samardaca Polhemus & Polhemus, 1988: 200- 201, figs. 194-200, 226. Material. — EAST MALAYSIA: Sabah, Lahad Datu, 60 km W of Danum Valley field Centre at junction of Sg. Segama and Sg. Palum Tambun, 4°58’N 117°48’E, 150m, bridge of Segama, 26. IV. 1987, 18.30-21.00h, clearing edge of un- touched evergr. lowl. rainforest, leg. van Tol & Huisman, 46 29 apterous (RMNH, 1d 1 NC). Distribution. — Sabah. Rhagovelia silau Polhemus & Polhemus Rhagovelia silau Polhemus & Polhemus, 1988: 184-186, figs. 99-105, 227. NIESER & CHEN, Rhagovelia of Sulawesi Material. — EAST MALAYSIA: Sabah, 105 km S of Beaufort: Long Pa Sia area, Sg. Ritan, 4°24’N 115°42’E, 1160m, un- disturbed evergreen tropical forest, 8.iv.1987, J. van Tol & J. Huisman, 18 39 (1d 19 apt, 19 macr. RMNH, 19 apt. NC). Distribution. — Previously recorded from Sabah in mountainous terrain, localities 900-2100m asl. Rhagovelia tawau Polhemus & Polhemus Rhagovelia tawau Polhemus & Polhemus, 1988: 187-188, figs. 15-22, 224. Material. — EAST MALAYSIA: Sabah, 70 km West of Lahad Datu, Main Trail West 12, 180m, narrow creek, 2.xii.1989, M. J. & J. P. Duffels, sample Sab. 52, 54 19; same, 4 km S Main Trail West 5, near Sungai Segama, 150m, middle si- zed stream and waterfall, 3.xii.1989, M. J. & J. P. Duffels, sample Sab. 54, 5d 29 (ZMA, 3d 19 NC); 60 km W Lahad Datu, Danum Valley Field Centre, road DVFC-Kg Silam nr km 68.3, 4°58 N 117°48’E, at junction of Sg. Sagama and Sg. Palum Tambun, 150m, disturbed evergreen lowland rainforest, 26.iii.1987, leg. van Tol & Huisman, 1d. All ap- terous. Distribution. — So far only known by the type se- ries from Tawau (near Kalabakan) which is also in the eastern coastal region of Sabah, close to the border with Kalimantan Timur. 281 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 BOOK REVIEW Kuchlein, J. H. (with collaboration of J. H. Donner), 1993. De Kleine Vlinders. Handboek voor de Faunistiek van de Nederlandse Microlepidoptera. - PUDOC, Wageningen. 715 pp., 1372 maps, 8 colour plates, textfigs. [ISBN 90- 220-1038-4] [The small moths. Manual for the Faunistics of Netherlands Microlepidoptera, in Dutch, with 5 pp. English summary]. Price Hfl. 150.- (ca US $ 80). This book deals with the Microlepidoptera of the Netherlands, the main body being distribution maps of all 1372 species (4 missing maps are provided on a separate er- ratum sheet). The first 170 pages contain lengthy general chapters, dealing respectively with Morphology of adult, larva, feed- ing traces (mines), Ecology, Taxonomy and zoogeography, Human influence, Faunistics, and a key to families based on adults with a bibliography of key-works. Apart from the maps, the main part of the book contains a checklist with annotations about faunistic status (incor- rectly referred to as ecological status), indication of occur- rence in the periods 1850-1900, 1900-1950 and later than 1950, number of provinces, number of grid squares, indica- tion of abundance and codenumbers for hostplants. Further there are annotations for part of the species, usually explain- ing past identification problems, or indication of first find- ings. A final chapter contains some general results and conclu- sions. Finally there is a long English summary, making the book also accessible for foreign readers. The book is attractively made, well printed, and the colour plates, with mounted moths, mostly from the.collec- tion of J. C. Koster, are well made, although the black back- ground sometimes tends to make the colours of the moth too dark. One colour plate depicts leaf-mines. The book contains a wealth of interesting information, and there lies in my opinion the main problem of the book: it is as if the author could not make a choice whether to write a general manual about microlepidopteran studies or to present the results of faunistic research on the species in the Netherlands. The book would have been much nicer and easier to use if a choice had been made before writing. The general introduction chapters are really more a student textbook, certainly with interesting information, but to be found in many other textbooks as well, albeit with fewer ex- amples from the Lepidoptera. The connection between the lengthy introduction to ecology for instance and the individ- ual species is difficult to find. In fact there are hardly any notes about the habitats and ecology of individual species. The only ecology found is a list of hostplant species (and then with numbers, which have to be searched in the appen- dix for the correct name), without any details. The most interesting and original part of the introducto- ry chapters is in my opinion the historical chapter on Dutch Microlepidopterists. The key to families could be useful, and the colour plates certainly are helpful. It is a pity that the family names are not given in the captions for the plates. One photograph has mistakenly been used twice: plate 5: 4 (Mompha epilobiella) shows the photograph of Bucculatrix ulmella, the same as plate 3: 1. The bibliography of key works is very long, for new work- ers disappointingly long. Nowhere a choice has been made, nor annotations given about the usefulness of a book or arti- 282 cle. Some references seem completely misplaced, like keys for New Zealand species for groups where recent European keys are available as well. On the other hand some important titles are missing. The checklist is an important summary of present know- ledge of Dutch faunistics: it is therefore to be regretted that here and in the species annotations there are many mistakes or inaccuracies. Despite a well written introduction to zoo- logical nomenclature, the author nowhere indicates his source for the nomenclature used. Several recently proposed changes are not incorporated, but some are (like the generic name Luquetia Leraut). Notable mistakes are the use of Episcardia for Haplotinea, shown to be incorrect by Robinson in 1984, the use of the generic name Glyphipteryx Curtis, rejected by the ICZN in 1986 and the use of the re- placement name Nemophora violaria (Razowski), which is only needed (because of secondary homonymy) when used in the genus Adela. Such mistakes could easily have been avoided by closer contact with specialists. The maps show the distribution in 5 X 5 km grid cells with a single symbol. Although the databank used contains information about the date of the records, all records have the same symbol, so a well dotted map may also show the distribution of about 100 years ago. It would have been bet- ter to indicate periods, using different symbols as is general practice in similar atlasses. Some rapidly declining species can not be identified from these maps, nor from the text. In some cases there is considerable doubt about the correctness of some dots: the large number of dots in the species Stigmella samiatella and S. ruficapitella could only be ex- plained by the inclusion of records based on mines. However, there are no reliable characters separating these mines, nor the larvae. In some cases I am missing previously published records, in others there are dots for records which have been shown to be incorrect in literature. The presentation chosen is partly impractical because in order to find all information about a species, one needs to switch from the map to the list, for which one needs the ap- pendix to understand the hostcodes, and the introduction to understand the other codes. The list also indicates whether there is an annotation for the species, but the caption to the map does not give this reference. Finally, I would like to comment on the choice of the spe- cies treated: it is unfortunate that this book chooses to in- clude only the Microlepidoptera in the old sense and exclud- ing such related groups as Psychidae or Sesiidae. On the other hand one could wonder about the inclusion of single import events of species found on Bonsai trees imported from Japan and China. These species have been found by the Plant Protection Service, and the trees were destroyed subsequently. Such species do not belong on the list of spe- cies occurring in The Netherlands, otherwise all species in- troduced alive in our country could be listed as well with the same argumentation, including those introduced on pur- pose by scientists or breeders! The maps of such species do not give any information (some of the maps even do not contain any dot, yet they are printed). In conclusion: a book with much faunistic information, useful for Microlepidopterists in Europe, but to be used with care. The long introduction is only of use for readers with knowledge of Dutch. In my opinion too expensive de- spite the excellent printing. [E. J. van Nieukerken] O. A. SETHER & T. ANDERSEN Museum of Zoology, University of Bergen LOBOSMITTIA, A NEW GENUS OF ORTHOCLADS FROM TANZANIA AND TURKEY (DIPTERA: CHIRONOMIDAE) Sæther, O. A. & T. Andersen, 1993. Lobosmittia, a new genus of orthoclads from Tanzania and Turkey (Diptera: Chironomidae). — Tijdschrift voor Entomologie 136: 283-287, figs. 1-10. [ISSN 0040-7496]. Published 10 December 1993. The genus Lobosmittia is erected for a new species from the West Usambara Mts in NE Tanzania, L. basilobata and for Pseudosmittia invaginata Caspers & Reiss, 1989 from Turkey. The genus differs from other orthoclads with bare eyes, wings and squama; by completely lack- ing acrostichals, median microtrichial tuft or less sclerotized median area on scutum; by having costa not or only slightly extended; fine, triangular, strongly pointed anal point bare of micro- trichia; and gonostylus with either a basal or preapical lobe. The Parakiefferiella group of gene- ra or Acamptocladius Brundin and related genera are the most likely closest relatives. Department of Systematic Zoology, Museum of Zoology, University of Bergen, Musépl. 3, N-5007 Bergen, Norway. Key words. — Chironomidae, Orthocladiinae, new genus, Tanzania, Turkey. INTRODUCTION During an expedition by the Museum of Zoology, University of Bergen, to the montane evergreen fo- rests in the West Usambara Mountains, NE Tanzania in the autumn of 1990, several interesting new gene- ra and species of chironomids were collected together with a few genera new to the African continent (Andersen & Szther 1993, in press a, b, Sæther & Wang 1992). One of the new genera had what in some specimens looked like a double gonostylus. However, other views showed that it was a basal lobe. Other characteristics made it clear that the species could not be placed in any presently recognized ge- nus. The senior author, together with L. C. Ferrington jr., Lawrence, Kansas, presently is revising the genus Pseudosmittia Goetghebuer. Many species in that genus are tentatively placed since the genus has not been well defined. When examining types it was found that Pseudosmittia invaginata described by Caspers & Reiss (1989) could be placed in the same genus as the species from Tanzania. The field work, which included an extensive use of Malaise traps and sweep nets, was mainly conducted in the Mazumbai Forest Reserve in the eastern part of the West Usambara Mts. A thorough description of the vegetation in these mountains is given by Iversen (1991). Our Malaise trap localities along the Kaputu Stream near Mazumbai are described in Andersen & Johanson (in press). METHODS AND TERMINOLOGY The material examined was mounted on slides fol- lowing the procedure outlined by Sæther (1969). The general terminology follows Sæther (1980). The me- asurements are given as ranges followed by mean and the number measured (n) in parentheses. The types of Lobosmittia basilobata sp. n. are de- posited in the Museum of Zoology, University of Bergen, Norway (ZMBN). Lobosmittia gen. n. Type species. — Lobosmittia basilobata sp. n. by present designation. Other included species: Lobosmittia invaginata (Caspers & Reiss, 1989: 123, Figs. 11-13, as Pseudosmittia invaginata), comb. n. Diagnostic characters. — The bare eyes, wing mem- brane and squama, combined with complete lack of acrostichals, microtrichial tuft on median hump or sclerotized area on scutum, not or only moderately extended costa, and pointed, triangular anal point, free of microtrichia will separate the genus from oth- er orthoclads. Etymology. — From the Greek lobos, a lobe, and Smittia, a related genus of Orthocladiinae and a com- mon ending among orthoclads, referring to the basal or subapical lobes of the gonostyli of the two includ- ed species. 283 TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Description Small species with wing length 0.7 - 1.1 mm. Eyes bare, without dorsomedial elongation. Coronal suture well developed ending in frontal pro- jection. Temporal setae consisting of weak inner ver- ticals and strong outer verticals. Tentorium and stipes normal. Antennae of male with 13 flagellomeres; an- tennal groove starting on flagellomere 3; sensilla chaetica on flagellomeres 2, 3 and 13; apex without apical setae. Antepronotum with median lobes narrowed medi- ally and slightly separated, with 1 lateral seta. Dorsocentrals several; acrostichals, microtrichial tuft, hump or unsclerotized median area of scutum absent; prealars few, in one or two groups; supraalars present or absent. Scutellum with few setae. Humeral pit not distinctive. Wing membrane bare, with very fine punctuation of microtrichia; veins brown. Anal lobe weak or ab- sent. Costa not to moderately extended; R,,, running in the middle between R, and R,,,, ending close to Rs Ry, ending clearly proximally of end ofM,,;; Cu, sinuate; FCu far distally of RM; postcubitus extend- ing beyond FCu; anal vein not reaching FCu. Brachiolum with 1 seta, other veins bare. Squama bare. Sensilla campaniformia about 8-10 on base of brachiolum, 3 below seta, and about 8-10 at apex of brachiolum; 1 at base of subcosta, 1 on FR and 1 at base of R.. Tibial spurs and tibial comb normal. Pulvilli ab- sent, at least in L. invaginata. Tergite IX with few to several setae at base of trian- gular, sharply pointed anal point which is bare of se- tae and microtrichia. Sternapodeme with weak or no oral projections. Phallapodeme normal. Virga well developed, consisting of few needle-like spines. Gonocoxite with long, relatively low inferior volsella. Gonostylus either with a basal crista dorsalis appear- ing as a basal appendage in some views or with a long apically angled crista dorsalis making the gonostylus appear to have a deep apical invagination. Female, pupae and larvae unknown. Systematics In the key to adult males of Orthocladiinae by Cranston et al. (1989) Lobosmittia will key to couplet 98. It will not key further since the pulvilli apparent- ly are absent or vestigial, while the anal point extends from the posterior margin of tergite IX. Caspers & Reiss (1989: 12) discussed the placement of their spe- cies as Pseudosmittia invaginata on the base of the si- milarity of Pseudosmittia Goetghebuer and Prosmittia Brundin. Cranston & Oliver (1988) even synonymi- zed the two genera based on the fact that Prosmittia nanseni Kieffer (Kieffer 1926: 82) is a good Pseudosmittia. Oliver (1963: 177) first placed nanseni in the genus Prosmittia apparently based on the simi- larities in the hypopygia between nanseni and Prosmittia jemtlandica (Brundin, 1947). Reexamin- ation of the type of P. nanseni by Sæther et al. (1984: 270) did not change the placement of the species. However, the wing venation could not be discerned on the damaged male. The senior author together with Dr. L. C. Ferrington jr., Lawrence, Kansas, pre- sently is revising the genus Pseudosmittia and we have examined P. jemtlandica as well as a large amount of P. nanseniand most types of Pseudosmittia. All species of Pseudosmittia have 2 or occasionally 4-11 (as in P. nanseni) short, but relatively strong acrostichals, with distinct sockets, in a pale, less sclerotised median area on the scutum. There is never a median microtrichial tuft as in Parakiefferiella Thienemann, and no me- dian tubercle, but occasionally there is a hump in so- me teneral specimens. On the wing, vein R,,, may end slightly distal to end of M,,,, but mostly ends well proximal of the end of M... In Prosmittia jemtlandi- ca there is no sign of acrostichals, microtrichal tuft, hump or pale area on scutum, and R,,, ends far dis- tally of the end of M,,, almost reaching the wing ap- ex. The illustration by Brundin (1947: fig. 16) of Prosmittia jemtlandica (as Pseudosmittia jemtlandica) is correct and not erroneous as previously presumed. The genus Prosmittia Brundin (1956: 58, 165) clear- ly has to be resurrected. The wing venation, however, appears to place it close to Unniella Sether, but not to Pseudosmittia nor to Lobosmittia. Lobosmittia, however, has an anal point placed on the posterior margin of tergite IX and does not appear closely related to any of the above mentioned genera. Boreosmittia Tuiskunen appears to be the most simi- lar genus. However, without knowledge of the imma- tures a more definite placement is as yet not possible. Lobosmittia basilobata sp. n. (figs. 1-10) Type material. — Holotype 6, TANZANIA, Tanga region, West Usambara Mts, Mazumbai, Kaputu Stream, 1640 ma. s. l., sweep net at waterfall, 28.x.1990, ZMB's Tanzania Expedition (ZMBN No. 156).— Paratypes: 4d, as holotype; 1d as holotype except Malaise trap loc. 2, 1650 m a.s. |., 2.-6.xi.1990; 26 as holotype except Malaise trap loc. 10, 1420 m a. s.1., xii.1990. Figs. 1-10. Male imago of Lobosmittia basilobata gen. n., sp. n. — 1, head; 2, thorax; 3, third palpal segment; 4, tentorium and stipes; 5, wing; 6, hypopygium, with dorsal aspect to the left, ventral to the right; 7, anal point, lateral view; 8-10, gonosty- lus, different views. 284 SÆTHER & ANDERSEN: Lobosmittia gen. n. 285 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Diagnostic characters. — The basal lobe of the go- nostylus easily separates L. basilobata from L. invagin- ata. Other characteristics of L. basilobata include ab- sence of supraalars, fewer prealars and inner verticals and more numerous sensilla clavata at apex of the third palpal segment. Etymology. — From the Latin basis, base, pedestal, and New Latin, /obatus, lobed, referring to the shape of the gonostylus. Description Male imago (n = 7-8, except when otherwise sta- ted). — Total length 1.34-1.46, 1.40 mm (6). Wing length 0.75-0.83, 0.81 mm. Total length / wing length 1.57-1.87, 1.73 (6). Wing length / length of profemur 2.73-3.08, 2.88. Coloration brownish black with yellowish shoulders and anepisternum. Legs yellowish brown. Head (Fig. 1). AR 0.39-0.49, 0.42. Ultimate fla- gellomere 123-171, 141 um long. One specimen with division between flagellomere 12 and 13 indis- tinct, if antenna regarded as 12-segmented AR 0.56. Temporal setae consisting of 2-3, 2 mostly widely separated, weak inner verticals; and 3 strong outer verticals. Clypeus with 4-6, 5 setae. Tentorium (Fig. 4) 80-94, 85 um long, 13-18, 15 um wide. Stipes (Fig. 4) 57-66, 62 um long, 21-27, 25 um wide. Palp segments lengths in um: 14-18, 15; 18-23, 21; 27-39, 34; 32-41, 38; 46-66, 59. Third palpal segment (Fig. 3) with 6-7, 6 (6) sensilla clavata. Coronal suture end- ing in frontal projection. Thorax (fig. 2). Antepronotum with 1 lateral seta. Dorsocentrals 8-11, 9; prealars 3; supraalars absent. Scutellum with 2 setae. Wing (fig. 5). VR 1.31-1.45, 1.40. Costal exten- sion 23 -62, 35 mm long. Legs. Spur of front tibia 29-39, 34 im long; spurs of middle tibia 14-18, 16 um (6) and 13-18, 15 um long; of hind tibia 32-39, 35 [um and 11-16, 14 um. Width at apex of front tibia 16-18, 17 um; of middle tibia 16-21, 19 um; of hind tibia 25-34, 30 um. Hind tibial comb of 13-15, 14 setae; shortest setae 18-25, 21 pm long; longest setae 26-34, 31 um long. Lengths of front, middle and hind femora (in Um) as: 252- 306, 282; 279-315, 306; 288-324, 308; of tibiae: 266-315, 293; 288-360, 325; 315-360, 339. Tarsomeres 1 and 2 of hind legs 171 um (1) and 90 Um (1) long, other tarsomeres lost. Hypopygium (fig. 6). Anal point 18-27, 21 pm long; with 9-14, 10 setae at base (fig. 7); laterosternite IX with 3 setae. Phallapodeme 39-48, 45 um long, transverse sternapodeme 34-43, 41 um long. Virga 14-19, 16 um long. Gonocoxite 91-101, 97 um long; with long inferior volsella apically hooked in some views. Gonostylus 39-45, 42 pm long; basal lobe end- 286 ing 21-27, 24 um from base (figs. 8-10); megaseta 9- 14, 10 pm long. Lobosmittia invaginata (Caspers & Reiss) comb. n. Pseudosmittia invaginata Caspers & Reiss, 1989: 123, figs. 11-13. — Holotype d, TURKEY: Kars province, W. Sarikamis, Soganli railway station, 2100 m a. s. l., 5.vii. 1985, W. Schacht (Zoologischen Staatssammlung, Miinchen). [examined] Diagnostic characters. — The subapical invagina- tion of the gonostylus easily separates L. invaginata from L. basilobata. Other characteristics of L. inva- ginata includes the presence of 1 supraalar, 6 inner verticals and 1 sensillum clavatum at the apex of the third palpal segment. Description The species is well described by Caspers & Reiss (1989: 123, figs. 11-13). However, this description contains a few mistakes. There are 6 weak inner verti- cals reaching from near the middle of the head to the position of the outer verticals. There also are 3 stron- ger outer verticals, apparently 8 setae on the clypeus, 1 lateral antepronotal and 1 supraalar. The first palp segment is shorter than the second, our measure- ments are 23 Mm and 30 um. The leg segments lengths appear to have been measured as total lengths, not following Schlee (1966), as especially the femora and ta, are about 15% too long compared with our measurements, while ta, to ta, are quite close. ACKNOWLEDGEMENTS The expedition to Tanzania was funded by The Norwegian Council for Science and the Humanities (NAVE). Financial support was also given by Nansenfondet. Thanks are also due to the staff at the Department for Forest Biology, Sokoine University of Agriculture (SUA), Morogoro, Tanzania and to the Norwegian Agency for Development (NORAD), Dar es Salam, Tanzania, for their cooperation and sup- port. Gladys Ramirez made the slide preparations of Lobosmittia basilobata. REFERENCES Andersen, T. & K. A. Johanson, (in press). Caddis flies (Trichoptera) from a mountain rain forest in NE Tanzania. — In: Otto, C. Proceedings of the 7th International Symposium on Trichoptera. — Universal Book Services, The Netherlands. Andersen, T. & O. A. Sæther, 1993. Lerheimia, a new genus of Orthocladiinae from Africa (Diptera, Chironomidae). — Spixiana 16: 105-112. Andersen, T. & O. A. Sæther, (in press a). Usambaromyia nigrala gen. n., sp. n., and Usambaromyiinae, a new sub- family among the Chironomidae (Diptera). — Aquatic Insects 15. Andersen, T. & O. A. Sæther, (in press b). Colosmittia clav- ata gen. n., sp. n., a new orthoclad from the Usambara Mts, Tanzania (Diptera: Chironomidae). — Journal of the Kansas Entomological Society 66. Brundin, L., 1947. Zur Kenntnis der schwedishen Chironomiden. — Arkiv for Zoologi 39A (3): 1-95. Brundin, L., 1956. Zur Systematik der Orthocladiinae (Dipt. Chironomidae). — Report, Institute of Freshwater Research, Drottningholm 37: 5-185. Caspers, N. & F. Reiss, 1989. Die Chironomidae der Türkei. Teil I: Podonominae, Diamesinae, Prodiamesinae, Orthocladiinae (Diptera, Nematocera, Chironomidae).— Entomofauna 10 (8/1): 105-160. Cranston, P. S. & D. R. Oliver, 1988. Additions and correc- tions to the Nearctic Orthocladiinae (Diptera: Chironomidae). — Canadian Entomologist 120: 425- 462. Cranston, P. S., D. R. Oliver, & O. A. Sæther, 1989. 9. The adult males of Orthocladiinae (Diptera: Chironomidae) of the Holarctic region - Keys and diagnoses. — In: Wiederholm, T. Chironomidae of the Holarctic region. Keys and diagnoses. Part 3. Adult males. — Entomologica scandinavica Supplement 34: 165-352. Iversen, S. T., 1991. The Usambara Mountains, NE Tanzania: Phytogeography of vascular plant flora. — Acta Universitatis Upsalensis, Symbolae Botanicae Upsaliensis 29 (3): 1-234. SÆTHER & ANDERSEN: Lobosmittia gen. n. Kieffer, J. J., 1926. Chironomiden der 2. Fram - Expedition (1898-1902). — Norsk entomologisk Tidsskrift 2: 78-89. Oliver, D. R., 1963. Entomological studies in the Lake Hazen Area, Ellesmere Island, including lists of species of Arachnida, Collembola and Insecta. — Arctic 16: 175- 180. Sæther, O. A., 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). — Bulletin of the Fisheries Research Board of Canada 107: 1-154. Sæther, O. A., 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Entomologica scandinavica Supplement 14: 1-51. Sether, O. A., J. E. Sublette & E. Willassen, 1984. Chironomidae (Diptera) from the 2nd Fram Expedition (1898-1902) to Arctic North America described by J. J. Kieffer. — Entomologica scandinavica 15: 245-275. Sæther, O. A. & X. Wang, 1992. Euryhapsis fuscipropes sp. n. from China and Tokyobrillia anderseni sp. n. from Tanzania, with a review of genera near /risobrillia Oliver (Diptera: Chironomidae). — Annales de limnologie 28: 209-223. Schlee, D., 1966. Präparation und Ermittlung von Messwerten an Chironomiden (Diptera). Gewässer und Abwässer 41/42: 169-193. Received: 6 January 1993 Accepted: 20 September 1993 287 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Cr ia: XX INTERNATIONAL CONGRESS SE. OF ENTOMOLOGY oe Florence (Italy), August 25 - 31, 1996 N INTENT FORM Please send it to: (ai) | O.I.C. igh Via A. La Marmora, 24 20121 Florence (I) N | no later than December 31, 1993 A : | FanulyanamegRgee ce e Reet NES MAME AAE ene, AA RR I AE LA GES tall ip Institution earste { PATES Shen on vib,” e. CHOY BING! OSA COGS Re EE ( Bj || COUNTY ENS EE IE en EER bun de: @}| © ram interested in the Congress, please send me further a circulars : ry CJ I will probably attend the Congress “| © Iwill probably submit a contribution on the following (à topic (please indicate the number referring to ® PIOPOSEISE CH OIS) RETE li Nr en EE i e @ 288 BERNHARD J. van VONDEL Natuurmuseum Rotterdam REVISION OE THE LAPATLUS SPECIES @F THE ORIENTAL REGION EXCLUDING CHINA (COREORTENA: HALIPEIDAE) Vondel, B. J. van, 1993. Revision of the Liaphlus species of the Oriental region excluding China (Coleoptera: Haliplidae). — Tijdschrift voor Entomologie 136: 289-316, figs 1-130. [ISSN 0040-7496]. Published 10 December 1993. The Oriental species of the Haliplidae are discussed. The species of the subgenus Liaphlus Guignot are revised, while the few remaining species of the subgenus Halipluss. str. (in revision by Holmen) and the genus Peltodytes Régimbart (already revised by Vondel 1992) are only dis- cussed in general. Twenty-six species of Haliplidae are known from the Oriental region. H. shanicus Guignot is established as a junior synonym of H. angustifrons Régimbart. Four new species are described: Haliplus javanicus, H. nedungaduensis, H. samosirensis and H. srilankanus. Most primary types have been studied. Lectotypes are designated for H. angustifrons Régimbart, H. oceanicus Régimbart, H. pulchellus Clark and H. pulchellus var. indicus Régimbart. The lat- ter is raised to specific rank. A check-list of all Oriental Haliplidae is provided, as well as a key to the Liaphlus species in the Oriental region excluding China, and distribution maps of most species. B. J. van Vondel, Roestuin 78, 3343 CV Hendrik-Ido-Ambacht, The Netherlands. Key words. — Haliplidae, Haliplus, Liaphlus, Oriental region, new species. Since Zimmermann (1924) published his key to the Haliplidae of the world, no revisional work has been done regarding the Oriental species. The status of the species described since is not always clear, so a revision is necessary. The type material of most spe- cies was studied, except for the three species described by Vazirani, whose types appear to be completely un- available for examination. Female genitalia are not described, because they usually are very uniform. Further research is necessary to check if there are characters sufficiently reliable to separate the species. MATERIAL About 280 specimens, including type-material, we- re studied from the following collections: Natural History Museum, London, UK (BMNH); B. P. Bishop Museum, Honolulu, Hawaii, USA (BPBM); Canadian National Collection, Ottawa, Canada (CNCI); Institut Royal des Sciences Naturelles, Brussels, Belgium (ISNB); Museum of Comparative Zoology, Cam- bridge, Massachusetts, USA (MCZC); Muséum Natio- nal d'Histoire Naturelle, Paris, France (MNHN); Museum of Zoology, Lund, Sweden (MZLU); Natur- historisches Museum Wien, Vienna, Austria (NHMV); Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRS); Nationaal Natuurhistorisch Museum (Rijks- museum van Natuurlijke Historie), Leiden, Nether- lands (RMNH); United States National Museum of Natural History, Washington D.C., USA (USNM); Instituut voor Taxonomische Zoölogie (Zoologisch Museum), Amsterdam, Netherlands (ZMAN); Zoolo- gisk Museum, Copenhagen, Denmark (ZMUC); Zoo- logische Staatssammlung, Munich, Germany (ZSMC). Private collections: L. Hendrich, Berlin, Germany; P. Mazzoldi, Brescia, Italy; B. J. van Vondel, Hen- drik-Ido-Ambacht, The Netherlands. METHODS Dissecting, preparing and drawing the aedaegi as well as the use of morphological terms follows the methods described by Vondel (1991: 76). Locality names are given, as far as possible, accord- ing to the Times Atlas of the World (Comprehensive edition 1983), but when different from original la- bels, the modern name is given in parentheses. Names of countries are used with their present-day boundar- ies. Distribution maps are based on material exam- ined. Literature records are included using different symbols. 289 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 SYSTEMATIC SECTION Checklist of Oriental Haliplidae, including Taiwan and China south of the river Chiang Jiang Haliplus Latreille subgenus Haliplus s.str. japonicus Sharp, 1873 (*) sauter: Zimmermann, 1924 (*) subgenus Liaphlus Guignot angustifrons Régimbart, 1892 shanicus Guignot, 1956 syn.n. arrowi Guignot, 1936 chinensis Falkenström, 1932 (**) diopus Guignot, 1955 diruptus Balfour-Browne, 1946 eximius Clark, 1863 modestus Zimmermann, 1924 emmerichi Falkenström, 1936 holmeni Vondel, 1991 (**) indicus Regimbart, 1899 stat. n. Javanicus sp. n. kapuri Vazirani, 1975 kotoshonis Kano & Kamiya, 1931 (**) manipurensis Vazirani, 1966 nedungaduensis sp. n. perroti Guignot, 1950 philippinus Chapin, 1930 pruthii Vazirani, 1966 pulchellus Clark, 1863 oceanicus Régimbart, 1886 samosirensis sp. n. sharpi Wehncke, 1880 (**) srilankanus sp. n. Peltodytes Régimbart coomani Peschet, 1923 (***) wui Gschwendtner, 1934 intermedius (Sharp, 1873) (***) sinensis (Hope, 1845) (***) variabilis (Clark, 1863) koreanus Takizawa, 1931 sumatrensis Régimbart, 1885 (***) The Oriental region as defined in this revision is according to fig. 122. Species of the subgenus Liaphlus, which are present in the south of China or in Taiwan and not further in the Oriental region (marked ** in the species-list) are already revised by Vondel (1991) and not included in the following key. The Oriental species of the genus Peltodytes (marked *** in the species-list) were revised together with the Palaearctic species (Vondel 1992). The few species of the subgenus Haliplus s. str. (marked with * in the species-list), which are present in the southwest of 290 China (A. japonicus Sharp) and in Taiwan (H. saute- ri Zimmermann) are subject of a revision of that sub- genus by Holmen (in prep.) and not further treated here. Some species are present both in the Orien- tal and Palaearctic regions. These species are included in the following key, but only treated here in a gener- al way, except H. pulchellus Clark, which is fully treat- ed. Key to the genera of Oriental Haliplidae 1. Elytra with sutural stripe on about apical half. Last segment of maxillair and labial palpi longer FAN PERL ATE SEP ments. E A CCnUS Pee Oder Pin bart - Elytra without sutural stripe or at most a weak trace at the apex. Last segments of maxillair and labial palpi much shorter than penultimate seg- TIDE AICS) IE Me genus Haliplus Latreille Remark. — Species of the genus Peltodytes are al- ready revised (Vondel 1992) and are only mentioned in the species-list. Key to the subgenera of Haliplus 1. Hind tibia with setiferous striole on dorsal face. Pronotum without basal plicae ....subg. Liaphlus — Hind tibia without setiferous striole on dorsal face. Pronotum with basal plicae … linate da ole subg. Haliplus s.str. Remarks. — Species of the subgenus Haliplus s.str. are in the Oriental region present in the southwest of China: Yunnan (7. japonicus) and in Taiwan (A. sauteri). The species concerned are only mentioned in the species-list, because they are subject of a revision by Holmen. Guignot (1955b) proposed a division of the subge- nus Liaphlus into species-groups. In some cases I do not agree with his interpretation of species. As far as Oriental species are concerned the following notes can be made. Haliplus diopus and H. pulchellus are closely related, but placed by Guignot in different groups. The latter, however, is placed by him with the clearly different H. testudo (from Australia) in one group. H. arrowi and H. angustifrons are closely related, but, again, placed by him in different groups. Key to the Oriental species of the subgenus Liaphlus: Species known only from China or Taiwan are not included in the key, but can be identified with Vondel (1991). VAN VONDEL: Oriental Haliplidae Figs. 1-9. Haliplus angustifrons, lectotype. — 1, dorsal view; 2, antenna; 3, punctures near elytral base and suture; 4, dorsal side of hind tibia; 5, prosternal process; 6, lateral view of prosternal process; 7, left paramere; 8, penis; 9, right paramere. 291 12. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Prosternal process with two longitudinal grooves (OL CR 2 Prosternal process flat or impressed in the middle (ie DA) n i 9 Éyiraimaculare (he 19) ren. 3 Elytra only narrowly darkened along suture (ie OS) ere Nae H.nedungaduensis Margins of elytra serrate in basal part (fig. 46) .4 Margins of elytra not clearly serrate in basal part Distance between eyes 0.9-1.0X width of one VE e specimens of H. pulchellus Distance between eyes 1.1-1.5X width of one eye Metasternal process with transverse impression, in which two pits can be recognised (fig. 50) RE dou meen EL SN ree H. indicus _Metasternal process with two separated pits (fig. DID ORE SIRIA E Di TA eee H. diopus Mesepimeron with about 4 strong punctures (fig. 92). Length less than 2.6 mm … … H. philippinus Mesepimeron without or rarely with one or two punctures. Length more than 2.6 mm Pronotum usually not continuously margined, sometimes traces of a margin in posterior part, posterior corners usually clearly protruding. Basal black band on elytra reaching fifth primary punc- ture-row (fig. 98) … … specimens of H. pulchellus Pronotum usually finely margined except near anterior corner, posterior corners not or hardly protruding. Basal dark band on elytra reaching at least sixth primary puncture row Elytral basal band brown, reaching margins when seen from above, black marks very irregular. Pronotum slightly darker in the middle (fig. 107). Length 3.3-3.5 mm SE I ia H. samosirensis Elytral basal band black, reaching sixth primary puncture-row, black marks well defined. Pronotum clearly darkened in the middle (fig. 55). Length 2.8-3.0 mm 10 Elytra without any markings Elytra with markings . Prosternal process broad and flat, narrowed beteen procoxae, not narrowed between meso- (elo) etal bite, Al) en nen. H. eximius Prosternal process narrowed between pro- and mesocoxae (see remarks below) ........... H. kapuri . Prosternal process without anterior margin (fig. Prosternal process pitted posteriorly (fig. 5) EE specimens of H. angustifrons Prosternal process canaliculate (see remarks be- 292 13. 16. 17 low ra aie an ae A N H. manipurensis Two longitudinal markings on disc of pronotum (see remarks below) nn H. pruthii At most small markings on disc of pronotum .. . Elytral markings not connected with darkened steuren ion E I I re H. perroti Elytral markings connected with darkened suture (fig. 1) . Prosternal process strongly grooved and with a deep pit in apical part (fig. 5), elytral base usually darkened Prosternal process flat or weakly impressed in ap- ical part (fig. 32), elytral base usually not dark- ened (HE IS) er H. diruptus Distance between eyes 0.8-0.9X width of one eye. Base of pronotum usually wider than elytral base, posterior corners rounded. Pronotum short, width at base 2.1-2.3 X length in the middle, strong and in basal corners coarse punctures. Margins thicker than elytral margin near shoul- ders. Basal dark band usually reaching to fifth puncture-row (fig. 1). Anterior edge of proster- num only margined near prosternal process. About 38 sutural punctures, about 32 punctures HV HITS CP EMATYÉTO WE H. angustifrons Distance between eyes 1.0-1.1X width of one eye. Width of pronotum at base about 2.0 X length in the middle, margins about as thick as elytral margins near shoulders . … … 17 Pronotal posterior corners clearly protruding (fig. 113). Anterior edge of prosternum completely margined. About 33 punctures in first primary row. Suture darkened to first primary puncture- row. (felt 3) i ml Ai H. srilankanus Pronotal posterior corners not protruding. Anterior edge of prosternum only margined near prosternal process. About 24 punctures in first primary row. Suture darkened to first secondary Pune (i 400) e H. arrowi Remarks. — The species described by Vazirani (AH. kapuri, H. manipurensis and H. pruthii) are only kn own to me by the original description. As I do not always know how to interpret some parts of these de- scriptions, especially concerning the shape of the prosternal process, some uncertainty remains regard- ing Vazirani s species. DESCRIPTION OF THE ORIENTAL LIAPHLUS Haliplus angustifrons Régimbart (figs. 1-9) Haliplus angustifrons Régimbart, 1892: 112. Lectotype d, ‘Konbir [Konbira, Ranchi distr., Bihar, India], P. VAN VONDEL: Oriental Haliplidae Figs. 10-18. Haliplus arrowi, holotype. — 10, dorsal view; 11, antenna; 12, punctures near elytral base and suture; 13, dorsal side of hind tibia; 14, prosternal process; 15, lateral view of prosternal process; 16, left paramere; 17, penis; 18, right para- mere. 293 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 294 Cardon; Museum Paris, Coll. Maurice Régimbart, 1908; type’ (MNHN) [examined]. Haliplus shanicus Guignot, 1956: 477. Holotype d, ‘S. Shan States, Road 40 km E. of Taunggyi, 25.IX-13.X.[19]34, Malaise’ (NHRS) [examined]. syn. n. Haliplus angustifrons, Régimbart 1899: 187, Zimmermann 1920: 303, 1924: 140, Guignot 1936: 117, 1954: 563, 1955b: 295, Vazirani 1966: 128. Haliplus shanicus, Vazirani 1966: 133. Diagnosis This species can be distinguished from the other species with a posteriorly well impressed prosternal process by the characters mentioned in the key. Description Length 2.9-3.8 mm, width 1.6-2.0 mm. Body oval, widest in the middle (fig. 1). Head: Yellow-brown to dark brown, vertex some- times darkened, moderately strongly punctured. Distance between eyes 0.8-1.0X width of one eye. Antennae yellow-red (fig. 2), palpi yellow-red. Pronotum: Yellow-red to rust-coloured. Lateral borders straight to slightly concave in posterior half and convex in anterior half, strongly margined. Posterior corners clearly protruding. Moderately strongly, in anterior corners and along base strongly punctured. Impressed along base. Elytra: Yellow-red to brown-red with extensive brown markings (fig. 1). Primary puncture-rows moderately strong, 25-33 punctures in first row. Secondary punctures relatively strong. All punctures darkened with yellow centre (fig. 3). Completely margined. Ventral side: Body yellow-brown to rust-coloured or brown, elytral epipleura yellow-brown, legs yellow- red to brown. Prosternal process strongly impressed in posterior part, anterior edge usually not clearly margined, strongly punctured, narrowed near coxae, anteriorly about as wide as posteriorly (fig. 5), lateral plicae complete (fig. 6). Metasternal process widely and strongly impressed in the middle, moderately strongly to weakly punctured (fig. 5). Metacoxal plates strongly punctured, along suture with a weakly punctured area. Setiferous striole on dorsal side of hind tibia about 15 X length of tibia, longest of the two tibial spurs about 23 X length of first tarsal seg- ment (fig. 4). Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on ventral side. Penis and parameres as in figs. 7-9. VAN VONDEL: Oriental Haliplidae Biology and distribution (fig. 123) In rivers. In mixed primary forest. In watertank. Up to an altitude of 1500 m. India: Armachal Pradesh, Bihar, Karnataka, Madhya Pradesh, Maharashtra, Uttar Pradesh, West Bengal; Vazirani (1966) also gives Himachal Pradesh and Orissa. Sri Lanka. Nepal. Burma. Material examined. — India: Armachal Pradesh: 1 ex., Assam, Patkai Mts, Doherty (BMNH); Bihar: 1d, Lectotype; Id, 12, Konbir [Konbira], P. Cardon, paralectotype (MNHN); 16, 12, Konbir [Konbira, Ranchi distr.], P. Cardon, Haliplus pulchellus Régimbart det. 1890 (ISNB); 42 ex., Chota-Nagpore [Chota Nagpur], Nowatoli, R. P. Cardon, viii-ix.1896, ix-x.1896, iv-v.1897 (MNHN, RMNH); Karnataka: 18, 19, Shimoga, Maissour, vi-vii.1897 (MNHN); Madhya Pradesh: 1 ex., Barway [Barwah?], P. Cardon (BMNH); Maharashtra: 1 ex., Lonaula, 80 km E. Bombay, 13.ix.1991, R. Schuh (NHMV); Uttar Pradesh: 16, O, Almora Dn., Kumaun, vii.1920, H.G.C (BMNH); West Bengal: 12, Sarda, F.W.C. (BMNH); no province known: 1d, Indes Or., 22-53 (MNHN). — Nepal: 22, Kathmandu Valley, 7.u.1981, M. Jäch (NHMV, Vondel); 29, Kathmandu Distr., Gokarna, 1.viii.1983, M.G. Allen, 4500’, mixed primary forest (BMNH); 12, Ktmd [Kathmandu], Godavari [?], 15.viii.1967, 5000’; 2d, 29, nr Birganj, Lothar, 12.ix.1967, 450 Ft. (CNCI); 8 ex., Godawari, 15 km S. Kathmandu, 1500 m, 29.iv.1984, Wewalka (Mazzoldi). — Sri Lanka: 1d, Inginiyagala, 14.x11.1979, V. Mahler Jensen (ZMUC). — Burma: Holotype 3, H. shanicus, 1% allotype, 16 paratype, data as holotype (NHRS); paratypes of H. shanicus, 16, 29, data as holotype (MNHN); 1d, 19, Shan States, road 40 km E. of Taung-gyi, 25.ix-13.x.1934; 16, Shan States, Namhkok [?], 20. viii.1934, Malaise, 900 m.; 1d, Shan States, Taung-gyi, 25.x.1934, Malaise, 1500 m, watertank (MNHN). Haliplus arrowi Guignot (figs. 10-18) Haliplus arrowi Guignot, 1936: 117. — Holotype d, “Type, 3, Calcutta, Atkinson 92-3., Det. Dr. Guignot Haliplus Arrowi Guign., Type’ (BMNH) [examined]. Haliplus arrow, Guignot 1955b: 294, Vazirani 1966: 131, 1975: 317. Diagnosis This species is very closely related to Haliplus an- gustifrons, but has sparser elytral puncture-rows, the eyes less close to each other, the posterior corners of the pronotum less protruding and some other slight differences, mentioned in the key. Description Length 2.8-3.1 mm, width 1.5-1.6 mm. Body oval, widest in the middle (fig. 10). Figs. 19-27. Haliplus diopus, holotype. — 19, dorsal view; 22, antenna; 21, punctures near elytral base and suture; 22, dorsal side of hind tibia; 23, prosternal process; 24, lateral view of prosternal process; 25, left paramere; 26, penis; 27, right para- mere. 295 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 30 een . © © : bo 296 Head: Brown, moderately punctured. Distance between eyes 1.1-1.2 X width of one eye. Antenna (fig. 11) and palpi yellow to yellow-brown. Pronotum: Yellow-brown. Lateral borders convex, very finely margined, posterior corners not protrud- ing. Width at base about 2.0 X length in the middle. Fairly strongly, along the base very strongly and densely punctured, punctures not darkened. In the midddle a transverse somewhat elevated unpunctured band. Elytra: Yellow-brown, brown marking along suture and part of base and on disc, marks in even intervals (fig. 10). Primary puncture-rows sparse and moder- ately strong, about 25 punctures in first row. Secondary punctures moderately strong along suture and rather weak in intervals. All punctures darkened with a yellow centre. Ventral side: Yellow-brown to brown, elytral epi- pleura yellow, legs yellow-brown, femora brown to dark-brown near coxae. Prosternal process strongly impressed in posterior part and less impressed in an- terior part, slightly narrowed near coxae, moderately strongly and densely punctured (fig. 14), lateral plicae complete (fig. 15). Anterior border of prosternum not completely margined, margin near prosternal process accompanied by four punctures in a row. Metasternal process with a wide and very deep pit, weakly punc- tured (fig. 14). Coxal plates densely punctured, ex- cept near suture, posterior part of plates dilated. Setiferous striole on dorsal side of hind tibia about 1/3 X length of tibia and with about 8 punctures, longest of the two tibial spurs almost as long as first tarsal segment (fig. 13). Male: First three tarsal segments of fore and mid- legs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 16-18. Biology and distribution (fig. 124) Biology unknown. India: Bihar, Madras, Uttar Pradesh, West Bengal. Burma. Laos. Material examined. — India: Bihar: 1d, Lectotype (BMNH); 2d, Calcutta, Atkinson, paratype (BMNH, MNHN); 2 ex., Pusa, Andrewes (BMNH); Uttar Pradesh: 1 ex. Sirampur [Sitapur?], Elberling (ZMUC); West Bengal: 16, Calcutta, Haliplus pulchellus det. dr Guignot (BMNH). — Burma: 1 ex., Maymyo, H.I. Andrewes (BMNH). — Laos: 1 ex., betw. Vientiane and Luang Prabang, end of 1919, R. V. de Salvara (BMNH). — Unlabelled: 3 ex. (BMNH). van VONDEL: Oriental Haliplidae Haliplus chinensis Falkenstròm Haliplus chinensis Falkenström, 1932: 191. Fully redescribed by Vondel (1991). In the Oriental region only known from China. Haliplus diopus Guignot (figs. 19-27) Haliplus diopus Guignot, 1955a: 270. Holotype d, ‘Tonkin, Hoa-Binh [Vietnam], VII-1940 (Coom.)’ [A. de Cooman] (MNHN) [examined]. Haliplus diopus, Guignot 1955b: 294. Diagnosis This species can be distinguished from H. pulchel- lus by the wider distance between the eyes and from H. indicus by the clearly separated pits on the metas- ternal process. Description Length 2.8-3.1 mm, width 1.6-1.7 mm. Body oval, widest in or just before the middle (fig. 19). Head: Brown, sparsely and weakly punctured, between eyes partly unpunctured. Distance between eyes 1.1 X width of one eye. Antennae yellow to yel- low-red (fig. 20), palpi yellow-red. Pronotum: Yellow to yellow-red, longitudinal dark blotch along the middle, this blotch can be vague or reduced to a blotch on anterior part (fig. 19). Lateral borders slightly convex, concave just behind anterior corners, weak margins not allways clearly visible be- cause of coarse punctures. Strongly, on the disc more weakly punctured, widened punctures in rounded ap- ical corners (fig. 19). Usually transverse impression along base. Width at base 1.8-2.1 length in the middle and 1.6-1.8 X width in front. Elytra: Yellow with an extensive dark marking along suture and along base to fifth puncture-row, central mark connected to small mark in third inter- val, a various number of irregular marks spread over elytra (fig. 19). Rather strong primary puncture-rows, 26-30 punctures in first row. Secondary punctures fairly strong, accompanied by some very small punc- tures. All punctures darkened, but distinctly yellow in centre of punctures (fig. 21). Margins of elytra slight- ly serrate in basal part (fig. 19). Ventral side: Yellow-red to rust-coloured, elytral epipleura yellow-red, legs red-brown, darkened to- wards coxae. Prosternal process strongly grooved along margins, slightly narrowed near coxae, strongly Figs. 28-36. Haliplus diruptus, holotype. — 28, dorsal view; 29, antenna; 30, punctures near elytral base and suture; 31, dor- sal side of hind tibia; 32, prosternal process; 33, lateral view of prosternal process; 34, left paramere; 35, penis; 36, right par- amere. 297 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 © © © ® e © ‚© € de ce © © ve ju ce © 9 © © SOS eos Ee © 8 0 © © e 0° eo ee © 0. et ee" Siero Ie ele e @ 0 ve ® ee eee 02 . 9 © © 0 0 06060 es % © ° cor © 298 punctured in both grooves, clearly margined anterior- ly (fig. 23), lateral plicae complete (fig. 24). Metaster- nal process flat with strong pit on both sides of the middle, between pits not or hardly impressed, sparse- ly, laterally densely punctured (fig. 23). Metacoxal plates strongly, along suture weakly and sparsely punc- tured, punctures without a hair, apical sutural corner slightly margined. Fifth and sixth abdominal segment with complete row of strong punctures. Setiferous stri- ole on dorsal side of hind tibia very short, about 1/8 X length of tibia, longest of two tibial spurs about 4/5 X length of first tarsal segment (fig. 22). Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on ventral side. Penis and parameres as in figs. 25-27. Biology and distribution (fig. 128) Biology unknown. Vietnam. Material examined. — Vietnam: Holotype d, 2 allotype, 4 d paratypes, 69 paratypes, Tonkin, Hoa Binh, viii.1940 & viii.1941, A. de Cooman, Haliplus diopus n. sp. dr. F. Guignot det. (ISNB, MNHN). Haliplus diruptus Balfour-Browne (figs. 28-36) [Haliplus simplex sensu Régimbart 1899, nec Clark 1863. Misidentification]. Haliplus diruptus Balfour-Browne, 1946: 436. This species was fully redescribed by Vondel (991095) Probably also in India: see under unnamed material. Guignot records this species from Burma (1954: 563), but later (1956) changed his view and based the new H. shanicus on that material. Additional material exam- ined: 12, SW China, Yunnan, Western Hills nr. Kunming, 2100 Mt., 7.viii.1940, J. L. Gressitt (BPBM). Haliplus eximius Clark (fig. 37-45) Haliplus eximius Clark, 1863: 418. Haliplus modestus Zimmermann, 1924: 139. Haliplus hiogoensis Kano & Kamiya, 1931: 1. Haliplus emmerichi Falkenstrôm, 1936: 79. This species was fully redescribed by Vondel (1991: 97), but erroneously treated under the name H. eximis. Additional material examined: China: 2 ex. VAN VONDEL: Oriental Haliplidae Chusan Isl, J. J. Walker (BMNH); 1 ex., Fukien, Yungan, 5.ix.1940, T. C. Man (BPBM). Haliplus holmeni Vondel Haliplus holmeni Vondel, 1991: 109. This species is only known from China: Yunnan. Haliplus indicus Régimbart stat. n. (figs. 46-54) Haliplus pulchellus var. indicus Régimbart, 1899: 189. Lectotype d (here designated), ‘Chota-Nagpore [India, Ranchi Distr., Bihar], Nowatoli, R. P. Cardon, viii- ix.1896, pulchellus var. indicus Rég., Dr. Régimbart vidit 1898, Museum Paris, ex coll. R. Oberthur’ (MNHN) [ex- amined]. Haliplus pulchellus var. indicus, Zimmermann 1920: 314. Haliplus pulchellus ab. indicus, Zimmermann 1924: 140. Haliplus pulchellus indicus, Vazirani 1966: 133. Diagnosis This species, belonging to the group with double grooved prosternal process, can be distinguished from H. pulchellus by the wider distance between the eyes. It is here considered a distinct species. Description Length 2.8-3.1 mm, width 1.6-1.7 mm. Body oval, widest in or just before the middle (fig. 46). Head: Brown, sparsely and weakly punctured, between eyes partly unpunctured. Distance between eyes 1.1-1.2 X width of one eye. Antennae yellow to yellow-red (fig. 47), palpi yellow-red. Pronotum: Yellow to yellow-red, dark mark in an- terior middle, vague central mark before base can be reduced or absent (fig. 46). Lateral borders slightly convex, concave just behind anterior corners, weak margins usually not clearly visible because of coarse punctures. Strongly punctured, on disc more weakly so, with widened punctures in rounded posterior cor- ners. Usually a transverse impression along base op- posite fourth to sixth elytral puncture-row. Elytra: Yellow with an extensive dark marking along suture and along base to fourth or fifth punc- ture-row, central mark connected with small mark in fourth interval, a variable number of irregular marks spread over elytra (fig. 46). Rather strong to strong primary puncture-rows, 26-35 punctures in first row. Secondary punctures fairly strong. All punctures darkened, but distinctly yellow in centre of punctures (fig. 48). Margins of elytra slightly serrate in basal part (fig. 46). Figs. 37-45. Haliplus eximius, China. — 37, dorsal view; 38, antenna; 39, punctures near elytral base and suture; 40, dorsal side of hind tibia; 41, prosternal process; 42, lateral view of prosternal process; 43, left paramere; 44, penis; 45, right paramere. 299 ‘TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 er‘ ; 2 ima ces a ne — 300 Ventral side: Yellow-red to rust-coloured, elytral epipleura yellow-red, legs red-brown, darkened to- wards coxae. Prosternal process strongly grooved along margins, slightly narrowed near coxae, strongly punctured in both grooves, clearly margined along anterior edge (fig. 50), lateral plicae complete (fig. 51). Metasternal process with strong transverse impres- sion, in which two pits can be recognised, sparsely punctured (fig. 50). Metacoxal plates strongly, along suture weakly and sparsely punctured. Fifth and sixth sternite with complete transverse row of strong punc- tures, last sternite strongly and densely punctured. Setiferous striole on dorsal side of hind tibia about 1/5 X length of tibia, longest of two tibial spurs about 4/5 X length of first tarsal segment (fig. 49). Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on, ventral side. Penis and parameres as in figs. 52-54. Biology and distribution (fig. 127) Biology unknown. India: Bihar, West Bengal, Tamil Nadu. Material examined. — India: 4d, 29, Chota Nagpore [Ranchi distr. Bihar], Nowatoli, R.P. Cardon, viii-ix.1896 (MNHN, RMNH); 5d, 49, W. Bengal, Santiniketan, Birbhum District, x.1937, T. C. Maa; 19, Madras State, Coimbatore, 420 m, iv.1962, P. S. Nathan (BPBM). Haliplus javanicus sp. n. (fig. 55-63) Type material. — Holotype d, ‘Tjilatjap, Java, Drescher, 12.1915; coll. F. C. Drescher’ (ZMAN). — Paratypes, 2d: 1d, data as holotype (coll. Vondel); 16, ‘Java, 1915, Weltevreden, Neytzell de Wilde; coll. Dr. D. MacGillavry (ZMAN). Diagnosis This species can be distinguished from other spe- cies with a double grooved prosternal process by the combination of the central pronotal blotch, the smooth elytral margin near shoulders and the elytral basal darkening reaching sixth puncture-row. Description Length 2.8-3.0 mm, width 1.5-1.7 mm. Body oval, widest in the middle (fig. 55). Head: Brown to dark brown, moderately strongly, on vertex stronger punctured. Distance between eyes 0.7-1.0 X width of one eye. Antennae yellow-red (fig. 56), palpi yellow-red. VAN VONDEL: Oriental Haliplidae Pronotum: Yellow to yellow-brown with large cen- tral longitudinal mark. Lateral borders straight and finely margined, near anterior corner concave and not margined. In posterior corners strongly and densely punctured, basally strongly punctured, on the disc more weakly and sparsely punctured. Impressed op- posite fourth to sixth elytral puncture-rows (fig. 55). Elytra: Yellow to yellow-red, dark marks along su- ture and along base to sixth puncture-row, various marks on intervals (fig. 55). Primary puncture-rows moderately strong, about 28 punctures in first row. Secondary punctures strong. All punctures darkened with yellow centre (fig. 57). Ventral side: Body yellow-red to brown, elytral epi- pleura yellow-red, legs yellow-red to brown-red. Prosternal process with two strong grooves, margined at anterior edge, narrowed near coxae, moderately strongly and sparsely punctured (fig. 59), lateral pli- cae complete (fig. 60). Metasternal process with two strong pits, weakly punctured (fig. 59). Metacoxal plates strongly punctured, along suture weakly punc- tured. Setiferous striole on dorsal side of hind tibia very short, consisting of about 4 small punctures, longest of the two tibial spurs about 2/3 X length of first tarsal segment (fig. 58). Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on ventral side. Penis and parameres as in figs. 61-63. Biology and distribution (fig. 130) Biology unknown. Indonesia: Java. Haliplus kapuri Vazirani (fig. 64-65) Haliplus kapuri Vazirani, 1975: 317. Holotytpe d, India, Maharashtra, Fritzgerald, Satara District, 5.x.1972, A. P. Kapur (in Zoological Survey of India, Calcutta)[not ex- amined] Original description Length 3.5 mm. Head testaceus; punctation moderate and irregular, stronger on the vertex; antennal segments 3 and 4 rather thinner than others, 7th segment slightly long- er than 8th segment. Pronotum testaceus; punctation larger than on the head, larger and stronger anteriorly and posteriorly, less strong on the disc, generally irregular, two longi- tudinal rows of fine punctures discernible on the disc; Figs. 46-54. Haliplus indicus, lectotype. — 46, dorsal view; 47, antenna; 48, punctures near elytral base and suture; 49, dorsal side of hind tibia; 50, prosternal process; 51, lateral view of prosternal process; 52, left paramere; 53, penis; 54, right para- mere. 301 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 55-63. Haliplus javanicus, holotype. — 55, dorsal view; 56, antenna; 57, punctures near elytral base and suture; 58, dor- sal side of hind tibia; 59, prosternal process; 60, lateral view of prosternal process; 61, left paramere; 62, penis; 63, right par- amere. 302 65 VAN VONDEL: Oriental Haliplidae 66 67 Figs. 64-65. Haliplus kapuri, (after Vazirani, 1975). — 64, ventral view of thorax; 65, penis. Figs. 66. Haliplus manipurensis, penis (after Vazirani 1966). Figs. 67. Haliplus pruthii, penis (after Vazirani 1966). lateral margin of the pronotum rebordered. Elytra pale testaceus without any markings; punc- tation quite regular except in the apical portion, both strial and interstrial punctures black; row of sutural punctures moderate, less strong but more dense than the strial punctures, interstrial punctures less strong than the strial punctures but not more numerous than them. Ventral side testaceus with legs ferruginous, meso- sternum darker, metacoxal plates and epipleurae pal- er. Prosternal process not rebordered laterally but canaliculate, strongly punctate more so on the lateral borders, a little narrowed in between the procoxae and the mesocoxae (fig. 64), anteriorly bordered by a fine ridge; anterior part of the metasternum (behind the prosternal process) distinctly swollen/inflated, with a large median pit behind it and two smaller pits on the anterolateral corner; punctation on the meta- sternum irregular except a transverse row of strong punctures behind the large median pit and a row of fine punctures along the posterior border of the met- asternum; metacoxal plates moderately punctate, punctures irregular and separated by 2-3 X its own diameter. Penis (fig. 65) rather strongly curved, rounded at the apex and narrowed in the middle. Biology and distribution (fig. 129) Biology unknown. Only known from the type-locality. Remarks. — As I have been unable to borrow the type for examination I only know this species from the original description. According to Vazirani it comes close to H. arrowi. Haliplus kotoshonis Kano & Kamiya Haliplus kotoshonis Kano & Kamiya, 1931: 2. This species is fully redescribed by Vondel (1991: 113). Probably in Laos: see under unnamed material. Additional notes: Satô (1984) recorded this species from the Ryukyu Islands. Additional material examined. — 19, Ryukyu Isl., Ishigaki I., 20-30.x1.1952, G.E. Bekart (BPBM). Haliplus manipurensis Vazirani (fig. 66) Haliplus manipurensis Vazirani, 1966: 132. Holotype d, [India], Manipur, Imphal, 880 m, 24°44’N, 93°58’E, 8.vii.1960, F. Schmid coll. (in Zoological Survey of India) [not examined]. Original description Length 3.2-3.5 mm. Form, oval, feebly attenuated posteriorly. Head testaceus, finely and moderately punctate, punctures on the vertex separated by their own diam- 303 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 È di OO OCs O5 e e ose CE o ee: ss ui Pe ee og x “oee : °° a ed ceo e o ee 5 © e © Le = .o avo "o Do . Figs. 68-76. Haliplus nedungaduensis, holotype. — 68, dorsal view; 69, antenna; 70, punctures near elytral base and suture; 71, dorsal side of hind tibia; 72, prosternal process; 73, lateral view of prosternal process; 74, left paramere; 75, penis; 76, right paramere. Figs. 77-86. Haliplus perroti, holotype. — 77, dorsal view; 78, antenna; 79, punctures near elytral base and suture; 80, dorsal side of hind tibia; 81, prosternal process; 82, lateral view of prosternal process; 83, metacoxal plate; 84, left paramere; 85, pe- nis; 86, right paramere. 304 VAN VONDEL: Oriental Haliplidae ee 78 \ ur DE a je e 5 79 | bia a | A ur of i 200 0 0 © 6 66 0 e o 8 9 ee » ®® DE MD e co RISO, © DH thea ° À © Oee » ee © dis ere oo 0° 5 305 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 87-97. Haliplus philippinus (87, 89-91, 93, 95-97, holotype; 88, 92, 94, Laos). — 87, dorsal view; 88, antenna; 89, punc- tures near elytral base and suture; 90, dorsal side of hind tibia; 91, prosternal process; 92, mesepimeron; 93, lateral view of prosternal process; 94, head and pronotum; 95, left paramere; 96, penis; 97, right paramere. 306 eter. Antennal segments 7-11 subequal and broader than the sixth. Pronotum testaceous, paler towards sides; puncta- tion moderate and irregular, stronger along anterior and posterior margins, sparse on the disc; a few punc- tures on postero-lateral margins strong and confluent. Anterior angles not projecting beyond the middle of anterior margin. Sides moderately rebordered, slight- ly convex, posterior angles slightly prominent, not projecting beyond the thoraco-elytral angles. Elytra testaceous with irregular ferruginous mark- ings, more so along the suture and extending laterally at base. Sutural line of punctation irregular, not so closely punctate; punctures separated by twice their own diameter. Strial punctures moderate, shallow, black; on the disc quite close and not separated by their own diameter. Each interstrial space with a row of much smaller and deeper, but not more numerous black punctures. Ventral side testaceous, epipleurae paler. Prosternal process and legs darker. Prosternal process margined laterally but not anteriorly, canaliculate, slightly nar- rowed adjacent to the front coxae; anterior end a little wider than apex; lateral margins punctate but not roughened, punctures fine, separated by more than their own diameter. Apical part of metasternum al- most flat, pitted, moderately but irregularly punctate. Metacoxal plates with moderate, shallow and dense punctures, punctures separated by half of their own diameter. Male with anterior tarsi moderately broadened and furnished with ‘ciliae’ underneath; claws broken. Penis very slightly curved, rounded at the apex (fig. 66). Biology ne inbucon (fig. 129) Biology unknown. Only known from the type-locality. Remarks. — As I have been unable to borrow the type for examination I only know this species from the original description. Haliplus nedungaduensis sp. n. (figs. 68-76) Type material. — Holotype à , ‘South India, Nedungadu, 7.1934, Hathach [?]’ (MNHN). Diagnosis This species can be distinguished from the other species with double grooved prosternal process by the absence of elytral markings. Description Length 2.7 mm, width 1.5 mm. Body oval, widest in the middle (fig. 68). VAN VONDEL: Oriental Haliplidae Head: Yellow-brown, vertex slightly darkened, rather weakly punctured, unpunctured band between the eyes not always present. Distance between the eyes 1.2-1.4 the width of one eye. Antennae yel- low-red (fig. 69), palpi yellow-red. Pronotum: Yellow to yellow-red. Lateral borders about straight, fine margins a little undulate. Modera- tely strongly and sparsely punctured. Base opposite fourth to sixth elytral puncture-row impressed and strongly punctured (fig. 68). Elytra: Yellow to yellow-red, suture slightly dark- ened to secondary punctures (fig. 68). Completely margined, margin serrate in anterior part. Primary puncture-rows fairly weak, about 29 punctures in first row. Secondary punctures strong. All punctures darkened with a yellow centre (fig. 70). Ventral side: Body yellow to yellow-red, elytral epi- pleura yellow, legs yellow-red to brown-red, slightly darkened towards coxae. Prosternal process with two weak grooves, margined at anterior edge, narrowed near coxae, anteriorly wider than posteriorly, moder- ately strongly punctured (fig. 72), lateral plicae com- plete (fig. 73). Metasternal process with deep trans- verse impression in which two pits are recognisable, weakly punctured (fig. 72). Metacoxal plates moder- ately strongly punctured, along suture with a weakly punctured area. Setiferous striole on dorsal side of hind tibia about 1/3 X length of tibia, longest of the two tibial spurs about 2/3 X length of first tarsal seg- ment (fig. 71). | Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on ventral side. Penis and parameres as in figs. 74-76. Biology and distribution (fig. 127) Biology unknown. Only known from the type-locality. Haliplus perroti Guignot (figs. 77-86) Haliplus perroti Guignot, 1950: DI Holotype 6, “Tam Dao, Tonkin, H. Perrot, 6, Type’ (MNHN) [examined]. Diagnosis This species can be recognised by the longitudinal elytral marks, which are not connected to the darke- ned suture. Description Length 3.2-3.4 mm, width 1.7-1.9 mm. Body oblong oval, widest in the middle (fig. 77). Head: Yellow-brown, vertex slightly darkened, rather weakly punctured, unpunctured band between eyes not always present. Distance between eyes 1.2- 1.4X width of one eye. Antennae yellow-red (fig. 307 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 78), palpi yellow-red. Pronotum: Yellow with two small marks on disc before base. Lateral borders convex, finely margined. In anterior corners strongly and densely punctured, basally with a continous row of darkened punctures getting strong towards sides, on disc more weakly and sparsely punctured. Slightly impressed in front of central base (fig. 77). Elytra: Yellow to yellow-red, suture completely darkened to secondary punctures, oblong marks on even intervals (fig. 77). Primary puncture-rows fairly weak, 31-35 punctures in first row. Secondary punc- tures weak. All punctures darkened with yellow centre (fig. 79). Ventral side: Body yellow-red to yellow-brown, elytral epipleura yellow, legs brown-red, darkened to- wards coxae. Prosternal process impressed in the mid- dle, less impressed at apex, margined at anterior edge, narrowed near coxae, moderately strongly punctured (fig. 81), lateral plicae complete (fig. 82). Metasternal process widely and strongly impressed in the middle, weakly punctured (fig. 81). Metacoxal plates densely and rather strongly punctured, along suture with a wide weakly punctured area, slightly impressed to- wards apical point (fig. 83). Setiferous striole on dor- sal side of hind tibia about 1/3 X length of tibia, con- sisting of about 12 small groovelike connected punctures, longest of the two tibial spurs about 2/3 X length of first tarsal segment (fig. 80). Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on ventral side. Penis and parameres as in figs. 84-86. Biology and distribution (fig. 125) Biology unknown. Vietnam. India: West Bengal, Rajasthan. Material examined. — Vietnam: Holotype d; 19, allo- type, 2 d, paratype, Tam Dao, Tonkin, H. Perrot (MNHN). — India: 1 ex., Pusa, Bengal, R. D. D., xi.1908 (BMNH); 19, Rajasthan, Sariaka, 16.viii.1989, A. Riedel (Hendrich). Haliplus philippinus Chapin (figs. 87-97) Haliplus philippinus Chapin, 1930: 9. Holotype d, ‘Bacoor Nov PI, P. L. Stangl Collector, Type No. 41757 U.S.N.M., Haliplus philippinus Type Chap.’ (USNM) [examined]. Diagnosis This species can be distinguished from the related H. pulchellus by the four punctures on the mesepimeron. Description Length 2.4-2.7 mm, width 1.3-1.5 mm. Body broad, more or less parallel in basal half of elytra (fig. 87). Head: Brown-red, margins dark, weakly and sparsely punctured. Usually with dark arrowlike mark (fig. 94). Distance between eyes about 1.2 X width of one eye. Antennae yellow-red (fig. 88), palpi yellow- red. Pronotum: Yellow-red to yellow in lateral parts. Usually wich dark central mark (fig. 94). Lateral bor- ders slightly convex, not margined. Posterior corners broadly rounded, anterior corners bent outwards. Anterior margin clearly protruding in the middle. Rather weakly punctured, two rows of strong punc- tures along the slightly impressed base, coarse punc- tures together forming a strong impression in each hind corner (fig. 87). A Elytra: Yellow-red, brown markings along base and suture and a clear mark in posterior half between fourth and sixth puncture-row, often additional marks on intervals (fig. 87). Primary punctures mod- erately strong, standing in irregular sometimes diffi- _ cult to recognise rows. About 28 punctures in first row. Secondary punctures relatively strong, some- times nearly as strong as primary punctures. All punc- tures slightly darkened with a yellow centre (fig. 89). Ventral side: Body brown-red with several strongly darkened parts, elytral epipleura yellow-red, legs yel- low-red to brown-red. Mesepimeron with about 4 strong punctures (fig. 92). Prosternal process parallel, slightly narrowed near coxae, sparsely punctured, margins ridgelike, about flat in the middle (fig. 91). Metasternal process flat, small clear pit on each side, sparsely punctured. Metacoxal plates strongly, near suture very sparsely and weakly punctured, apical cor- ner of suture sharp. Last sternite largely covered with strong punctures, fifth and sixth sternite with a con- tinuous row of strong punctures. Setiferous striole on dorsal face of hind tibia about % X length of tibia, longest of tibial spurs about 4 X length of first tarsal segment (fig. 90). Male: First three segments of front and midlegs slightly widened, scaly hairs on ventral side. Claws of forelegs equal in shape and length. Penis and para- meres as in figs. 95-97. Biology and distribution (fig. 128) Attracted to light. Up to an altitude of 920 m. Philippines. Indonesia: Java. Thailand. Laos. Vietnam. Burma. Figs. 98-106. Haliplus pulchellus, lectotype. — 98, dorsal view; 99, antenna; 100, punctures near elytral base and suture; 101, dorsal side of hind tibia; 102, prosternal process; 103, lateral view of prosternal process; 104, left paramere; 105, penis; 106, right paramere. 308 van VONDEL: Oriental Haliplidae 309 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Material examined. — Philippines: Holotype d (USNM). — Vietnam: 19, Cochin China, Travinh, Travinh Prov, vii.1932, K.M. Tsui (BPBM). — Laos: 2d, Vientiane, 31.v- 3.vi.1960, S. Quale & L. Quale, light trap; 18, Muong Sing, NW of Luang Prabang, 6-10.vi.1960, L.W. Quale, light trap, 650 m.; 2 ex, 39 km S. Djiring, 810 m, 29.iv.1969, J.W. Quale; 2 ex., Di Linh, Djiring, 920 m, light trap, 22-28.iv.1960, S. & J. Quale; 1 ex., Cochin China, French Indo China, Travinh, Travinh Province, vii.1932, K. H. Tsui (BPBM). — Thailand: 2d, 19, Bangkok (BMNH, Vondel); 2 ex., Bangkok, Aynttaya, 28.viii.1988, S. Schödl (NHMV, Vondel); 2 ex., Bangkok (MNHN); 2 ex., 25 km NW Lan Sak, 210 km NW Bangkok, viii.1991, W. Thielen (RMNH). — Burma: 1 2, Victoria Point, Tenasserim, viii. 1887, E.T. Atkinson (BMNH); 8 ex., Rangun, 16.v.1976, Wewalka (Mazzoldi). — Indonesia: 16, Batavia [Jakarta], ii.1949, C. v. Nidek (ZMAN). Haliplus pruthii Vazirani (fig. 67) Haliplus pruthii Vazirani, 1966: 130. Holotype d, Madhya Pradesh (Narmada Survey); Rewa State (=Rewa District), Khetagao, stn. 45, c 880 m, 24°31N, 81°19E; Mar.1927, H.S. Prithi coll. (in: Zoological Survey of India) [not examined]. Diagnosis The character of the two longitudinal marks on pronotum seems to offer a possibility to distinguish this species from related species. Original description Length 3.0-3.2 mm. Head testaceus; punctation on the vertex fine, moderately close; antennal segments 7-11 subequal, segment 7 not significantly broader than preceding segment. Pronotum testaceus, with two ferruginous longitu- dinal parallel lines on the disc coinciding with punc- tures; punctation irregular, moderate, rather effaced on the disc (excepting those coinciding with the fer- ruginous markings); sides narrowly rebordered, a lit- tle oblique, very slightly convex; posterior angles ob- tuse. Anterior angles almost in line with middle of the anterior margin. Elytra testaceus, with irregular and indeterminate ferruginous markings, more prominent along the su- ture and at base. Sutural line of punctures regular, punctures almost separated by their own diameter, more numerous than the strial punctures, Strial punc- tures moderate, rather close and shallow, narrowly black. Each interstrial space with a row of small deep punctures not more than the strial punctures. Ventral side testaceus; sternum and legs ferrugi- nous. Prosternal process moderately margined lateral- ly, narrowly margined anteriorly, moderately nar- rowed adjacent to the front coxae, anterior end and apical end subequal, lateral margins punctate, median 310 area hardly inflated, punctation indistinct. Apical part of metasternum slightly inflated and pitted. Metasternum weakly and sparsely punctate. Abdominal sternites with a single transverse row of punctures. Apical abdominal sternite with a few pos- tero-lateral punctures (less than in angustifrons or ar- rowi). Metacoxal plates more or less regular and fine- ly punctate, separated by two or three times their own diameter. Male with protarsi moderately dilated and fur- nished with cilia underneath. Protarsal claws thin, slightly curved and more than half the apical seg- ments length. Penis (fig. 67) of paratype moderately curved. Biology and distribution (fig. 129) Up to an altitude of 880 m. Probably attracted to light. India: Madhya Pradesh, probably also in Uttar Pradesh and Bihar (see under unnamed material). Remarks. — As I have been unable to borrow the type for examination I only know this species from the original description. Haliplus pulchellus Clark (fig. 98-106) Haliplus pulchellus Clark, 1863: 418. Lectotype d (here des- ignated), ‘Browning, Siam [Thailand], Co-type’ (BMNH) [examined]. Haliplus oceanicus Régimbart, 1886: 139. Lectotype (here designated), ‘Ed. Everts, W. Dates, Medan, Sumatra; Haliplus oceanicus Régb., type; Type; Museum Leiden, Haliplus pulchellus det. Clark (RMNH) [examined]. Haliplus pulchellus, Régimbart 1889: 147, 1892: 112, 1899: 188, Zimmerman 1920: 314, 1924: 140, 1927: 2, Guignot 1955b: 295, Vazirani 1966: 133. Haliplus oceanicus, Zimmermann 1927: 3. Haliplus pulchellus var. oceanicus, Régimbart 1899: 189, Zimmermann 1920: 314, 1924: 140. Diagnosis This species can be distinguished from the other species with a double grooved prosternal process by the characters mentioned in the key. Description Length 2.5-3.4 mm, width 1.5-2.0 mm. Body oval, widest just before the middle (fig. 98). Head: Brown to dark brown, darkened along the eyes and near antennae, sometimes with-a weak ar- row-like darkening in the middle. Sparsely and weak- ly, behind eyes stronger punctured. Distance between eyes 0.9-1.1 X width of one eye. Antennae yellow to brown-yellow (fig. 99), palpi yellow-red to brown. Pronotum: Yellow to yellow-red, large medial dark blotch that can be reduced to a round blotch on ante- VAN VONDEL: Oriental Haliplidae Figs. 107-112. Haliplus samosirensis, holotype. — 107, dorsal view; 108, antenna; 109, punctures near elytral base and suture; 110, dorsal side of hind tibia; 111, prosternal process; 112, lateral view of prosternal process. rior half with a partly reduced or vague blotch on ap- ical half (fig. 98). Lateral borders straight to slightly convex, anterior corners concave, clearly, vaguely or not visibly margined, margin never reaching anterior corner. Apical corners rounded, middle of base usual- ly finely impressed. Strongly, anteriorly weaker punc- tured, near basal corners with one or two stronger punctures in a slight depression. Basal punctures darkened. Elytra: Yellow to yellow-red, clear and extensive markings along suture and along base untill fifth pri- mary puncture-row, a central mark in third interval connected to suture and various small marks in later- al part (fig. 98). Primary punctures of average strength, stronger towards base, about 30 punctures in first row. Secondary punctures relatively strong ac- companied by some very small punctures. All punc- tures darkened except the central hole. Lateral bor- ders sometimes slightly serrate in basal part. Ventral side: Body yellow-brown to dark brown, elytral epipleura yellow to yellow-brown, legs yellow- brown with brown to dark-brown femora and coxae. Prosternal process with two strong grooves, anterior edge margined, punctured, sometimes strongly, espe- cially in the grooves, weakly narrowed near coxae, sligthly diverging towards apex (fig. 102, 103). Metasternal process flat with two pits, these pits sometimes very strong and than almost forming one confluent transverse pit, sparsely punctured (fig. 102). Coxal plates rather sparsely, near suture weakly a TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Figs. 113-121. Haliplus srilankanus, holotype. — 113, dorsal view; 114, antenna; 115, punctures near elytral base and suture; 116, dorsal side of hind tibia; 117, prosternal process; 118, lateral view of prosternal process; 119, left paramere; 120, penis; 121, right paramere. 52 punctured, sutural corners more or less rectangular. Sternites with complete puncture-rows. Last sternite almost completely covered with strong punctures, ap- ical point with a very short ridge. Setiferous striole on dorsal side of hind tibia very short, very close to the lateral row of setae and often hard to distinguish from it, longest of the tibial spurs about 4 X length of first tarsal segment (fig. 101). Male: First three tarsal segments of fore- and mid- legs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 104-106. Biology and distribution (fig. 126) Collected up to an altitude of 920 m. Attracted to light. China along the coast north to Xiamen, Thailand, Vietnam, Indonesia: Sumatra, Java, Sulawesi, India: West Bengal, Sri Lanka. Material examined. — China: 1 d, 12, Amoy [Xiamen], Galatea (ZMUC). — Thailand: 16, Lectotype; 19, Paralectotype, Browning, Siam; 1d, Siam; 2 ex., Bangkok, Castlenau (BMNH); 16, Cotype, Wallace, Mal.’ [Malacca], 67-56, Paralectotype (BMNH); 1 ex., Pattaya, 4.x11.1979 (MZLU); 16, Banna, Chawang nr. Nabon, 70 m., 5.ix.1958, J.L. Gressitt, light trap (BPBM); 6 ex., 25 km NW Lan Sak, 210 km NW Bangkok, viii.1991, W. Thielen (RMNH). — Vietnam: 1 2, Annam, Chuan-an [nr. Hue], Museum Paris, Coll. Maurice Régimbart, 1908, TYPE, oceanicus Rég. [not belonging to syntype-series]; 12, Annam, Chuan-an [nr. Hue]; 16, Cochin Chine, Harmand 1872; 1 m, Annam, Rég. de An-Ninh [?], R. P. M. Maunier 1903 (MNHN). — Sri Lanka: 16, Polonnaruwa, 28.xi.1980, M. Jäch (NHMV); 16, Maha-illerpulma [Madakalapuwa ?], 30.xi.1957, C. H. Fernando, light (BMNH). — India: 19, Calcutta, Atkinson coll. (BMNH). — Indonesia: Sumatra: 16, Lectotype of H. oceanicus (RMNH); 6 ex., Fort de Kock [Bukittinggi], 920 m, E. Jacobson, 1.1922, 1924, 1925, 1926 (BMNH, RMNH, ZMAN); 1 ex., Solok, exped. Kleiweg de Zwaan 1907; 1 ex., Deli, de Bussy (ZMAN); 8 ex., Medan, vii. 1904, 1906 (- RMNH); Java: 22 ex., Weltevreden [nr. Jakarta], 1915, Neytzell de Wilde (RMNH, ZMAN); 4 ex., G. Oengaran [G. Ungaran], v.1915, xii.1907, F. C. Drescher; 39 ex, Batavia [Jakarta], C. v. Nidek, 25.ix.1947, xii.1948, 11.1949, v.1949, vi.1949, xii.1949, 1.1950, v.1950; 2 ex., Tengger Mountains, Drescher; 1 ex., Tjilatjap, xii.1914, F. C. Drescher (ZMAN); 1 ex., Batavia [Jakarta], v.1913, J. B. Corporaal; 1 ex., Weltevreden [nr. Jakarta], 1.1919, P. B. Buytendijk (RMNH); Sulawesi: 1d, Makassar, I-74 [1.1874 ?] O. Beccari (Vondel); 3 ex., Makassar, L. de Vos (RMNH); 19, Sharp (MNHN). Haliplus samosirensis sp. n. (fig. 107-112) Type material. — Holotype 9, ‘Sumatra [Indonesia], Samosir Isl., 2°28’N 98°49’E, Tuk-Tuk, L. Toba, 2800 Ft. 4.11.1978, M. Holmen; 13.’ (zMuc). — Paratypes, 22: 19, same data as holotype (coll. Vondel); 12, ‘J. Corporaal, Sumatra’s O.K., Medan, 1.10.1921, 20 M, Haliplus pul- chellus v. oceanicus det. A. Zimmermann (ZMAN). VAN VONDEL: Oriental Haliplidae Diagnosis This species can be distinguished from related spe- cies by the elytra, which are provided with a basal dark band reaching margins and very irregular dark marks. Description Length 3.3-3.5 mm, width 1.9-2.0 mm. Body oval, widest in the middle (fig. 107). Head: Red-brown, frons vaguely marked, moder- ately strongly, on vertex stronger punctured. Distance between eyes about 1.1 X width of one eye. Antennae yellow-red (fig. 108), palpi yellow-red. Pronotum: Yellow-brown, gradually darkening to- wards the middle and base. Lateral borders convex, finely margined, anterior corner not margined and bending outwards. Strongly but not very densely punctured, all punctures darkened. Impressed oppo- site fourth to sixth elytral puncture-row (fig. 107). Elytra: Yellow to yellow-red, suture weakly dark- ened to first secondary punctures, base darkened to seventh row, darker towards shoulders, various dark marks (fig. 107). Completely margined, margin sometimes invisible from above. Primary puncture- rows strong and somewhat irregular, 22-26 punctures in first row. Secondary punctures strong, sometimes almost as strong as adjacent primary punctures. All punctures darkened with a yellow centre (fig. 109). Ventral side: Body yellow-red to brown, elytral epi- pleura yellow to yellow-brown, legs brown-red, dark- ened towards coxae. Prosternal process with two grooves, margined at anterior edge, narrowed near coxae, weakly punctured (fig. 111), lateral plicae complete (fig. 112). Metasternal process with trans- verse impression, in which two pits are recognisable, weakly punctured (fig. 111). Metacoxal plates dense- ly and rather strongly punctured, along suture with a weakly punctured area. Setiferous striole on dorsal side of hind tibia very short, longest of the two tibial spurs about % X length of first tarsal segment (fig. 110). Sternites strongly punctured. Male: Unknown. Biology and distribution (Be. 130) Up to an altitude of 850 m. Indonesia: Sumatra. Haliplus sharpi Wehncke Haliplus sharpi Wehncke, 1880 Haliplus tsukushiensis Yoshimura, 1932 This species is fully redescribed by Vondel (1991: 129). Additional notes: also recorded from Taiwan. Additional material examined: 16, Taiwan, Taihoku, 4.vi.1928, F. C. Hadden (BPBM). 515 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 Fig. 122. Oriental region as treated in this revision. Fig. 124. Distribution of Haliplus arrowi (dots: examined, Fig. 123. Distribution of Haliplus angustifrons (dots: exam circles: literature record). Fig. 125. Distribution of Haliplus ined, circles: literature record). perroti (dots) and Haliplus srilankanus (triangles). Fig. 126. Distribution of Haliplus pulchellus (dots: exam- ined, circles: literature records). Fig. 127. Distribution of Haliplus indicus (dot) and Haliplus nedungaduensis (triangle). Fig. 128. Distribution of Haliplus philippinus (dots) and Haliplus diopus (square). Fig. 129. Distribution of Haliplus pruthii (dots: examined, circle: literature record), Haliplus kapuri (triangle) and Haliplus manipurensis (square). Fig. 130. Distribution of Haliplus samosirensis (dot) and Haliplus javanicus (triangles). 314 Haliplus srilankanus sp. n. (figs. 113-121) Type material. — Holotype à, ‘Sri Lanka, Tissawewa, 10.xii.1979, V. Mahler Jensen leg. (ZMuc). Paratype 6, same data as holotype (Vondel); Paratype 2, ‘Sri Lanka, Tissawewa, 10.xii.1979, V. Mahler Jensen leg., Haliplus (Liaphlus) pulchellus Clark det. T. G. Vazirani 1981, SLO’ (ZMUC). Paratype, 2d, ‘India, Rajasthan, Bharatpur, 11. viii.1989, A. Riedel’ (Hendrich, Vondel). Diagnosis This species is related to A. angustifrons and H. ar- rowi, but can be distinguished from them by the char- acters given in the key. Description Length 3.2 mm, width 1.8 mm. Body oval, widest in the middle (fig. 113). Head: Red-brown, moderately strongly punctured. Distance between eyes about 1.1 X width of one eye. Antennae yellow-red (fig. 113), palpi yellow-red. Pronotum: Yellow-red, slightly and vaguely dark- ened in the middle near anterior side and before base. Lateral borders straight, finely margined, posterior corner protruding (fig. 113). In posterior corners strongly and densely punctured, impressed along base. Strongly and except on disc densely punctured, punctures not darkened except in the darkened cen- tral areas near base and anterior edge. Elytra: Yellow, suture completely darkened to first primary punctures, base darkened to fifth row, hooked marks on disc and in apical part connected to suture, marks in sixth interval (fig. 113). Completely margined. Primary puncture-rows moderately strong and not very dense, stronger in basal part, about 30 punctures in first row. Secondary punctures moder- ately strong, sparse or partly absent in even intervals. All punctures darkened with a yellow centre (fig. 11159). Ventral side: Body yellow-red to red-brown, elytral epipleura yellow, legs yellow-brown, darkened to- wards coxae. Prosternal process impressed in the mid- dle, margined at anterior edge, slightly narrowed near coxae, strongly and coarsely punctured (fig. 117), lat- eral plicae complete (fig. 118). Metasternal process widely and strongly impressed in the middle, weakly punctured (fig. 117). Metacoxal plates moderately strongly punctured, along suture more weakly punc- tured. Setiferous striole on dorsal side of hind tibia about 4 X length of tibia, consisting of about 12 small punctures, longest of two tibial spurs about 7 X length of first tarsal segment (fig. 116). Sternites weakly punctured. Male: First three tarsal segments of fore- and mid- legs widened, scaly hairs on ventral side. Penis and parameres as in figs. 119-122. VAN VONDEL: Oriental Haliplidae Biology and distribution (fig. 125) Biology unknown. Sri Lanka. India: Rajasthan. Unnamed material The specimens I failed to determine are listed be- low. Some of them might belong to still undescribed species. Species near H. diruptus. — 16, [India], Calcutta, Type, Atkinson Coll., BM 92-3, Haliplus falsificus n. sp. [MS-name] Type! J. Balfour-Browne det. (BMNH) Species near H. kotoshonis. — 19, Laos, Vientiane, 9.v.1965, P.D. Ashlock, at light (BPBM) Species close to H. indicus, but with unmaculated pronotum. — 1®, Sarda, Bengal, F. W. C. (BMNH) Specimens probably representing H. pruthii.—1°, [India], Uttar Pradesh, Haldwani Divn., Kumaun, vii.1923, H. G. C., at light, 3956 (BMNH); 19, (India, Bihar], Chota-Nagpore, Nowatoli, R. P. Cardon, viii-ix.1896 (MNHN); 19, N. India, U.P., Rishikesh, 450 m, viii.1988, Werner (NHMV). ACKNOWLEDGEMENTS I wish to express my sincere thanks to the following persons for placing material or information at my dis- posal: R. Angus (Egham), L. Baert (Brussels, ISNB), M. J. D. Brendell (London, BMNH), B. Brugge (Amsterdam, ZMAN), S. P. Cover (Cambridge, MCZC), R. Danielsson (Lund, MZLU), K. Desender (Brussels, ISNB), J. P. Duffels (Amsterdam, ZMAN), M. Hansen (Copenhagen, ZMUC), L. Hendrich (Berlin), S. J.Hine (London, BMNH), M. Holmen (Copenhagen, ZMUC), J. Huijbregts (Leiden, RMNH), M. Jäch (Vienna, NHMV), J. Krikken (Leiden, RMNH), P. Lindskog (Stockholm, NHRS), E. Matsui (Hondo City), P. Mazzoldi (Brescia), H. Perrin (Paris, MNHN), G. A. Samuelson (Honolulu, BPBM), G. Scherer (Munich, ZSMC), A. Smetana (Ottawa, CNCI), P. J. Spangler (Washington DC, USNM). The Uytten- boogaart-Eliasen Foundation is acknowledged for fi- nancial support. REFERENCES Balfour-Browne, J., 1946. The aquatic Coleoptera of Manchuria (Weymarn Collection). — Annals and Magazine of Natural History (11) 13: 433-460. Chapin, E. A., 1930. New species of Haliplus Latr. (Col.). — Proceedings of the Biological Society of Washington 43: 9-12. Clark, H., 1863. Descriptions of new East-Asiatic species of Haliplidae and Hydroporidae. — Transactions of the Entomological Society of London 3: 417-428. Falkenstrôm, G., 1932. Vorläufige Mitteilung über die neuen Halipliden und Dytisciden, von Dr. D. Hummel >15 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 in den Jahren 1927-30 während Dr. Sven Hedins China- Expedition eingesammelt. — Entomologisk Tidskrift 53: 191-192, 1 pl. Falkenstróm, G., 1936. Schwedisch-chinesische wissens- chaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof.-Sü Ping-chang. Insekten, 15, Coleoptera. 1. Haliplidae und Dytiscidae. — Arkiv för Zoologi 27A: 1-7. Gschwendtner, L. 1934. Aquatic insects of China. Article XVI. Eine neue Art von Haliplidae aus China. — Peking Natural History Bulletin 9: 107. Guignot, F., 1936. Description d’Haliplus nouveaux (Col. Haliplidae). — Bulletin de la Société entomologique de France 41: 115-118. Guignot, F., 1950. Trente-et-unieme note sur les Hydrocanthares. — Bulletin mensuel de la Societe Linneenne de Lyon 19: 25-28. Guignot, F., 1954. Entomological results from the Swedish expedition 1934 to Burma and British India. — Arkiv för Zoologi 6: 563-567. Guignot, F., 1955a. Quarante-deuxieme note sur les Hydrocanthares. — Revue Frangaise d’Entomologie 22: 270-276. ' Guignot, F., 1955b. Sur la systématique des Haliplus. — Mémoires de la Société Royal Entomologique de Belgique 27: 289-296. Guignot, F., 1956. Un nouvel Haliplus de Birmanie. — Arkiv for Zoologi 9: 477. Hope, F. W., 1845. On the entomology of China, with de- scriptions of the new species sent to England by Dr. Cantor from Chusan and Canton. — Transactions of the Entomological Society of London 4: 4-17. Kano, T. & K. Kamiya, 1931. Two new species of Haliplidae from Japan. — Transactions of the Kansai Entomological Society 2: 1-4, 1 pl. Peschet, R., 1923. Dytiscidae et Haliplidae nouveaux (Col.). — Bulletin de la Société entomologique de France1923: 175-181. Régimbart, M., 1885. Description d’une espèce nouvelle de Haliplides. — Notes from the Leyden Museum 7: 55-56. Régimbart, M., 1886. Dytiscides et Gyrinides nouveaux de la collection du Musée de Leyde. — Notes from the Leyden Museum 8: 139-144. Régimbart, M., 1889. Contributions à la faune Indo- Chinoise. — Annales de la Société Entomologique de 316 France (6) 9: 147-156. Régimbart, M., 1892. Insectes du Bengale Occidental. — Annales de la Société Entomologiques de Belgique 36: 112-121. Régimbart, M., 1899. Révision des Dytiscidae de la région Indo-Sino-Malaise. — Annales de la Société Entomolo- gique de France 68: 186-367. 4 Satô, M., 1984. Taxonomic notes on the aquatic Coleoptera of Japan I. — Coleopterists’ News 65: 1-4. Sharp, D., 1873. III. The water beetles of Japan. — Transactions of the Entomological Society of London 21: 45-67. Takizawa, M., 1931. The Haliplidae of Japan. — Insecta Matsumarana 5: 137-143. Vazirani, T. G., 1966. A review of the Indian Haliplidae (Insecta: Coleoptera), with descriptions of two new spe- cies. — Proceedings of the Zoological Society, Calcutta 19: 127-134. Vazirani, T. G., 1975. A new species of Haliplidae (Insecta: Coleoptera) from India. — Oriental Insects 9: 317-318. Vondel, B. J. van, 1991. Revision of the Palaearctic species of Haliplus subgenus Liaphlus Guignot (Coleoptera: Haliplidae). -Tijdschrift voor Entomologie 134: 75-144, figs 1-312. Vondel, B. J. van, 1992. Revision of the Palaearctic and Oriental species of Peltodytes Régimbart (Coleoptera: Haliplidae). — Tijdschrift voor Entomologie 135: 275- 267, figs 1-101. Wehncke, E., 1880. Neue Haliplus. — Stettiner Entomolo- gische Zeitung 41: 72-75. Yoshimura, S., 1932. On a new species of Haliplus from Japan. — Mushi 5: 102-103. Zimmermann, A., 1920. Dytiscidae, Haliplidae, Hygrobii- dae, Amphizoidae. — Coleopterorum Catalogus 71: 1- 325. Zimmermann, A., 1924. Die Halipliden der Welt. — Entomologische Blatter (fiir Biologie und Systematik der Kafer) 20 (1): 1-16; (2): 65-80; (3): 129-144; (4): 193- 213. Zimmermann, A., 1927. Fauna sumatrensis. Beitrag Nr. 45. Revision der Haliplidae et Dytiscidae von Sumatra. — Supplementa Entomologica 16: 1-44. Received: 3 March 1993 Accepted: 4 May 1993 Volume 136 1775 Tijdschrift voor | Entomologie A journal of systematic and evolutionary entomology since 1858 Published by the Nederlandse Entomologische Vereniging Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. 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 Published with index of volume 136 (1993). Graphic design Ontwerpers B.V., Aad Derwort, 's-Gravenhage Tijdschrift voor Entomologie Contents of volume 136 Articles 113 125 133 137 147 235 23 55 257 Andersen, T.: see Saether Beccaloni, G. W. A new species of Extatosoma Gray (Phasmatodea: Phasmatidae) from Papua New Guinea, with remarks on the species in the genus. Chen, P. P.: see Nieser Cuppen, J. G. M. Distribution and ecology of Hydraena in The Netherlands (Coleoptera: Hydraenidae). Donnelly, T. W. Two new genera of isostictid damselflies from New Britain, Bougainville, and the Solomon Islands (Odonata: Zygoptera). Haitlinger, R. A new species of the genus Diplothrombium Berlese (Acari, Prostigmata, Johnstonianidae) from Poland, based on the larva. Hämäläinen, M. Description of Neurobasis daviesi sp.n. from Palawan, with taxonomic notes on other species of the N. chinensis group (Odonata, Calopterygidae). Häuser, C. L. An annotated checklist of the species of the Parnassiinae (Lepidoptera: Papilionidae). Hurkmans, W. A monograph of Merodon (Diptera: Syrphidae). Part |. Jong, H. de The phylogeny of the Nephrotoma flavescens species group (Diptera: Tipulidae). Lansbury, I. Strongylovelia (Veliidae) and Metrobatopsis (Gerridae) and associated pleus- ton Hemiptera of West New Britain. Lansbury, I. Rhagovelia of Papua New Guinea, Solomon Islands and Australia (Hemiptera - Veliidae). Mathis, W. N. & T. Zatwarnicki Revision of the shore-fly genus Chlorichaeta Becker (Diptera: Ephydridae). Neboiss, A. New species of the genus Molanna Curtis from Sulawesi (Trichoptera: Molannidae). 259 Nieser, N. & P. P. Chen The Rhagovelia (Heteroptera: Veliidae) of Sulawesi (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), IV. 77 Pape, T. & H. Shima A new genus of Tachinidae from the Phillipines (Diptera). 83 Roth, L. M. Revision of the cockroach genus Ctenoneura Hanitsch (Blattaria, Polyphagidae). 283 Sæther, O. A. & T. Andersen Lobosmittia, a new genus of orthoclads from Tanzania and Turkey (Diptera: Chironomidae). - Shima, H.: see Pape 289 Vondel, B. J. van Revision of the Liaphlus species of the Oriental region excluding China (Coleoptera: Haliplidae). Zatwarnicki, T.: see Matthis Announcements and book reviews 14 Ossiannilson, F., 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. [J. van Tol]. 82 Scoble, M. J., 1992. The Lepidoptera: Form, function and diversity. [E. J. van Nieukerken]. | Fibiger, M., 1993. Noctuidae Europaeae. Volume 2, Noctuinae Il. [E. J. van Nieukerken]. Ill Publications Netherlands Entomological Society. 112 Announcement: 5th European Congress of Entomology, University of York, U. K. 124 Announcement: XX International Congress of Entomology. 282 Kuchlein, J. H. (with collaboration of J. H. Donner), 1993. De Kleine Vlinders. Handboek voor de Faunistiek van de Nederlandse Microlepidoptera. [E. J. van Nieukerken]. 288 Announcement: XX International Congress of Entomology, intent form. Referees for volume 136 R. A. Angus (London), P. Ashe (Dublin), D. A. L. Davies, P. L. T. Beuk (Leiden), P. E. Bragg (Nottingham), P. Goeldlin (Lausanne), R. W. Holzenthal (St Paul), D. K. McAlpine (Sydney), N. Méller Andersen (Copenhagen), R. V. Southcott (Adelaide), M. C. D. Speight (Dublin), I. Tangelder (Leiden), J. A. L. Watson (Canberra) Dates of Publication Volume 136 (1), pages 1-112, | July 1993 Volume 136 (2), pages 113-316, 10 December 1993 © Nederlandse Entomologische Vereniging, Amsterdam _IssN 0040-7496 NEW TAXA DESCRIBED IN TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136 ACARI Diplothrombium ludwinae Haitlinger .................... 12 BLATTARIA Ctenoneura crassistyla Roth ......... 107 Ctenoneura gigantea ROLE 0014 RE AU 87 Ctenoneura kinabaluana Roth … … nn. 104 Ctenoneura luma Roth... 101 Ctenoneura murudensis Roth................................ 106 Ctenoneura parascutica Roth … … … … 103 @ienoneurayparınga Roth. 2.7. 100 Ctenoneura propannulicornis Rother wert settee ae 88 Ctenoneura scutica Roth... 102 Cicnoncuratspinasiyla Roth: ne inn 98 Ctenoneura triprocessa Roth … … … … … n 106 Ctenoneura uncata Roth ........................................ 99 COLEOPTERA Haliplus javanicus van Vondel … … 301 Haliplus nedungaduensis van Vondel..................... 307 Haliplus samosirensis van Vondel … … 313 Haliplus srilankanus van Vondel … … 315 DIPTERA vaponcormyia Pape 82 Shninmas seated senaat: 77 Aporeomyia antennalis Pape & Shima .................... VAI Azpeytia shirakii Hurkmans............................... 208 nom. nov. for Merodon scutellaris Shiraki, 1968) Chlorichaeta africana Mathis & Zatwarnicki ......... 62 Chlorichaeta mais Mathis & Zatwarnicki ............... 66 Chlorichaeta orba Mathis & Zatwarnicki ............... 70 Lobosmittia Sæther & Andersen... 283 Lobosmittia basilobata Sæther & Andersen........... 284 Merodon aberrans isperensis Hurkmans ................ 177 Merodon altinosus Hurkmans............................... 164 Merodon ankylogaster Hurkmans ........................ 169 Merodon aureotibia Hurkmans … … 203 Merodon auronitens Hurkmans............................ 170 Merodon bequaerti Hurkmans … ne 194 Merodon cupreus Hurkmans …............................. 179 Merodon elegans Hurkmans ...................... 195 Merodon hypochrysos Hurkmans........................... 165 Merodon isperensis Hurkmans (ssp. of aberrans) … 177 Merodon kaloceros Hurkmans .......... 187 Wierodon lucas: AUTRMANS 2... 198 Merodon lusitanicus Hurkmans … … 181 Merodon marginicornis Hurkmans............. 166 Merodon mariae Hurkmans … … un 160 Merodon nitidifrons Hurkmans.........................- 199 Merodon oidipous Hurkmans … … … iii 171 Merodon ottomanus Hurkmans … … 161 Merodon persicus Hurkmans ............ n 171 Merodon satdagensis Hurkmans............................ 200 Merodon schachti Hurkmans … … … 201 Merodon sophron Hurkmans … annae 168 Merodon splendens Hurkmans....................... 182 Merodon tangerensis Hurkmans............................ 172 Merodon taniniensis Hurkmans ........................... 201 Merodon testaceoides Hurkmans … … … … … …… … 162 Merodon toscanus Hurkmans … … … 202 Merodon vandergooti Hurkmans … … 188 Merodon warnckei Hurkmans … … i 184 Merodon xanthipous Hurkmans … nennen 175 HEMIPTERA Rhagovelia amnicus Lansbury … … nennen eneen 31 Rhagovelia aureospicata Lansbury........................... 41 Rhagovelia blagiokommena Nieser & Chen … … 268 Rhagovelia browni Lansbuty … nennen 33 Rhagovelia caesius Lansbury ........................2..........I0 Rhagovelia chrysomalla Nieser & Chen … 268 Rhagoveliatenmtallansbury an. 46 Rhagovelia daktylophora Nieser & Chen .............. 270 Rhagovelia fulvus Lansbury............22e22220222.20220202200e 36 Rhagovelia herzogensis Lansbury........................ 43 Rhagovelia hirsuta Lansbury......... i 39 Rhagovelia horaia Nieser & Chen ....................... 271 Rhagovelia kalami Nieser & Chen ....................... 273 Rhagovelia kastanoparuphe Nieser & Chen ......... 274 Rhasovelapriordbansbun sees eee eee 29) Rhagovelia pseudocelebensis Nieser & Chen … … 275 Rhagovelia robina Nieser & Chen........................ 275 Rhagovelia thysanotos Lansbuty … nennen 48 Rhagovelia trichota Nieser & Chen....................... 277 Rhagovelia tropidata Nieser & Chen.................... 278 Strongylovelia priori Lansbuty ............. 15 ODONATA @OnenvsticaDonnell ge CREO 128 Cnemisticta angustilobata Donnelly...................... 128 Cnemisticta latilobata Donnelly … … 129 Neurobasis daviesi Hämäläinen … 135 Ian: Donne 126 Titanosticta macrogaster Donnelly … … 126 PHASMATODEA Extatosoma carlbergi Beccaloni … … … 115 TRICHOPTERA Molanna jolandae Neboiss.................. 257 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 136, 1993 INSTRUCTIONS TO AUTHORS The Tijdschrift voor Entomologie publishes original papers dealing with systematic and evolutionary ento- mology. 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Vii | | i Lan un SR Er TI el Pr ve Bani (up He VON AN Cap itis ab var "e | | | Ù J | i | My ia mf | I nr VISI Laie) CO - SA RR Mikael tat eat L oo Dette STU Se hi dar ob Mat US h / I ’ i raven ball Ù Ni | FR i eee fade vaer 4 I y k Ath Doh a sti | alu \ ATOM / {| al a \ VE Cat the “OPEN Ten STONE Le HAUTS pi We a si ed 3 Anke | 1 HR Pag ; i r CAMERA seh VT ARM pd bled ‘ \ vasi m i ae hy nied di pi Hd En wre: ai sr Fl 2 - PARAT AIAR DT - Soldi A tra ro “Me é i ss à pa crap ein’ MA du ni “din ¢ a } Nee \ il fe ua verf Paes È 4 L «ii Ye i whe 7 ta ki if DITO, rl ara ‘hé i oll: wis nd nr athe NT ae té CN i AN: ARE Ni hand Me | GAVIN Wik ung Wa Ù Pl Thy bhi) a! wi iV ut Le ve ver KL ij mn! u perni, i" | ue eher us n x uranio ® i Tijdschrift voor Entomologie Volume 136, no. 2 Articles 113 G. W. Beccaloni A new species of Extatosoma Gray (Phasmatodea: Phasmatidae) from Papua New Guinea, with remarks on the species in the genus. 125 T. W. Donnelly Two new genera of isostictid damselflies from New Britain, Bougainville, and the Solomon Islands (Odonata: Zygoptera). 133 M. Hämäläinen Description of Neurobasis daviesi sp.n. from Palawan, with taxonomic notes on other species of the N. chinensis group (Odonata, Calopterygidae). 137 C. L. Häuser An annotated checklist of the species of the Parnassiinae (Lepidoptera: Papilionidae). 147 W. Hurkmans A monograph of Merodon (Diptera: Syrphidae). Part |. 235 H. de Jong The phylogeny of the Nephrotoma flavescens species group (Diptera: Tipulidae). 257 A. Neboiss New species of the genus Molanna Curtis from Sulawesi (Trichoptera: Molannidae). 259 N. Nieser & P. P. Chen The Rhagovelia (Heteroptera: Veliidae) of Sulawesi (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), IV. 283 O. A. Sæther & T. Andersen Lobosmittia, a new genus of orthoclads from Tanzania and Turkey (Diptera: Chironomidae). 289 B. J. van Vondel Revision of the Liaphlus species of the Oriental region excluding China (Coleoptera: Haliplidae). Announcements and book reviews 124 Announcement: XX International Congress of Entomology. 282 Kuchlein, J. H. (with collaboration of J. H. Donner), 1993. De Kleine Vlinders. Handboek voor de Faunistiek van de Nederlandse Microlepidoptera. — PUDOC, Wageningen. [E. J. van Nieukerken]. 288 Announcement: XX International Congress of Entomology, intent form. © Nederlandse Entomologische Vereniging, Amsterdam Published 10 December 1993 ISSN 0040-7496 x fi 5 5 ay A 4 / 4 © FN A | \ | 1 x 19 \ es \ \ ; | | | / ack cover um su | ‘i Lion 4 dla (67770 ORN Nur D À 7 ” 4 ERNST MAYR LIBRARY DEMCO INC 38-2931 pr ap MO hy PANNE TROIA COMICA 8 ben abri OD Ba SI Ventura es pre USER), Gast yh tes su BPA vga ator Yasue oh WELT ety) Cut orb rund, REA MEN AKA Rn vrt Ward ydeurirery AEG Wht Wen 0015 ete it N vlek Dif ; Ù quel uf AMARA) HOME \ t beurt Hoban PIANTE SERIA TI MILA i H PW eho ay AT DENT % x \ Hi TELE Ot one) i; ne Hi D A TE | ACC wein die vk do Wie tH