M te 4 M ft ANNALES de la SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSÉUM D'HISTOIRE NATURELLE de la Ville de Genève tome 1 1 6 fascicule 3-4 2009 o o N LU o < Z O C/Ì M i ël GENEVE DECEMBRE 2009 ISSN 0035 - 41 8 X Q/i LU u o o o N LU û LU oc/5 LU > LU REVUE SUISSE DE ZOOLOGIE TOME 116— FASCICULE 3-4 Publication subventionnée par: Académie suisse des Sciences naturelles (SCNAT) Ville de Genève Société suisse de Zoologie Comité de rédaction DANIELLE DECROUEZ Directrice du Muséum d'histoire naturelle de Genève ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d'histoire naturelle de Genève Comité de lecture A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons), B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes), M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger (arthropodes excl. insectes). Le comité soumet chaque manuscrit pour évaluation à des experts d'institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. Administration MUSÉUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm Prix de l'abonnement: SUISSE Fr. 225.— UNION POSTALE Fr. 250.- (en francs suisses) Les demandes d'abonnement doivent être adressées à la rédaction de la Revue suisse de Zoologie, Muséum d'histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse ANNALES de la SOCIÉTÉ SUISSE DE ZOOLOGIE et du MUSÉUM D'HISTOIRE NATURELLE de la Ville de Genève tome 1 1 6 fascicule 3-4 2009 LU O o o o N û o O N CE O en > JeL GENEVE DECEMBRE 2009 ISSN 0035 -418 X ro > ex. Peduncle of sensillus short, curved, its head small, asymmetric, with some very small spicules on its distal margin. MOSS MITES FROM SWITZERLAND 327 Figs 1-3 Topobates helveticus sp. n. (1) Body in dorsal view. (2) Body in ventral view. (3) Podosoma in lateral view. 328 S. MAHUNKA & L. MAHUNKA-PAPP Notogaster. Dorsosejugal suture distinct, convex, pteromorphae small, blade- like. Whole surface finely punctate. Thirteen pairs of simple, distinctly and finely barbed, long notogastral setae and four pairs of small sacculi present, Sa slightly larger than the others. Lateral part of podosoma: Sublamellar area porosa small. Lateral region under prelamella ornamented with weak polygonal pattern (Fig. 3). Pedotecta 1 narrow. Ventral parts: Surface of infracapitulum, epimeral and ventral plates nearly smooth. Epimeral fields well framed laterally and medially (Fig. 2). Discidium weakly developed, not protruding. All ventral parts with the usual numbers of setae, i.e. epimeral setal formula: 3-1-3-3, anogenital setal formula 4-1-2-3. Except for setae ad x and ad 2 , all others short and simple. Legs: All legs tridactylous and heterodactylous. Remarks: The new species is well characterised by the thirteen pairs of very long notogastral setae and by the finely punctate notogastral sculpture. On this ground it stands nearest to T. comatus (Pérez-Inigo, Herrero & Pérez-Inigo, 1987). Thus, the currently known European Topobates-fauna. comprises eight species. On the basis of their main characters (length of notogastral setae, number of noto- gastral setae and sculpture of notogaster), they can be arranged into three groups. Weigmann & Miko (1998) compiled an identification key for the previously known species, we here add the two Swiss species T. umbraili and T. helveticus sp. n. Key to European Topobates species la Surface of notogaster granular granifer Grandjean, 1958 lb Surface of notogaster smooth, or finely punctate 2 2a Twelve pairs of notogastral setae present carpathicus Weigmann & Miko, 1998 2b Thirteen or fourteen pairs of notogastral setae present 3 3a Fourteen pairs of notogastral setae and five pairs of sacculi present alvaradoi Pérez-Inigo, 1969 3b Thirteen pairs of notogastral setae and four pairs of sacculi present 4 4a Notogastral setae long, distance between the insertion of dm and dp about as long as the length of setae dm 5 4b Notogastral setae short, distance between the insertion of dm and dp much longer than the length of setae dm 6 5a Interlamellar setae very long, reaching far over the rostral cusp. Setae dp much longer than the distance between setae dm and dp comatus (Pérez-Inigo, Herrero & Pérez-Inigo, 1987) 5b Interlamellar setae ending far from the rostral cusp. Setae dp shorter than the distance between setae dm and dp helveticus sp. n. 6a Sensillus long, reaching lateral margin of pteromorphae umbraili (Schweizer, 1956) 6b Sensillus short, not reaching lateral margin of pteromorphae 7 7a Notogaster with a semicircular band running parallel to margin of body. Setae c 2 shorter than posterior notogastral setae circumcarinatus Weigmann & Miko, 1998 MOSS MITES FROM SWITZERLAND 329 7b No semicircular band on notogaster. Setae c 2 not shorter than posterior notogastral setae holsaticus Weigmann, 1969 Family Microzetidae Grandjean, 1936 Miracarus similis Subias & Iturrondobeitia, 1978 Figs 4-8 Locality: VS-48: Switzerland: Valais: Vouvry, layer of mosses from a rocky slope on the trail to the cave „Grotte de la Pierre Perret" (B), 460 m; 10. Vili. 1989; leg. B. Hauser. Remarks: The genus Miracarus Kunst, 1959 so far comprises five species, all from the Mediterranean region of Europe. However, according to Subias (2004, 2008), M. abeloosi Lions, 1979 is conspecific with the above species, therefore the genus would have only four species. As far as we are concerned, the synonymy of similis and abeloosi needs further proof, also because Pérez-Inigo (1997) made no mention of this opinion. By comparing the published figures of the two species, significant differences can be established, although the figure of similis is highly simplified, and we do not know whether the authors have studied the types of both species. The lamellar apices of M. similis described from Arratia (Viczaya) are very wide and touching each other medially. A well-developed sejugal line is present, which is not interrupted medially. M. abeloosi was described by Lions (1979) from specimens collected in France, in the region of 'Alpes Maritimes", in the environs of "Le Chens'" and "Sainte Baume". His description is perfect, covering all the details, but differs from the original description of M. similis. The Swiss specimens, the figures of which are given hereunder (Figs 4-8), may be conspecific with the ones from France. The shape of the lamellar apices in the Swiss specimens is different, and the dorsosejugal line is clearly interrupted. The known localities in France are quite close to those of the Swiss specimens, therefore it is not surprising that they belong to the same species. We accept the opinion of Subias, with the reservation that a further study of the types is necessary. Key to the species of Miracarus la Outer cusps of lamellae four to five times longer than the inner cusps 2 lb Both lamellar cusp nearly equal in length 3 2a Head of sensillus long, with spinifom distal end liurkai Kunst, 1958 2b Head of sensillus rounded distally discrepans Mahunka, 1966 3a Distal end of lamellae wide, lamellar setae located in the middle similis Subias & Iturrondobeitia, 1978 3b Distal end of lamellae narrowed, lamellar setae located laterally senensis Bernini, 1975 Family Nosybeidae Mahunka, 1993 Lamellocepheus personatus (Berlese, 1910) Figs 9-14, 19 Localities: VS-40: Switzerland: Valais: above Vernayaz, 700 m; 7.X.1980; leg. C. Besuchet (221). - TI-42: Switzerland: Ticino: Cortascio above Brissago, sifting of mosses, 900 m; 20.V.1998; leg. C. Besuchet (229). Remarks: This species was studied and redescribed by Weigmann & Murvanidze (2003) on the basis of Georgian specimens. They gave a thorough histori- 330 S. MAHUNKA & L. MAHUNKA-PAPP FlGS 4-8 Miracarus similis Subias & Iturrondobeitia, 1978. (4) Body in dorsal view. (5) Body in ventral view. (6) Sensillus and pteromorpha. (7) Body in lateral view. (8) Lamellar cusps. MOSS MITES FROM SWITZERLAND 331 Figs 9-14 Lamellocepheus personatus (Berlese, 1910). (9) Body in dorsal view. (10) Rostral apex. (11) Body in ventral view. (12) Podosoma in lateral view. (13) Anterior part of notogaster. (14) Trichobothrium. 332 S. MAHUNKA & L. MAHUNKA-PAPP cal survey of the morphological and nomenclatural problems of the genus and species. The species, now also known from Switzerland, has been found in the Mediterranean region, particularly at alpine localities. These specimens are easily identifiable with the description and figures published by Weigmann and Murvanidze (cf. our Figs 9-14 made from the Swiss specimens,). The present study gave us the opportunity to review the taxa belonging to the genera Lamellocepheus Balogh, 1961 and Nosybea Mahunka, 1993. Soon after the publication of the paper of Weigmann and Murvanidze (2003), Subfas (2004) accepted the validity of the family Nosybeidae, but synonymised the two genera, without having studied their types. A fairly recent study and the re-examination of the type specimens of Nosybea genavensis Mahunka, 1993 and Lamelloceppheus personatus by Mahunka (1993) clearly show that Subfas made a mistake, so the validity of the genus name Nosybea is herewith re-established. Distinction between Lamellocepheus and Nosybea Lamellocepheus 1. Middle of anterior half of notogaster with a U-shaped, undivided suture. 2. Epimeral region without annulate structure. 3. A pair of ridges along the genital aperture. 4. Five pairs of genital setae present. Nosybea 1 . Anterior part of notogaster in the middle with a longitudinal crest. 2. Epimeral region with 12 robust structures. 3. No pairs of ridges along the genital aperture. 4. Four pairs of genital setae present. Family Ceratozetidae Jacot, 1925 Jugatala ungulata (C. L. Koch, 1839) Figs 15-18 Material examined: VS-44: Switzerland: Valais: Saas-Almagell, waterlogged mosses, 1650 m; 5.VII.1997; leg. C. Besuchet (225). Measurements: Length of body: 505-520 pm, width of body: 351-363 /mi. Description: Prodorsum: Rostrum rounded, without rostral apex. Surface of rostral part striated longitudinally. Other parts of surface polygonate. A distinct trans- versal, arched lath present in front of the lamellae. Lamellar complex well developed, wide, typical for this genus. Lamellae and translamella nearly equal in width, lamellar setae arising on distinct lamellar cusps. Interlamellar setae strong, straight, distinctly pilose. Bothridium cup-shaped, covered by dorsosejugal tectum. Sensillus short, rounded in dorsal view, hemispherical in lateral view, its surface finely aciculate. Notogaster. Dorsosejugal suture distinct, strongly convex, whole surface with distinct polygonate ornamentation. Pteromorphae small, triangular, a hinge line partly observable. Eleven pairs of short, mostly bacilliform, dark notogastral setae present, all finely barbed. Four pairs of small and round porose areas and 5 pairs of lyrifissures present. All porose areas nearly equal in length. MOSS MITES FROM SWITZERLAND 333 Figs 15-17 Jugatala ungulata (C. L. Koch, 1839). (15) Body in dorsal view. (16) Body in ventral view. (17) Posteromarginal part of notogaster. Lateral part of podosoma: Genal tooth wide, elongate. Tutorium narrow, striate, conspicuously arched ventrally, with small apex. Rostral setae arising on rostral surface. Pedotectum I large, wide, distinctly striate. Pedotecta II-III covering posterior 334 S. MAHUNKA & L. MAHUNKA-PAPP FlGS 18-19 (18) Jugatala ungulata (C. L. Koch, 1839). lateral part of podosoma. (19) Lamellocepheus personatus (Berlese, 1910), ventral plate and posterior part of epimeral region. MOSS MITES FROM SWITZERLAND 335 half of acetabulum. Three humerosejugal porose areas present. Sublamellar porose area distinctly visible. Circumpedal carina present. Ventral regions: Apodemes 2 and ap. sej. connected medially. Surface of infra- capitulum smooth, epimeres and genital plate with very fine striation. Ventral and anal plate with polygonate sculpture. All epimeral setae thin, setiform, finely barbed. Custodian short, discidium large. Genito-anal setal formula: 6-1-2-3, anterior 3 pairs arising close to each other, along the anterior margin. Genital, aggenital and anal setae thin, simple, setiform, except for the anterior adanal setae ad^; all other setae bacilli- form, barbed, like the notogastral setae. Lyrifissures iad located at anterior corner of anal plates. Postanal porose area narrow, divided into 2-4 small parts. Legs: All legs tridactylous, with equally large claws. Remarks: The type species of the genus Jugatala Ewing, 1913 is J. tuberosa Ewing, 1913. Sellnick (1928, 1960) transferred the species Oribata ungulata CL. Koch, 1939 to this genus. The position of the genus Jugatala was several times mis- interpreted by different authors, so at times it was placed in the families Ceratozetidae and Mycobatidae. Having studied immature specimens, Behan-Pelletier (2000) placed Jugatala in the family Ceratozetidae. On the other hand, Subias (2004, 2008) ranged ungulata again in the subgenus Calyprozetes Thor, 1930 of the genus Mycobates in the family of Mycobatidae, although Jugatala tuberosa was placed by him in the family Ceratozetidae. This latter point of view obviously needs to be revised. Although Behan- Pelletier (2000) redescribed the type species, he did not deal with J. angulata, neither did Subias carry out a new study. On the basis of our recent study of the Swiss specimens, we state that angulata belongs in the genus Jugatala, and we accept the conclusions of Behan-Pelletier that this genus belongs in the family Ceratozetidae. We cannot deal with the other described species. J. (?) rotunda Willmann, 1953, described from Europe, is possibly a member of the genus Jugatala, but this supposition needs confirmation as noted by Willmann himself in 1953. A part of the species listed by Subias may not be members of the genus Jugatala, but rather of Calyptozetes. Of course, the taxonomic position of that genus is dubious. On the basis of the Swiss specimens examined we can provide the complemen- tary description given above. Jugatala angulata is well distinguished from the type species of the genus by the form of the body, by the form of the notogastral setae and, most importantly, by the number and form of the porose areas of the notogaster. The only specimen of this species previously known from Switzerland was collected by Schweizer (1948) in the canton of Graubünden. A recent redescription of this species was given by Bayartogtokh & Schatz (2008). ACKNOWLEDGEMENTS We like to thank the collectors of the material examined, mainly Dr C. Besuchet (Geneva), and Dr P. Schwendinger and Dr B. Hauser for the opportunity to study these specimens in the Geneva Museum. For linguistic revision and editorial help with the manuscript we extend our sincere thanks to Dr P. Schwendinger and Dr L. Zombori. This project was partly sponsored by the Hungarian Scientific Research Fund (OTKA, number T45889). 336 S. MAHUNKA & L. MAHUNKA-PAPP REFERENCES Bayartogtokh, B. & Schatz, H. 2008. Trichoribates and Jugatala (Acari: Oribatida: Cerato- zetidae) from the Central and Southern Alps, with notes on their distribution. Zootaxa 1948: 1-35. Behan- Pelletier, V. M. 2000. Ceratozetidae (Acari: Oribatida) of arboreal habitats. The Canadian Entomologist 132: 153-182. Grandjean, F. 1936. Les Microzetidae nov. fam. (Oribates). Bulletin de la Société Zoologique de France dì: 60-93. Grandjean, F. 1962. Le genre Tegeocranellus Beri. 1913 (Oribates). Acarologia 4: 78-100. Lions, J. C. 1979. Sur le genre Miracarus Kunst 1959, Oribate: Miracarus abeloosi sp. n. l re partie: la stase adulte. Acarologia 20(3): 404-418. Mahunka, S. 1993. Oribatids from Madagascar I. (Acari: Oribatida). New and interesting mites from the Geneva Museum LXXVI. Revue suisse de Zoologie 100(2): 289-315. Mahunka, S. & Mahunka-Papp, L. 2003. Oribatids from Switzerland Vili (Acari: Oribatida: Ptyctima). (Acarologica Genavensia CII). Revue suisse de Zoologie 110(3): 453-481. Pérez-Inigo, C. 1997. Acari, Oribatei, Gymnonota 1. In: Ramos, M. A., Tercedor, J. A., Ros, X. B., Noguera, J. G., Sierra, Ä. G., Mayol, E. M., Piera. F. M., Marino, J. S. & Gonzalez, J. T. (eds). Fauna Ibèrica. Museo National de Ciencias Naturales CSIC, Madrid, 374 pp. Schweizer, J. 1948. Die Landmilben aus der Umgebung des schweizerischen Nationalparks. Ergebnisse der wissenschaftlichen Untersuchungen des schweizerischen Nationalparks II (N. F.) 20: 3-28. Sellnick, M. 1928. Formenkreis: Hornmilben. Oribatei. In: Brohmer, P, Ehemann, P. & Ulmer, G. (eds). Die Tierwelt Mitteleuropas 9: 1-42. Sellnick, M. 1960. Formenkreis: Hornmilben, Oribatei. Nachtrag. In: Brohmer, P, Ehemann, P & Ulmer, G. (eds). Die Tierwelt Mitteleuropas Ergänzung: 45-134. Subîas, L. S. 2004. Listado sistematico, sinonìmico y biogeogrâfico de los âcaros oribâtidos (Acariformes, Oribatida) del Mundo (1758-2002). Graellsia 60: 3-305. Subi'as, L. S. 2008. Listado sistematico, sinonimico y biogeogrâfico de los âcaros oribâtidos (Acariformes, Oribatida) del Mundo (exepto fósiles) (Originally published in Graellsia 60: 3-305, 2004, actualized in May 2008, in pdf), 540 pp.^" Weigmann, G. 2006. Hornmilben (Oribatida). Die Tierwelt Deutschland 76: 1-520. Weigmann, G. & MiKO, L. 1998. Taxonomy of European Scheloribatidae, 3. Remarks on Scheloribates Berlese 1908 with description of two new species of the subgenus Topobates Grandjean 1958 (n. stat.) (Arachnida: Acari: Oribatida). Senckenbergiana bio- logica 77(2): 247-255. Weigmann, G. & Murvanidze, M. 2003. Contribution to the oribatid mite fauna of Georgia. 2. Carabodes and Lamellocepheus (Acari, Oribatida). Spixiana 26(3): 221-226. Willmann, C. 1953. Neue Milben aus den östlichen Alpen. Sitzungsberichte der Öster- reichischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abt. I. 162(6): 449-519. Revue suisse de Zoologie 116 (3-4): 337-352; décembre 2009 Oribatids from Madagascar IV (Acari: Oribatida) Sândor MAHUNKA Department of Zoology, Hungarian Natural History Museum and Systematic Zoology Research Group of the Hungarian Academy of Sciences, H-1088 Budapest, Baross utca 13, Hungary. E-mail: mahunka@gmail.com Oribatids from Madagascar IV (Acari: Oribatida). - Oribatida material collected in Madagascar by a scientist of the Muséum d'histoire naturelle, Genève was studied. Twenty species are listed, six of them are new to science: Mesoplophora (P.) madegassica, Masthermannia hauseri, Caveremulus foliaceus, C. salicinus, Ambrobates translamellatus and Vilhenabates ambohitra. The new genus Ambrobates gen. n. is established in the family Scheloribatidae. Taxonomical notes on rare or little known species and a key for identification of Caveremulus species are given. Keywords: New taxa - list of identified species - taxonomical and zoogeo- graphical notes - key - Malagasy Republic. INTRODUCTION The special and peculiar fauna of Madagascar is nowadays quite well known, however, the soil mesofauna, characterised by species with minute size and unspecta- cular life style, and especially the mites are an exception. Research on them has been carried out with high intensity recently (Mahunka 2002), and lately Dr Csaba Csuzdi, a well-known expert of earthworms, collected soil samples there in 2008 (Mahunka 2009). Earlier, at the end of the 20 th century, the soil fauna of Madagascar was systematically studied by Swiss taxonomists (Dr Bernd Hauser and Dr Charles Lienhardt of the Muséum d'histoire naturelle, Genève) and Hungarian (the associate co-workers of the Hungarian Natural History Museum, Dr Dénes Balâzs and Dr Tamâs Pócs) scientists. Due to these exploratory activities, the oribatid fauna of Madagascar, the former Malagasy Republic, is now quite well known. However, some regions of the island and the taxa expected to be found there - which constitute the majority of oribatids - are still considerably less well known and hence severe gaps in our knowledge exist. In this paper I continue my study of the material collected by researchers of the Muséum d'histoire naturelle, Genève (Mahunka 1993, 1994, 1997), publish the results of my taxonomic investigation and give further notes on the geographic distribution of oribatid species. Hereunder I list twenty species, six of which are new to science. Six Manuscript accepted 21.07.2009 338 S. MAHUNKA species are for the first time indicated in the Madagascan fauna: Epilohmannia pallida Wallwork, 1962, Hoplophorella vitrina (Berlese, 1913), Hoplophora hamata (Ewing, 1909), Tectocepheus velatus velatus (Michael, 1880), Ramusella clavipectinata (Michael, 1885) and Berlesezetes africanus (Balogh,1958). I give notes on the Madagascan specimens of two species. MATERIAL AND METHODS As in my earlier papers, I follow the system of Marshall et al. (1987), based on that of Grandjean (1954, 1965), with some modifications introduced by Woas (2002), Weigmann (2006) and Subias (2004, 2008). In the descriptions the morphological terminology of Grandjean (in several publications) was used with some modifications concerning the studied groups or organs (e.g. Mahunka & Zombori, 1985; Norton et al, 1997; Mahunka & Mahunka-Papp, 2001; Niedbala, 2001, 2004) and the above mentioned publications). All material examined is deposited in the Muséum d'histoire naturelle de Genève (MHNG) and in the Hungarian Natural History Museum, Budapest (HNHM). LIST OF COLLECTING SITES Mad-89/8: Madagascar (Prov. Antsiranana [anciennement Diego-Suarez], Sous-préf. Antsiranana): Pare National „Montagne d'Ambre" (=Ambohitra), au début du chemin vers la "Petit Cascade", après la pénière, forêt primaire, prélèvement de sol dans les angles formés par les contreforts d'un grand arbre vivant et d'un grand arbre mort, 980 m; 23.XI.1989, leg. B. Hauser. Mad-89/29: Madagascar (Prov. Antsiranana [anciennement Diego-Suarez], Sous-préf. Andonay [anciennement Hell-Ville]): île Nosy Be, Réserve naturelle intégrale „Lokobe", forêt primaire près d'Ampasindava, prélèvement de sol dans les angles formés par les contreforts d'un grand arbre, 14 m; 28.XI.1989, leg. B. Hauser. Mad-89/45: Madagascar (Prov. Toliara [Tulear], Sous-préf. Tôlenaro [Fort-Dauphin]): 73 km de Tôlenaro route vers Amboasary, plantation de sisal, prélèvement de sol au pied d'un baobab mort, env. 100 m; 4.XII.1989, leg. B. Hauser (extraction Berlese à Genève). LIST OF STUDIED SPECIES Epilohmanniidae Oudemans, 1923 Epilohmannia pallida Wallwork, 1962 Locality: Mad-89/8. First record for Madagascar. Mesoplophoridae Ewing, 1917 Mesoplophora (Parplophora) madegassica sp. n. EUPHTHIRACARIDAE JaCOt, 1930 Microtritia tropica Märkel, 1964 Locality: Mad-89/8. ORIBATIDS FROM MADAGASCAR 339 Steganacaridae Niedbala, 1 986 Hoplophorella viirina (Berlese, 1913) Locality: Mad-89/29. First record for Madagascar. Hoplophora hamata (Ewing, 1909) Locality: Mad-89/45. First record for Madagascar. Nanhermanniidae Sellnick, 1928 Masthermannia hauseri sp. n. Nanhermannia milloti Balogh, 1960 Locality: Mad-89/8. Zetorchestidae Michael, 1898 Zetorchestes {Phyllorchestes) phylligerus Mahunka, 1983 Locality: Mad-89/8. Eremulidae Grandjean, 1965 Caveremulus foliaceus sp. n. Caveremulus cordisetus Mahunka, 1983 Locality: Mad-89/29. Caveremulus salicinus sp. n. Tectocepheidae Grandjean, 1954 Tectocepheus velatus velatus (Michael, 1880) Locality: Mad-89/8. First record for Madagascar. Oppiidae Sellnick, 1937 Ramusella (Ramusella) clavipectinata (Michael, 1885) Locality: Mad-89/8. First record for Madagascar. Ramusella (Insculptoppia) aepyornis Mahunka, 1994 Locality: Mad-89/45. Oppiella nova (Oudemans, 1902) Locality: Mad-89/8. Fossoppia calcarata Mahunka, 1994 Locality: Mad-89/8. Microzetidae Grandjean, 1 936 Acaroceras (Malgoceras) hellen Mahunka, 1993 Locality: Mad-89/29. Berlesezetes africanus (Balogh, 1958) Locality: Mad-89/8. First record for Madagascar. Scheloribatidae Grandjean, 1933 Ambrobates translamellatus gen. n., sp. n. Haplozetidae Grandjean, 1936 Vilhenabates ambohitra sp. n. 340 S. MAHUNKA DESCRIPTIONS AND REMARKS Hoplophorella hamata (Ewing, 1 909) Remarks: Judging from my study of types, the new specimens collected appear clearly conspecific with H. schauenbergi Mahunka, 1978 and H. ligulifera Mahunka, 1987, which were treated by Niedbala (2001) as synonyms of H. hamata. H. ligulifera and H. schauenbergi were described from the Ethiopian region, H. hamata from the USA. The main characteristics of this species are: a slightly dilated, lanceolate lamellar seta and spiniform interlamellar setae, notogastral ornamentation consisting of finely punctulate small foveolae, and the long peduncle of the median adanal setae (ad 2 ). Mesoplophora (Parplophora) madegassica sp. n. Figs 1-4 Material examined: Holotype: Madagascar, Prov. Antsiranana; 23. XI. 1989; leg. B. Hauser (Mad-89/8); 3 paratypes from the same sample. Holotype and 2 paratypes deposited in MHNG, 1 paratype (1785-PO-2009) in HNHM. Diagnosis: Rostrum triangular, apex pointed. Rostral and lamellar setae seti- form, cibate, interlamellar setae filiform, smooth. Sensillus long, head small, dilate. Among the notogastral setae c 3 shorter and thinner than others. Latter setae finely cibate. Eight pairs of ventral setae, one pair of them very long. Seven (5+2) pairs of genital, three pairs of anal setae present, all finely cibate. Measurements: Length of aspis 196-216 ^m, height of aspis 148-152 /mi, length of notogaster 270-286 /urn, height of notogaster 182-197 /urn. Description: Prodorsum: Rostral apex sharply pointed. Prodorsal surface dis- tinctly punctulate. Prodorsal setae of different lengths and sizes, rostral and lamellar setae much thicker than interlamellar and exobothridial setae, rostral and lamellar setae well cibate, setiform, interlamellar setae very thin, filiform, smooth, exobothridial setae also smooth and filiform, the latter shortest of all, but longer than the diameter of the cup-shaped bothridium. Peduncle of sensillus conspicuously long, head small, lanceolate, with irregular velum and some small cilia. Notogaster. With eight pairs of long setae, one (setae c 3 ) much shorter and slightly thinner than others, smooth. Remaining setae with small cilia or spines, nearly equal in length. Setae c^ and c 2 arising far from notogastral border, setae c 3 located very close to this border. Ventral parts: Genital plates with seven (5+2) pairs of simple setae, all setiform and smooth. Anal plates with three pairs of thinner but longer setae, finely roughened. Nine pairs of thin ventral setae of different lengths, all slightly cibate. Lateral setae lon- gest of all. Legs: Typical for the genus. Remarks: On the basis of the number of anal setae, the new species belongs to the subgenus Parplophora Niedbala, 1985. The new species is well characterised by the striking shape of the sensillus, which was previously unknown for Mesoplophora taxa from the Ethiopian Region (Niedbala 1985). Etymology: Named after its country of origin. ORIBATIDS FROM MADAGASCAR 341 Figs 1-4 Mesoplophora (Parplophora) madegassica sp. n. ( 1 ) Body in lateral view. (2) Genital plates (3) Ventral parts. (4) Aspis in dorsal view. 342 S. MAHUNKA Masthermannia hauseri sp. n. Figs 5-7 Material examined: Holotype: Madagascar, Prov. Antsiranana; 3.XI.1989; leg. B. Hauser (Mad-89/8); 2 paratypes from the same sample. Holotype and 1 paratype deposited in MHNG, 1 paratype (1786-PO-2009) in HNHM. Diagnosis: Rostrum rounded. Postbothridial condyles situated near to each other. Rostral setae dilated, lamellar and interlamellar setae T-shaped, exobothridial setae filiform. Sensillus bacilliform. Notogastral surface with five pairs of round elevations, ornamented with irregular foveolae. All setae T-shaped. Epimeral seta formula: 3 - 2 - 3 - 4, all except median setae setiform. Nine pairs of genital, two pairs of aggenital, two pairs of anal and three pairs of adanal setae. All legs monodactylous. Measurements: Length of body 418-433 /urn, width of body 191-202 p.m. Description: Prodorsum: Rostral part narrowing anteriorly. Prodorsal surface with anterolateral thickening, lamellar lines not visible. Median part punctate. Bothridium distinctly protruding, cup-shaped, sensillus comparatively short, bacilli- form. Its distal end roughened with a very narrow velum. Posterobothridial condyles triangular, situated close to each other. Their interdistance not longer than distance between lateral ridge. Rostral setae dilated basally (Fig. 7), narrowed anteriorly. Lamellar and interlamellar setae T-shaped, with very long, filiform branches. Notogaster. Dorsosejugal margin convex, one pair of posterolateral, and one pair of posterior projections present, these and three pairs of elevations on the antero- median surface well framed and separated from the other surface. All five pairs with round and smooth median field bearing one notogastral seta each. Remaining notogas- tral surface ornamented with irregular foveolae. Fifteen pairs of notogastral setae present, all T-shaped, size and length of the branches strongly variable. Their surfaces finely roughened. Gnathosoma: Typical for the family (Grandjean 1954). Setae h of the infra- capitulum very long, S-shaped. Ventral parts: Surface of the epimeres punctate. Epimeral setae (except the minute, spiniform la and 2a setae) long, setiform, distinctly roughened. Diagastry well visible. Ventral plate typical, as shown by Grandjean (1954), only ornamented with minor foveolae. Genito-anal setal formula: 9-2-2-3; four pairs of lyrifissures (ih, ips, iad and ion) present. Genital setae long, dilated basally. Aggenital setae also long, setiform. Anal setae minute, spiniform, adanal ones T-shaped, much smaller than notogastral setae. Legs: Well corresponding to Grandjean's drawings and descriptions, not studied in detail. Remarks: The new species is close to the type species (M. mammillaris Berlese, 1913) of the genus, which was redescribed by Grandjean (1954). It is distin- guishing from M. mammillaris by the interdistance of the posterobothridial condyles, the shorter sensillus, the absence of the wide epimeral setae (some bacilliform in M. mammillaris) and by the dilate genital and setiform, long aggenital setae (very short und minute in M. mammillaris). Etymology: Named after my friend Dr Bernd Hauser (Geneva), collector of this very interesting material, for his continuous help in my work on oribatids. ORIBATIDS FROM MADAGASCAR 343 Figs 5-7 Masthermannia hauseri sp. n. (5) Body in lateral view. (6) Body in ventral view. (7) Rostral setae. Caveremulus foliaceus sp. n. Figs 8-10 Material examined: Holotype: Madagascar, Prov. Antsiranana; 3. XI. 1989; leg. B. Hauser (Mad-89/8); 6 paratypes from the same sample. Holotype and 4 paratypes deposited in MHNG, 2 paratypes ( 1787-PO-2009) in HNHM. Diagnosis: Median part of prodorsal surface without costulae. Rostral setae arising from distinct tubercles, between the lamellar setae a weak transcostula present. Interlamellar setae minute, arising near to each other. Some small maculae present behind them. Sensillus very long, setiform. Notogaster with eight pairs of notogastral setae, among them three pairs large, phylliform. Epimeral setal formula 3 - 1 - 3 - 3. Six pairs of genital and two pairs of anal setae. Nine pairs of setae on ventral plate, mostly in preanal position. 344 S. MAHUNKA Figs 8-10 Caveremulus foliaceus sp. n. (8) Body in dorsal view. (9) Body in ventral view. (10) Anterior part of podosoma in lateral view. Measurements: Length of body 358-405 /urn. width of body 186-237 jjm. Description: Prodorsum: Rostral apex elongated, conical. Rostral setae arising laterally, from distinct tubercles, lamellar setae situated near to them, their bases connected by a transversal ridge. Both pairs of setae long, setiform. Median part of pro- dorsum smooth, in its posterior third 5-6 small, indistinct maculae present. In front of ORIBATIDS FROM MADAGASCAR 345 this sculpture a pair of minute interlamellar setae. Exobothridial setae fine, setiform, much longer than interlamellar ones. Bothridium round, a short crest directed inwards from it. Sensillus very long, ciliate, its distal end filiform, curved. Notogaster. Dorsosejugal suture concave. A pair of characteristic deep hollows present laterally. Setae c 2 short, setiform, arising at the anterior margin of the hollows. Only eight pairs of notogastral setae visible, three of them large, phylliform, covered by small cilia. Setae h\ very short (or broken), setae p represented only by their alveoli in a posteromarginal position. Lateral part of podosoma: Rostral part of prodorsum with deep hollow. Bothridium distinctly protruding from the prodorsal surface (Fig. 10). Pedotecta I large, pedotecta II-III absent. Ventral parts: Apodemes and borders weakly developed. Sternal ones only partly visible, sejugal ones well sclerotised, wide. Posterior border of epimeral region absent. Epimeral surface with irregular polygonal pattern. All epimeral setae long and thin, mostly setiform or filiform. Ventral plate with some round alveoli laterally, remaining surface smooth. Six pairs of genital setae arranged in longitudinal rows, two pairs of anal and three pairs of adanal setae well distinguishable, neotrichy present in aggenital region, consisting of six pairs setae. All setae simple, thin, epimeral setae much longer than the ventral ones. Remarks: See remarks after the next species. Etymology: The name refers to the form of the widened notogastral setae. Caveremulus salicinus sp. n. Figs 11-13 Material examined: Holotype: Madagascar. Prov. Antsiranana; 3. XI. 1989; leg. B. Hauser (Mad-89/8); 2 paratypes from the same sample. Holotype and 1 paratype deposited in MHNG, 1 paratype (1788-PO-2009) in HNHM. Diagnosis: Median part of prodorsal surface without costulae. Rostral and lamellar setae arising on small tubercles, lamellar setae located far from each other, between them no transcostula present. Interlamellar setae shorter than the preceding ones, arising also from small, separate tubercles. Sensillus very long, setiform. Notogaster with nine pairs of notogastral setae, six of them much longer than the others, three pairs in the posteromarginal position shortest. All notogastral setae more or less saliciform. Epimeral setal formula 3-1-3-3. Six pairs of genital and two pairs of anal setae. Ten pairs of setae on ventral plate, mostly in a preanal position. Measurements: Length of body 290-306 pm, width of body 157-173 pm. Description: Prodorsum: Rostral apex conical. Anterolateral part of prodorsum with short crest. All prodorsal setae, except exobothridial ones, arising from short tubercles, all setae setiform, finely ciliate. Exobothridial setae arising from short longitudinal crests, originating from anterior margins of bothridia. Interlamellar setae situated far from each other. Sensillus long, setiform, distinctly ciliate, distal part characteristically curved. Notogaster: Whole surface foveolate. Dorsosejugal suture straight. A pair of characteristic deep hollows present, these laterally connected by a distinct transversal line. Setae c 2 arising far medially from the hollows, short, setiform, much shorter than 346 S. MAHUNKA Figs 11-13 Caveremlus salicinus sp. n. (11) Body in dorsal view. (12) Body in ventral view. (13) Anterior part of podosoma in lateral view. setae la. Nine pairs of notogastral setae present, among them five pairs large, narrow, phylliform, resembling Salix leaves. Their surface smooth, a median vein well obser- vable. Setae p much shorter than the others. Lateral part of podosoma: Pedotectum I large, pedotecta II-III absent. Ventral parts: Apodemes and borders partly developed, a part of sternal apodemes reduced. Sternal apodemes forming a transversal band, ap. 3 also obser- ORIBATIDS FROM MADAGASCAR 347 vable. Posterior border of epimeral region absent. Epimeral surface smooth. All epimeral setae long and thin, mostly filiform, setae lc longest. Ventral plate smooth. Six pairs of genital setae arranged in longitudinal rows, two pairs of anal and three pairs of adanal setae distinguishable, neotrichy present in aggenital region, consisting of seven pairs of setae. All setae simple, thin, epimeral setae much longer than the ventral ones. Etymology: The new species is named after the form of its notogastral setae. Remarks: All species of the genus Caveremulus Mahunka, 1985 are known from Madagascar. They are well distinguishable from each other by the following key: 1 All notogastral setae, including setae c 2 wide, phylliform 2 Setae c 2 thin, setiform, much thinner than the other phylliform noto- gastral setae 3 2 All notogastral setae equal in length. Adanal setae short, thin, much shorter than the aggenital setae serratus (Mahunka, 1985) One pair of phylliform notogastral setae much smaller than the others. Adanal and aggenital setae equal in length cordisetus Mahunka, 1983 3 Interlamellar setae minute, arising very near to each other. Three pairs of well dilated notogastral setae present foliaceus sp. n. Interlamellar setae long, arising far from each other. Notogaster without wide phylliform setae, all setae saliciform salicinus sp. n. It is necessary to be aware that some Hymenobelba Balogh, 1962 species seem to be closely related to Caveremulus species. A further investigation is desirable. Tectocepheus velatus velatus (Michael, 1880) Remarks: The two studied specimens correspond to the nominate subspecies on the basis of the cuspis of the lamellae and the longitudinal pattern composed of lines in the interbothridial region, although according to Weigmann (2006) the longitudinal lines are characteristic of the subspecies sarekensis Trägardh, 1910. The position of the iad lyrifissures is also as in T. v. velatus, and so is the body shape. The only difference is the excavate anterior border of the aspis, but this alone does not justify the proposal of a new species or subspecies. Measurements: Length of body 250-255 /urn, width of body 146-150 ^urn. Ambrobates gen. n. Diagnosis: Belonging to the family Haplozetidae. Lamellae normal, anterior part of prelamella absent, sublamella present. Tutorium weakly developed. Sensillus short, head round. Pteromorphae large, partly movable. Nine pairs of small, notogastral setal alveoli and one pair of short distinct notogastral setae (/?j). Four pairs of round sacculi also well visible. Epimeral region without posterior border, sternal apodeme reduced. Epimeral setal formula: 3 - 1 -3-3. Three pairs of genital, one pair of aggenital, two pairs of anal and three pairs of adanal setae. All legs monodactylous. Type species: Ambrobates translamellatus sp. n. 348 S. MAHUNKA Remarks: The new genus is well charaterised by the form of the prelamella, the presence of a translamella, the reduced notogastral setae and by the three pairs of genital setae. This combination of features was previously unknown in the family. Ambrobates translamellatus sp. n. Figs 14-16 Material examined: Holotype: Madagascar, Prov. Antsiranana; 23.XI.1989; leg. B. Hauser (Mad-89/8); 10 paratypes from the same sample. Holotype and 6 paratypes deposited in MHNG, 4 paratypes (1789-PO-2009) in HNHM. Diagnosis: Rostral apex blunt. Lamellae not reaching the prodorsal margin. Translamella present. Sensillus short, its head round. Epimeral borders and apodemes weakly developed, all epimeral setae minute, simple. Genito-anal setal formula: 3-1 - 2 - 3. All legs monodactylous. Measurements: Length of body 241-295 pm, width of body 148-182 ^um. Description: Prodorsum: Rostral apex blunt at tip. Lamellae in normal position, narrow, with small rounded apices bearing the lamellar setae. A narrow, but distinctly developed translamella connecting the lamellar apices. Rostral, lamellar and interlamellar setae long, setiform; rostral setae densely pilose, lamellar and inter- lamellar setae moderately pilose. Sensillus conspicuously short, directed laterad, head round, surface with rounded alveoli. Notogaster. Dorsosejugal suture distinct. Pteromorpha large, movable, suture (hinge) sometimes hardly visible, especially in the anterior part. Nine pairs of alveoli (vestigial setae), one pair of true setae (/?j) and four pairs of minute, round sacculi present. All four pairs equal in size. Lateral part of podosoma: Pteromorpha large, rounded. Tutorium very simple. Pedotecta I normal, dorsal margin nearly straight. Exobothridial setae short, fine. Porose area al located very near to sublamella. Ventral parts: Apodemes and borders weakly developed, sternal apodeme absent or hardly observable. Ap. 2 and ap. sej. not reaching the genital aperture and not connected with each other. Ap. 4 absent. Epimeral surface smooth, only some large foveolae present. All epimeral setae short and fine. Three pairs of simple genital, one pair of aggenital, two pairs of anal, three pairs of adanal setae present. Setae ad^ in preanal position. Lyrifissures iad located near the anal aperture. Legs: All legs monodactylous. Remarks: See the remarks after the genus description. Etymology: The name refers to the presence a distinct translamella. Vilhenabates ambohitra sp. n. Figs 17-19 Material examined: Holotype: Madagascar, Prov. Antsiranana; 23. XI. 1989; leg. B. Hauser (Mad-89/8); 10 paratypes from the same sample. Holotype and 6 paratypes deposited in MHNG, 4 paratypes (1790-PO-2008) in HNHM. Diagnosis: Rostral apex tripartite. Lamellae running laterally, their apices bearing lammellar setae. Sensillus extraordinarily long, directed backwards, head fusiform. Pteromorphae movable, sejugal margin convex, with a pair of porose areas. Notogaster with ten pairs of alveoli and with three pairs of porose areas. Epimeral ORIBATIDS FROM MADAGASCAR 349 Figs 14-16 Ambrobates translamellatus gen. n., sp. n. (14) Body in dorsal view. (15) Body in ventral view. (16) Anterior part of podosoma in lateral view. borders and apodemes weakly developed, partly absent. Genito-anal setal formula: 5 - 1 - 2 - 3. All legs monodactylous. Measurements: Length of body 291-312 /jm, width of body 206-221 /um. Description: Prodorsum: Rostral part wide, with trifurcate rostral apex. Lamellae long, narrow, slightly dilated distally, directed inwards, lamellar setae located 350 S. MAHUNKA Figs 17-19 Vilhenabates ambohitra sp. n. (17) Body in dorsal view. (18) Body in ventral view. (19) Anterior part of podosoma in lateral view. ORIBATIDS FROM MADAGASCAR 351 on their apices. Prelamellae absent, sublamellae short, weakly developed. All prodorsal setae comparatively short, finely ciliate. Sensillus very long, directed backwards, with small, lanceolate head, ciliate along the whole length. One pair of very small dorso- sejugal porose areas present basally, near the lamellae. Notogaster. Dorsosejugal suture well developed, convex. Pteromorphae large, round, tongue-shaped. Three pairs of small, round porose areae well observable. Ten pairs of alveoli of notogastral setae, probably two pairs in posteromarginal position. Lateral part of podosoma (Fig. 19): Tutorium short, without true apex. Pedotectum 1 long, with straight margin. A small porose area lamellaris {la) visible. Circumpedal carina long, reaching lateral margin of ventral plate. Ventral parts: Apodemes and borders weakly developed. Bo. 2 and bo. sej. reaching the genital aperture and connected with each other. Epimeral surface smooth, without pattern. All epimeral setae short, not ciliate. Bo. 4 absent. Five pairs of simple genital, one pair of aggenital, two pairs of anal and three pairs of short adanal setae present. Setae ad^ and lyrifissures iad in preanal position, both located very near to the anterior corner of the anal aperture. Legs: All legs monodactylous. Femora of leg II-IV with distinct blade-like shape basally. Remarks: The new species is well distinguishable from the previously known Vilhenabates species by the shape of the rostral apex. Etymology: Named after the type locality of this species on Madagascar, the Pare National Montagne d'Ambre (= Ambohitra). ACKNOWLEDGEMENTS This research work was partly sponsored by the Hungarian Scientific Research Fund (OTKA 45889 and 72744). I thank Dr Bernd Hauser for collecting this very interesting material, Dr Peter Schwendinger for his help in preparing this manuscript, and my wife, Mrs Luise Mahunka-Papp, for doing the drawings. I also like to thank Dr Tibor Fuisz for translating part of the English text of this paper. REFERENCES Grandjean, F. 1954. Posthermannia nematophora n. g., n. sp. (Acarien, Oribate). Revue française d'Entomologie 21 : 298-3 1 1 . Grandjean, F. 1965. Complément a mon travail de 1953 sur la classification des Oribates. Acarologia 7: 713-734. Mahunka, S. 1993. Oribatids from Madagascar I (Acari: Oribatida). New and interesting mites from the Geneva Museum LXXVI. Revue suisse de Zoologie 100 (2): 289-315. Mahunka, S. 1994. Oribatids from Madagascar II (Acari: Oribatida). (New and interesting mites from the Geneva Museum LXXIX). Revue suisse de Zoologie 101 (1): 47-88. Mahunka, S. 1997. Oribatids from Madagascar III (Acari: Oribatida). (Acarologia Genavensia LXXXIII). Revue suisse de Zoologie 104 (1): 1 15-170. Mahunka, S. 2002. A survey of the Oribatida fauna of Madagascar (Acari: Oribatida). Folia entomologica hungarica 63: 5-16. Mahunka, S. 2009. Oribatid mites from the Vohimana reserve (Madagascar) (Acari: Oribatida). I. Acta Zoologica Academiae Scientiarum Hungaricae 55(2): 89-122. 352 s - MAHUNKA Mahunka, S. & Mahunka-Papp, L. 2001. Oribatids from Switzerland V (Acari: Oribatida: Suctobelbidae 2). (Acawlogica Genavensia XCVII). Revue suisse de Zoologie 108: 355-385. Mahunka, S. & Zombori, L. 1985. The variability of some morphological features in Oribatid mites. Folia entomologica Hungarica 46: 115-128. Marshall, V. G., Reeves, R. M. & Norton, R. A. 1987. Catalogue of Oribatida (Acari) of conti- nental United States and Canada. Memoirs of the Entomological Society of Canada 139: I-VI, 1-148. Niedbala, W. 1985. Essai critique sur Mesoplophora (Acari, Oribatida, Mesoplophoridae). Annales zoologici 39 (4): 93-1 17. NlEDBALA, W. 2001. Study on the diversity of ptyctimous mites (Acari, Oribatida) and quest for centres of its origin: the fauna of the Ethiopian Region. Monographs of the Upper Silensian Museum 3 : 1 -245 . Niedbala, W. 2004. Zoogeography of the ptyctimous mites (Acari: Oribatida) of Madagascar and other eastern Africa islands. International Journal of Tropical Insect Science 24 (4): 330-335. Norton, R. A., Alberti, G., Weigmann, G. & Woas, S. 1997. Porose integumental organs of oribatid mites (Acari, Otibatida). 1. Overview of types and distribution. Zoologica 146: 1-33. SubÎas, L. S. 2004. Listado sistematico, sinonimico y biogeogrâfico de los âcaros oribâtidos (Acariformes, Oribatida) del Mundo (1758-2002). Graellsia 60: 3-305. Subîas, L. S. 2008. Listado sistematico, sinonimico y biogeogrâfico de los âcaros oribâtidos (Acariformes, Oribatida) del Mundo (exepto fósiles) (originally published in Graellsia 60: 3-305, 2004, updated in May 2008, in pdf), 540 pp. Weigmann, G. 2006. Hornmilben (Oribatida). Die Tierwelt Deutschlands 76: 1-520. Woas, S. 2002. 4. 1. Acari: Oribatida (21-291). Adis, J. (ed.). Amazonian Arachnida and Myriapoda. Pensoft Publishers, Sofia-Moscow, 590 pp. Revue suisse de Zoologie 1 16 (3-4): 353-358; décembre 2009 Species of Hybos Meigen from Ningxia, Palaearctic China (Diptera, Hybotidae) Weihai LI 1 and Ding YANG 2 - 3 1 Department of Plant Protection, Henan Institute of Science and Technology, Xinxiang, Henan 453003, China. E-mail: lwh7969@163.com 2 Department of Entomology, China Agricultural University, 2 Yuanmingyuan West Road, Beijing 100193, China. E-mail: dyangcau@yahoo.com.cn 3 Correspondence author Species of Hybos Meigen from Ningxia, Palaearctic China (Diptera, Hybotidae). - Only one species of the genus Hybos was known to occur in Ningxia of Northwest China, Palaearctic Region. Here five species are reported. One species, Hybos liupanshanus sp. nov., is described as new to science. A key to the five species of the genus from Ningxia is presented for the first time. Keywords: Diptera - Hybotidae - Hybos - new species - Ningxia. INTRODUCTION Within the Hybotidae Hybos Meigen is characterized by the following traits: long spine-like proboscis, anal cell longer than basal cell, Rs rather short, R 4+5 and Mj divergent apically (Chvâla, 1983; Yang & Yang, 2004). It is distributed worldwide with 164 known species, of which 12 species are from the Palaearctic Region and 130 species from the Oriental Region (Yang et al., 2007). The species of Hybos from China were revised by Yang & Yang (2004) who mentioned 85 known species. The major references dealing with Hybos in the Old World are Brunetti (1920), Frey (1953), Smith (1965), Saigusa (1963, 1965), Chvâla (1983), Yang & Yang (2004), and Yang et al. (2006). Ningxia is a semi-humid to arid region with a temperate climate and belongs to Northwest China, Palaearctic Region. Up to now, only one Hybos species, Hybos grossipes (Linnaeus, 1767), has been described from this region (Yang & Yang, 2004). In the present paper, four species of Hybos are added to the fauna of Ningxia, supple- menting Yang & Yang (2004). One species is described as new to science. A key to separate these five species of the genus from Ningxia is presented for the first time. MATERIAL AND METHODS The specimens for this study are deposited in the following collections: CAU = Entomological Museum of China Agricultural University, Beijing, China. MHNG = Muséum d'histoire naturelle, Genève, Switzerland. Manuscript accepted 16.06.2009 354 W. LI & D. YANG Basic terminology follows McAlpine (1981) and Steyskal & Knutson (1981). The following abbreviations are used: acr = acrostichal, ad = anterodorsal, av = ante- roventral, dc = dorsocentral, h = humeral, npl = notopleural, oc = ocellar, pd = postero- dorsal, prese = prescutellar, psa = postalar, pv = postero ventral, v = ventral. KEY TO SPECIES (MALES) OF HYBOS FROM NINGXIA 1 Legs partly or mostly yellow 2 Legs entirely black 4 2 Fore and mid legs mostly black 3 Fore and mid legs mostly yellow Hybos liupanshanus sp. n. 3 Hind tibia with 2 dorsal setae at middle . Hybos hubeiensis Yang et Yang, 1 99 1 Hind tibia without dorsal setae at middle . Hybos joneensis Yang er Yang, 1988 4 Hind coxa with 3 spine-like anterior setae at tip; fore and mid tibiae and tarsomere 1 with long setulae Hybos grossipes (Linnaeus, 1767) Hind coxa without spine-like anterior setae at tip; only fore tarsomere 1 with long setulae Hybos wudanganus Yang e?Yang, 1991 TAXONOMIC PART Hybos liupanshanus sp. n. Figs 1-4 Material: Holotype male, Ningxia, Longde, Liupanshan Mountain, Sutai (2100 m), 24. VI. 2008, Jingxian Liu (CAU). Paratypes 3 males 1 female, same data as holotype (CAU & MHNG). Etymology: The species is named after the type locality Liupanshan. Diagnosis: Setulae and setae on body pale. Legs yellow except hind coxa dark yellow, hind femur black, hind tibia partly blackish; all tarsomeres 3-5 black (not in dried specimens: tarsomeres yellow or brownish). Hypandrium with long narrow api- cal portion and a finger-like lateral process near base. Description: Male. Body length 4.7-4.9 mm, wing length 4.7-5.0 mm. Head: black, pale gray microtrichose. Eyes brownish, contiguous on frons (but narrowly separated in dried specimens); upper facets indistinctly enlarged. Setae and setulae on head pale; ocellar tubercle weak, with one pair of long brownish oc. Antenna black; scape bare; pedicel with a circle of subapical setulae; first flagellomere without dorsal setulae, about 3 times as long as wide; arista black, about 3.5 times as long as first flagellomere, short pubescent except apical 1/4 thin and bare. Proboscis black; palpus black with 4 long ventral setulae. Thorax: black, pale grey microtrichose. Setae and setulae pale; postpronotal lobe with pale setulae; h absent, 2 npl (anterior npl short), acr more or less in two rows, 1 distinct posterior dc and 8 setulae anteriad, 2 weak prese, 1 long psa; scutellum with 6 or 8 short lateral setulae and 2 long subapical setae. Legs yellow except hind leg with coxa varying from yellow to blackish, femur black, tibia brown to blackish; tarsomeres 3-5 yellow or slightly brownish, but rarely tarsomeres 1-2 also dark brown. Setae and setulae on legs pale except tibiae and tarsi with some blackish setulae. Hind trochanter with 2 short black ventral spines apically. HYBOS FROM NINGXIA 355 Figs 1-4 Hybos liupanshanus sp. n. (male). (1) genitalia, dorsal view. (2) right surstylus. (3) left surstylus. (4) hypandrium, ventral view. 356 W. LI & D. YANG All femora with long setulae. All tibiae and fore and mid tarsomeres 1-2 with some long and nearly erect setulae. Fore femur 1.1 times as wide as mid femur; mid femur with 1 blackish preapical anterior seta; hind femur 2.4 times as wide as mid femur, with 1 brownish yellow preapical anterior seta, and with 9 av (longer than v and pv), 15 v (3 sparse v on basal 1/2 and 12 dense v on apical 1/2), 5 pv on basal 1/3, which are spine-like and inserted on weak tubercles. Fore tibia apically with 1 blackish ad and 1 brownish yellow av; mid tibia with 1 long blackish ad at base and 1 long blackish ad at middle, apically with 1 brownish yellow av; hind tibia apically with 1 brownish yellow av. Fore tarsomere 1 with 1 black av and 1 black pv which are located at same level near extreme base; mid tarsomere 1 with two rows of black v; hind tarsomeres 1-2 with short black ventral spines. Wing slightly tinged with grayish; stigma dark brown; veins dark brown, R 4+5 and M^ divergent apically. Squama pale yellow, bordered with pale setulae. Halter pale yellow. Abdomen: weakly curved downward, blackish, grey microtrichose. Setulae and setae on abdomen pale, but those on hypopygium sometimes black. Male genitalia (Figs 1-4): left epandrial lobe with convex inner margin, left surstylus slightly constricted basally and obtuse apically; right epandrial lobe with slightly concave inner margin, right surstylus short and broad, with a short finger-like inner process at base; hypandrium much longer than wide, basally wide, apically narrow and slightly curved, and with a finger-like lateral process near base. Female: Body length 3.8 mm, wing length 4.6 mm. Similar to male, but hind trochanter without short black ventral spines apically, fore and mid tarsomere 2 without long and nearly erect setulae. Distribution: China (Ningxia). Remarks: The new species is similar to Hybos minutus Yang & Yang, 1997 from Hubei in the coloration of the legs, but it can be easily separated from the latter by having the first flagellomere without any dorsal seta, hind femur with 3 rows of ventral spines and hypandrium much longer than wide without the furcate apex. In H. minutus, the first flagellomere has 1 dorsal seta, the hind femur has one row of ventral spines, the hypandrium is slightly longer than wide, with a furcate apex (Yang & Yang, 2004). Hybos grossipes (Linnaeus, 1767) Musca grossipes Linnaeus, 1767: 988. Type locality: "Europa". Material: 2 males 1 female, Ningxia. Jingyuan, Liangdianxia 2000 m, 4. VII. 2008, lingxian Liu (CAU). Distribution: China (Jilin, Ningxia, Gansu, Inner Mongolia, Shanxi, Shaanxi, Hebei, Henan, Sichuan); Europe. Hybos hubeiensis Yang et Yang, 1991 Hybos hubeiensis Yang et Yang, 1991: 3. Type locality: China: Hubei, Wudangshan Mountain. HYBOS FROM NINGXIA 357 Material: 2 males 1 female, Ningxia, Jingyuan, Longtan 1880 m, 5. VII. 2008, Jingxian Liu (CAU). Distribution: China (Ningxia, Henan, Gansu, Hubei). Hybos joneensis Yang et Yang, 1988 Hybos joneensis Yang er Yang, 1988: 284. Type locality: China: Gansu, Jone. Material: 1 male 1 female, Ningxia, Jingyuan, Heshangpu 2000 m, 26. VI. 2008, Jingxian Liu (CAU). Distribution: China (Ningxia, Gansu). Hybos wudanganus Yang et Yang, 1991 Hybos wudanganus Yang et Yang, 1991: 5. Type locality: Hubei, Wudangshan Mountain. Material: 1 male 1 female, Ningxia, Jingyuan, Longtan 1880 m, 5. VII. 2008, Jingxian Liu (CAU). Distribution: China (Ningxia, Henan, Hubei). ACKNOWLEDGEMENTS Our sincere thanks are due to Dr. Jingxian Liu (Zhejiang University, Hangzhou) and Ms. Weina Cui for their help during the study. The research was funded by the National Natural Science Foundation of China (No. 30770259, No. 30225009). REFERENCES Brunetti, E. 1920. Diptera Brachycera Vol. 1. The Fauna of British India, including Ceylon & Burma. London, 401 pp. Chvâla, M. 1983. The Empidoidea (Diptera) of Fennoscandia and Denmark. II. General Part. The families Hybotidae, Atelestidae and Microphoridae. Fauna Entomologica Scandinavica 12: 279 pp. Scandinavian Science Press, Kopenhagen. Frey, R. 1953. Studien über ostasiatische Dipteren. II. Hybotinae, Ocydromiinae, Hormopeza Zett. Notulae Entomologicae 33: 57-71. Linnaeus, C. 1767. Systema naturae per regna tria naturae. Ed. 12 (rev). Vol. 1, Pt. 2, pp. 533- 1327. Holmiae. McAlpine, J. F. 1981. Morphology and terminology - adults, (pp. 9-63). In: Mcalpine. J. F. Peterson, B. V, Shewell, G. E., Teskey, H. J., Vockeroth, J. R. & Wood, D. M. (eds). Manual of Nearctic Diptera. Vol. 1. Research Branch, Agriculture Canada, Monograph 27: 1-674. Saigusa, T. 1963. Systematic studies of the genus Hybos in Japan I. New species with yellowish legs (Diptera, Empididae). Sieboldia 3(1): 97-104. Saigusa, T. 1965. Studies on the Formosan Empididae collected by Professor T. Shirôzu (Diptera, Brachycera). Special Bulletin of Lepidoptera Society in Japan 1: 180-196. Smith, K. G. V. 1965. Diptera from Nepal: Empididae. Bulletin of the British Museum (Natural History), Entomology 17(2): 61-112. Steyskal, G. C. & Knutson, L. V 1981. 47. Empididae (pp 607-624). In: Mcalpine, J. F. Peterson, B. V, Shewell, G. E., Teskey, H. J., Vockeroth, J. R. & Wood, D. M. (eds). Manual of Nearctic Diptera. Vol. 1. Research Branch, Agriculture Canada, Monograph 27: 1-674. 358 W. LI & D. YANG Yang, C. & Yang, D. 1991. New species of Hybos Meigen from Hubei (Diptera: Empididae). Journal of Hubei University (Natural Science) 13(1): 1-8. Yang, D. & Yang, C. 1988. New species of Hybos Meigen from China (Diptera: Empididae). Acta Agriculturae Universitatis Pekinensis 14(3): 282-287. Yang, D. & Yang, C. 2004. Fauna Sinica Insecta Vol. 34. Diptera, Empididae, Hemero- dromiinae Hybotinae. Science Press, Beijing, 329 pp. Yang, D., Merz, B. & Grootaert, P. 2006. New yellow-legged Hybos from Nanling, Guangdong, China (Diptera, Empidoidea, Hybotidae). Revue suisse de Zoologie 1 13(4): 797-806. Yang, D., Zhang, K. Y, Yao, G. & Zhang, J. H. 2007. World catalog of Empididae (Insecta: Diptera). China Agricultural University Press, Beijing, 599 pp. Revue suisse de Zoologie 116 (3-4): 359-377; décembre 2009 Variations in sex ratio studied in a single breeding population of an endangered salamander: a long-term study Michael R. WARBURG Dept. of Biology, Technion-Israel Institute of Technology, Haifa 32000, Israel. E-mail: Warburg@tx.technion.ac.il Variations in sex ratio studied in a single breeding population of an endangered salamander: a long-term study. - A small population of a rare, xeric-inhabiting salamander Salamandra infraimmaculata Martens, 1885, was studied throughout 25 years at the breeding ponds on Mt. Carmel, Israel. The breeding period extended between October and January but most females visited the ponds during November and December. The number of salamanders fluctuated during the study period not showing any particular pattern. The sex ratio was male-biased during most of this period while female numbers dropped during the last few years. Only such long-term observations can illustrate this point in perspective of time leading to the conclusion that the observed drop in female numbers may eventually effect a decline in the species' presence in this fringe habitat. The subject is discussed and reviewed and a general comparison with variation in sex ratio in other urodele species is presented. Keyword: Amphibian decline - Endangered species - Long-term study - Salamandra - Sex ratio. INTRODUCTION The salamander, Salamandra infraimmaculata Martens, 1885, is a rare and pro- tected species inhabiting mountainous regions in northern Israel where it is limited in its distribution to three disjunct metapopulations. The main population is located in the mountains of the Western and Central Galilee (Degani & Warburg, 1978). In addition there are two smaller ones at about 50 km distance from, and unconnected to the main area. One is in the north-eastern part of Israel at the foot of Mt. Hermon at Tel Dan (Degani & Mendelssohn, 1982), the other is located southwest of the main area in the northern part of Mt. Carmel Warburg (1986, 1994). This metapopulation of salamanders on Mt. Carmel comprises the southeastern limit of the species' Palaearctic distribution. Therefore, it inhabits a fringe area where conditions are optimal only part of the time to the animals. It seems possible that sala- manders in this area may have to cope, at times, with unsuitable conditions not encountered by other populations inhabiting more favorable environments in the Galilee mountains (Degani & Mendelssohn, 1982; Degani & Warburg, 1978), or by other Salamandra species in the center of the species' distribution in Central Europe (Joly, 1968; Feldmann & Klewen, 1981; Klewen, 1985, 1988; Thiesmeier, 2004). Manoscript accepted 31.07.2008 360 M. R. WARBURG The adult salamander is largely a terrestrial animal returning to water only when mature at the age of 3-4 years, and then only the females for a few hours to breed (Warburg, 2009). Males usually remain out of water even during the breeding period as mating takes place on land. Its aquatic life during the larval period lasts only three to four months. Since as adult, the female enters water only for a few hours, the species' aquatic life totals about 1.25% of its lifetime. The uniqueness of this species is because of two main of reasons: 1 . It occupies a fringe habitat at the edge of its Palaearctic distribution. 2. It has to survive the weather in a xeric Mediterranean region characterized by a rather short rainy season unpredictable in its duration and magnitude between October and January when about 66% of the average annual rain falls. On Mt. Carmel the average annual rainfall ranged between 440- 1 1 60 mm during the 25 years of study (averaging 690 mm annually). Since the rainy season is followed by eight months of hot-dry weather, breeding has to take place by January at the latest or the metamorphosing larvae will die (Cohen et al., 2005, 2006). In these studies it was shown that through both differential growth and cannibalism, half-sib larval cohorts facilitate spacing-out the timing of metamorphosis resulting in spacing-out of both temporal and spatial dispersal in juvenile S. infraimmaculata (in preparation). This is an important mechanism for survival eventually enabling colonization of new breeding ponds (Degani et al., 2007). This long-term study was not planned as such but started as a family project during which the salamanders were observed in their breeding rock-pools on Mt. Carmel, on cold-rainy nights. It developed into this long-term study as ever-more questions arose regarding different aspects of their life history. The original objective of this long-term study was to learn about the life history and phenology of S. infraimmaculata by monitoring adult salamanders during the breeding season in a single breeding metapopulation. It later evolved into a study on breeding pond tenacity (Warburg, 2006), longevity (Warburg, 2007a), phenology (Warburg, 2007b), recapture rates (Warburg, 2008a), dimensions (Warburg, 2008b), growth (Warburg, 2008c), and reproduction (Warburg, 2009), as these salamanders were observed over a long period (25 years). Consequently new questions arose: does the sex ratio studied in this single population show changes over the years? When can a drop in the number of females be interpreted as a decline? MATERIALS AND METHODS The study area was located 12 km south of Haifa on the top of Mt. Carmel towards its western slopes. The study site (about 60x100 m) contains four shallow rock-pools which are one of the main breeding sites for the salamanders in this area. There are a few other far apart breeding ponds that are used by the Mt. Carmel Salamandra metapopulation. The average annual rainfall based on 25 years of meteo- rological observations is 690 mm. Since the breeding season when adult salamanders emerge from their aestivation sites starts earliest in September (generally October) and lasts till January, I used data only for these months. The study period lasted from 1974 to 1998 with the exception of one breeding season (1990/91) altogether 24 years of observations. SALAMANDER SEX RATIO 361 Adult Salamanders were observed near their breeding sites on stormy winter nights throughout the entire breeding season for 10-12 weeks starting at the onset of the rainy season (October or November), continuing until mid-January. The animals were identified individually by their typical yellow patterns on a black background on the dorsal side of the salamander. These hardly ever change throughout its lifetime (Warburg, 2006 see Fig.l, 2007a, b, 2008a). Consequently the salamanders could easily be identified individually by their photographs throughout the entire study period. Marking animals by toe clipping or subcutaneous insertion of PIT tags were not necessary especially since the first method is only adequate for short- term monitoring due to the high rate of regeneration in amphibians especially in urodeles, and both techniques involve rough handling and molesting this rare, endangered species. The sex was determined by cloacal examination (see Degani & Warburg, 1978). In this study I have used three different ways to express sex ratio: 1. Male/female ratio when male is always one therefore females may be either larger or smaller than one. 2. I have also used percentage (%) to express the ratio between the genders. 3. For comparing with other data I have also used the male/female ratio where male numbers are divided by the females'. The animals were then photographed and finally released back to their habitat either during the same night or on the following one. For the partial review on the subject of changes in sex ratio in urodeles, I was limited in the number of papers I could use since many of the papers describing popu- lation structure or dynamics did not provide the original data in the form of Tables but rather in Figures. The urodelan species cited in the text and Tables are listed in the Appendix. It is of interest to note here that the earliest paper on this subject providing quantitative data was published in this journal (Revue suisse de Zoologie 1952) by Fritz Ernst of Zurich Anatomical Institute. For statistical analysis I have used standard regression analyses since the number of salamanders sampled was rather small. RESULTS One hundred and thirty salamanders were captured and about 50% of them were recaptured during the 25 years of study. This paper is based on both captured and recaptured (more than once up to 40 times in males, see Warburg 2006, 2007b, 2008) salamanders. The number of both males and females did not show significant relation with either years (1974-1998 Fig. 1 A) or rain (Fig.l B). Nor was there a significant relation between years and rainfall (Fig.l C). During the long study the sex ratio (%) fluctuated (Fig. 2 A). Thus, during the first three years (1974-76) there was an increase in the relative number of females which became greater than that of males in the 3rd year (1976). After this, the numbers of males was always greater than those of females. In all but three years (1976, 1980, 362 M. R. WARBURG Number of male (line) and females (columns) captured 30 25 (0 E 15 à 10 k\ DO UUDUUD 12 10 8 e "«5 6 1 4 d z 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 A Years Number of male (triangle) and female (qudrates) salamanders as related to rainfall 30 25 A 8 20 « E 15 A A A A A A A A * A o io 5 A ■ A m ■ A m m • ■ A ■ É ■ 1 i m A A« " 100 200 300 400 500 600 700 B Rain (mm) Relationship between rainfall during the breeding season (X-l) and years 700 600 -g 500 E 400 = 300 £ 200 lull 100 o I. I, III J I f f f f ,«* d?* d?* d^ 0?* els' dP" dp <& C Year FlG.l Number of male and female salamanders captured over the years (A) and in relation to rain (B), and between rain and years (C). SALAMANDER SEX RATIO 363 1982) males comprised more than half the population and in most years there were more than 70% males (Fig. 2 A). Sex ratio (M:F ratio) did not relate significantly to years (Fig. 2. C). Sex ratio ranged from 1:0.12 (in 1985) to 1:2 (in 1976) averaging 1:0.4 in 24 years of observations (Table 1). There was no significant relationship between number of males and females (Fig. 2 B). When the 25-year study period was divided into six 4-year periods (excluding the year 1990 when the site could not be visited during the breeding season), no signi- ficant relation was noticed in either females (R 2 = 0.0495 Fig. 3 A), males (R 2 = 0.634 Fig. 3 B) or rainfall (R 2 = 0.1 127 Fig. 3 C). However, the M:F ratio was related signi- ficantly when arranged in 4-year periods (R 2 =0.9527) becoming increasingly male biased during the last two 4-year periods (1991-1998) (Fig. 4). The M:F ratio drops gradually from 1:0.64 (M:F) during the first eight years of observation (1974-1981), to 1:0.26 in later years (1995-1998) showing a decline in the percentage of females in this breeding population (Table 2). DISCUSSION The data presented here where assembled over a long period of time during which profound changes in the sex ratio within a single salamander population were noticeable. Two main points need to be discussed: 1. Are there any similar data on urodelan sex ratio that can be compared? 2. How can these temporal variations in the sex ratio be explained? Regarding comparable data: Similar studies on sex ratio in the genus Salamandra show that the sex ratio ranged between 1:0.29 to 1:1.14 M:F ratio (Table 3). In only three of the 12 studies reviewed, the sex ratio was female-biased. On the other hand, in one 5 -year long study of S. s. terre stris, there was a significant relation (R 2 =0.7174) between M:/F ratio and years. The proportion of males continuously increased with years (from 2.31 to 3.52 M:F ratio, see Klewen, 1985). Reviewing 5 1 studies on 34 urodele species studying sex ratio (Table 4) the sex ratio in 22 species was about 1:1 (M:F). In eight Ambystoma spp. and in five Plethodon spp. sex ratio ranged between 1:0.35 to 1:3. In seven Triturus spp. sex ratio ranged between 1:0.58 to 1:3.3. The reason for a higher proportion of males in the populations close to the breeding sites could be because of early maturity in males (Flageole & Leclair, 1992; Leclair & Caetano, 1997). Moreover, their greater activity extending over much of the breeding season arriving at the breeding ponds earlier than females and leaving later, can also contribute to apparent unbalanced sex ratios in adult urodeles (Caetano & Leclair, 1 996) thereby explaining some of the spatial and temporal variation in the sex ratio of newts during the season as was suggested by Arntzen (2002). Observations on S. salamandra (Linnaeus, 1758) similarly indicated the early activity of males (Warburg, 1994), and Beneski, Zalisko & Larsen (1986) found more males than female A. macrodactylum Baird, 1849 at the beginning of the breeding period, the females only being active later on. Previous studies were concerned with both spatial and temporal changes in sex ratio. 364 M. R. WARBURG Salamandra sex ratio (%) lllll'll I 'II' Sex ratio 3 O O O O O < 1974 1976 1978 1980 982 1984 1986 1988 1990 1992 1994 1996 1998 A Years ■ Females a Males 30 25 S 20 io E 15 é io 5 Male:Female ratio R 2 =03186 logarithmic ♦ 10 12 No. females Sex ratio (M:F) in S. infraimmaculata during the study period 8 R 2 =03514 exponential 7 7 7 ♦ ♦ s t 6 ♦ S w 5 O '•§ 4 2 •> 4 ♦ , 3.4 ♦ V 5 3S 3.33 3 2.75 ♦ ♦ ♦ ♦ x 3 2.33 2.5 ♦ ♦ 2.33 3 25 • (0 2 1 o \|3 1 l 7 * ♦ 0.5 1 At 1 71 1.91 1 43 A « ♦ * 143 1 1.11 14J ♦ 1970 1975 1980 1985 1990 1995 2000 c Years Fig. 2 Sex ratio (M:F) (A), the relationship between the number of males and of females (B) and changes in sex ration during the study period (C). SALAMANDER SEX RATIO 365 Table 1. Sex ratio. Year Males Females M:F 1974 7 3 1 : 0.43 1975 4 3 1 0.75 1976 1 2 1 2 1977 5 3 1 0.6 1978 5 2 1 0.4 1979 4 1 1 0.25 1980 6 6 1 1 1981 10 7 1 0.7 1982 9 10 1 1.1 1983 18 6 1 0.33 1984 12 7 1 0.58 1985 24 3 1 0.12 1986 22 8 1 0.36 1987 10 7 1 0.7 1988 26 11 1 0.42 1989 17 5 1 0.29 ==== the site was not visited 1991 15 2 1 0.13 1992 15 6 1 0.4 1993 8 3 1 0.37 1994 15 4 1 0.27 1995 7 1 1 0.14 1996 14 4 1 0.28 1997 14 4 1 0.28 1998 4 1 1 0.25 272 109 1 :0.4 Table 2. Sex ratio calculated in 4-year periods. Nos captured Years M F M:F 1974-77 17 11 1 : 0.65 1978-81 25 16 1 : 0.64 1982-85 63 26 1 : 0.41 1986-89 75 31 1 : 0.41 1991-94 53 15 1 : 0.28 1995-98 39 10 1 : 0.26 Total 272 109 1 :0.4 Spatial changes The sex ratio in salamanders varied between localities. In Salamandra atra Laurenti, 1768 it ranged between 1:0.49 and 1:0.84 M:F ratio (Klewen, 1986), in Batrachoseps attenuatus (Eschscholtz, 1833) M:F sex ratio ranged between 0.17-1.78, and in Aneides lugubris between 0.73-1 depending on the locality (Anderson, 1960). Likewise, in Notophthalmus viridescens (Rafinesque, 1820), differences in the sex ratio were found in populations from different localities ranging from 1:0.6 to 1:2 M:F 366 M. R. WARBURG 35 30 g 25 | 20 Ä "S 1 5 È 10 5 Number of females arranged in 4-year periods R 2 =0.0495 31 26 10 16 ■ 74-77 78-81 82-85 86-89 91-94 95-98 4-year periods Number of mates arranged in 4-year periods R 2 =0.634 80 70 60 in | 50 (0 I 40 O Ö 30 Z 20 10 D 75 63 53 39 74-77 78-81 It 82-85 86-89 4-year periods 91-94 95-98 Rainfall arranged In 4-year periods 1800 R 2 =0.1127 1678 1600 1610 1523 1400 1138 _1200 1100 836 | 1000 c" 800 2 600 400 200 74-77 78-81 82-85 86-89 91-94 95-98 C 4-year periods Fig. 3 Number of males, females and rainfall arranged in 4-year periods. SALAMANDER SEX RATIO 367 Table 3. Sex ratios in Salamandra. Species M:F Source Salamandra salamandra 1:0.55 S. salamandra 1:1 S. salamandra 1:0.6 S. salamandra 1:0.29 S. salamandra 1:1.08 S. s. terresîris 1 :0.55 S. s. terresîris 1:0.38 S. s. terrestris 1:0.81 S. s. gallaica 1:1.04 S. salamandra 1:1.14 S. infraimmaculata 1:0.41 S. atra 1:0.67 S. atra atra 1 : 1 Geiler, 1974 Degani & Warburg, 1978 Degani & Mendelssohn, 1982 Warburg, 1994 Rebelo & Leclair, 2003 Feldmann & Klewen, 1981 Klewen, 1985 Seifert (see Thiesmeier, 2004) Rebelo & Caetano, 1995 Catenazzi, 1998 (see Thiesmeier, 2004) This study Klewen, 1986 Luiselli et al, 2001 ratio Gill (1978, a,b) and 1:0.53 to 1:1.7 M:F ratio (Leclair & Caetano, 1997). Similar differences in sex ratios among populations of this species were also described by Hurlbert (1969). In the eastern Galilee population of S. salamandra, Degani (1980) found 161 specimens 54.4% of them were males, and in a later study 55% were males (Degani & Mendelssohn, 1982). Sex ratio in Desmognathus ochrophaeus Cope, 1859 changed with altitude: at high elevation it was 1:0.59 M:F ratio and at low elevation 1:0.74 M:F ratio (Tilley, 1973). Seasonal changes Seasonal changes in the sex ratio in a population of Desmognathus fuscus (Green, 1818) were studied by Danstedt (1975). He stressed that they may be a conse- quence of a decline in the frequency of females in the area he studied. Similar seasonal differences in sex ratio of Triturus vulgaris (Linnaeus, 1758) were described: in autumn and winter it was 1:2 M:F, in spring and summer 1:0.77 M:F. The terrestrial phase of the newt showed a 1:3.3 M:F averaging 1:1.26 (Griffiths, 1984). In breeding newts Harrison et al. (1983) found a sex ratio of 1:1.1 in Triturus helveticus Razumowski, 1789 and 1:2.6 in T. vulgaris. The sex ratio in outward migrating newts from the ponds was 1 : 1 .4 M:F in T. helveticus and 1 :0.7 in T. vulgaris. In N. viridescens the sex ratio was biased towards males inside the ponds (1:08 M:F), and towards females outside the ponds 1:1.2. During the spring migration it ranged from 1:0.88 to 1:2 M:F ratio; during the fall migration it ranged between 1:1.37 to 1:0 M:F ratio (Hurlbert, 1969). Finally, the sex ratio in Ambystoma texanum (Matthes, 1855) was different in migrating salamanders (1.5 males to 1 female), and in breeding ones (2.2 males to one female) (Petranka, 1984). Annual changes Not only spatial and seasonal variations in sex ratios were described in several urodeles, but sex ratio also appears to vary from year to year and in some cases this variability is rather high. 368 M. R. WARBURG Average sex ratio (M/F) in S. infraimmaculata arranged in 4-year periods 4.5 ü: 3.5 £ 3 | 2.5 8 1.5 R' =0.9527 linear 3.9 1.3 1.55 3.53 2.42 2.41 Uli 1974-77 1978-81 1982-85 1986-89 1991-94 1995-98 4-year periods Fig. 4 Average sex ratio (M:F) in S. infraimmaculata arranged in 4-year periods. Table 4. Sex ratios in other urodeles. Species M:F Source Cryptobranchus allageniensis 1 0.85 Humphries & Pauley, 2005 C. allageniensis 1 0.77 to 1.1.35 Nickerson & Mays, 1972 Amby stoma maculatum 1 0.53 Husting, 1965 A. maculatum 1 1 Blackwell et al, 2004 A. maculatum 1 0.5 Flageole & Leclair, 1992 A. macrodactylum 1 3 Fukumoto & Herrero, 1998 A. opacum 1 1 Parmelee, 1993 (vide Petranka, 1984) A. opacum 1 1 Stenhouse, 1987 A. talpoideum 1 1.06 Hardy & Raymond, 1980 A. talpoideum 1 1 Raymond & Hardy, 1990 A. texanum 1 0.45 Petranka, 1984 A. tigrinum 1 0.76 Rose, 1976 A. annulatum ~ 1 1 Briggler et al. 2004 A. cingulatum 1 0.35 Palis, 1997 Desmognathus fuscus 1 1.27 Orser & Shure, 1975 D. fuscus 1 1 Hall, 1977 D. ochrophaeus 1 0.6 Martof&Rose, 1963 D. ochrophaeus 1 .0.68 Tilley, 1973 D. ochrophaeus 1 1 Hall, 1977 Enastina eschscholtzii platensis 1 0.99 Staub et al, 1995 Plethodon. wehleri 1 0.35 to 1:1.3 Hall & Stafford, 1972 P websteri 1 1 Semlitsch & West, 1983 P. lar selli ~ 1 1 Herrington & Larsen, 1987 P. vehiculum ~ 1 1 Ovaska & Gregory, 1989 P. kentucki 1 0.8 Marvin, 2001 P. cinereus 1 3 Marsh & Goicochea, 2003 Gyrinophilus porphyriticus 1 0.77 to 1:1.19 Bruce. 1978 Eurycea l. longicauda 1 1 Anderson & Martino, 1966 SALAMANDER SEX RATIO 369 E. quadridigitata 1:1 Semlitsch & McMillan, 1980 Stereochilus marginatus 1:1 Bruce, 1971 Triturus vulgaris 1:0.66 Hagström, 1979 T. vulgaris 1:0.58 Bell, 1977 T. vulgaris 1:2.6 Harrison et al., 1983 T. vulgaris 1:2-3.3 Griffiths, 1984 T. vulgaris 1:0.92 Verrell & Francillon, 1986 T. vulgaris ~ 1:1 Arntzen, 2002 T. vulgaris meridionalis 1:1 Kleteäki, 1995 T. cristatus 1:0.61 Hagström, 1979 T. cristatus 1:1.03 Arntzen & Teunis, 1993 T. a. alpestris 1:0.8 Ernst, 1952 T. alpestris 1:1 Kleteäki, 1995 T. alpestris 1:1 Arntzen, 2002 T. helveticus 1:0.93 Gelder, 1973 T. helveticus ~ 1:1 Arntzen, 2002 T. helveticus 1:3 Harrison et al., 1983 T. marmoratus ~ 1:1 Arntzen, 2002 T. m. pygmaeus 1:1 Diaz-Paniagua, 1998 T. vittatus 1:0.91 Geffen et al., 1986/87 T. carnifex 1:1 Kleteäki, 1995 Chioglossa lusitanica 1:0.43 to 1: 1.14 Arntzen, 1981 Euproctus platycephalus 1:1 Bovero v ^ IRAN / e ' '5 i <ü \ -c 1 t£P 'x / / \ V" \ Persian^ — \Gulf \ Saudi Arabia \. 2 V) A La \J Caspian \* • *^ \ A** Iraq -*\ -A, * Turkmenistan IRAN \ -£ ) ,M> \Persian\ — Saudi Arabia A 4 X) /I L ©a NJ Caspian Turkmenistan \ ~Wk Sea U* AV Iraq >v+ ^ + / + IRAN / c S cd ) ^ ^— — — *-._ 1 I v- *i-j. y Y Persian^ — Saudi Arabia A 6 k] /I LINYPHIID SPIDERS OF IRAN 383 Agyneta mollis (O.P.-Cambridge, 1871) Material: IRAN: 1 3 [7598], Eastern Azarbayjan, NW of Sowfyan (38°21'N, 45°51'E), 21.IX.1975. - 1 3 [7504a], S of Khoy (38°28'N, 44°56'E), swamp, under stones and trunks, 1. VI. 1975. Remarks: This species is here reported for the first time for the Iranian fauna. Range: Palaearctic-Alaskan. Agyneta rurestris (C.L. Koch, 1836) Material: IRAN: 1 3 [7328], Tehran, Polur (35°51'N, 52°04'E), 2300 m a.s.l, meadow, 17.VII.1973. - 1 3 [7335], Firuz Kun (35°45'N, 52°46'E), 23.VII.1973. Remarks: This species is here reported for the first time for the Iranian fauna. Range: West Palaearctic. Araeoncus caucasicus Tanasevitch, 1987 Material: IRAN: 5 3, 8 9 [7302], Eastern Azerbaijan, Khoy (38°41'N, 45°08'E), 24.VI. 1973. - 1 3 [7504A], S of Khoy (38°28'N, 44°56'E), swamp, under stones and trunks, 1.VI.1975. - 3 3, 8 ? [7513], Zarjabad (37°38'N, 48°03'E), 8.VI.1975. - 2 o\ 4 9 [7503], Western Azarbayjan, N of Khoy (38°37'N, 45°02'E), l.VI. 1975. - 1 9 [7372], Mazandaran, near Tang-e-Ram (37°25'N, 55°45'E), 28.VII.1974. - 1 3, 7 5 (ZMMU) [7321], Tehran, N of Kendvan Canyon (36°10'N, 51°19'E), 2700 m a.s.l., 12.VII.1973. - 3 9 [7338], Tehran, Avaj (35°38'N, 49°13 , E), 27.VII.1973. - 4 9 [7362], Fasham (35°55'N, 51°32'E), 2000 m a.s.l., 27.VIJI.1973. - 2 9 [7434], Bakhtiyari, Kuhrang (32°28'N, 50°08'E), 19.VI.1974. - 1 3, 3 9 [7438], Esfahan, Nowghan (33°11'N, 50°04'E), 22.VI.1974. - 1 o\ 5 2 [7345], Lorestan, Ma'amulan (33°20'N, 47°54'E), 6.VIII.1973. - 17 9 [7441], Ma'amulan (33°20'N, 47°54'E). 24.VI.1974. - 2 9 [7596], Kordestan, Hoseynabad (35°33'N, 47°08'E), 17.IX.1975. - 7 3, 8 9 [7447], Ham, near Ham (33°37'N, 46°23'E), sifting of litter, 27.VI.1974. - 2 3, 1 9 [7451], Kermanshah, Kangavar (34°29'N, 47°55'E), 1.VII.1974. -2 3,1 9 [7452], Hamadan, NE of Asadabad, (34°51'N, 48°12'E), 2.VII.1974. -23,29 [7579], Fars, S of Fahlyan (30°00'N, 51°35'E), 4.IX.1975. Records from Iran: Western Azarbayjan: Qareh Zia-od-Din. - Tehran: Nesa, Pol-e-Djadjirad. - Khuzestan: Andimeshk, Masjed Soleyman. - Fars: Kavar (Band-e- Bahman), Serizjan, Sivand. - Esfahan: Pol-e-Kaleh. - Ham: Ham (Tanasevitch, 2008). Range: Eastern-Ancient Mediterranean. Figs 1-6 Localities in Iran. (1) (+) Collecting localities visited by A. Senglet, (#) Same, visited by Y. Marusik. (2) (*) Agyneta kopetdaghensis Tanasevitch, 1989, (+) A. mesasiatica Tanasevitch, 2000, (#) Araeoncus mitriformis Tanasevitch, 2008. (3) (+) Archaraeoncns alticola Tanasevitch, 2008, (*) Bolyphantes elburzensis sp. n., (#) Caviphantes dobrogicus (Dumitrescu & Miller, 1962), (A) Diplocephalus transcaucasicus Tanasevitch, 1990, (♦) Erigonoplus sen- gleti Tanasevitch, 2008, (■) E. zagros sp. n. (4) (♦) E. ninae Tanasevitch & Fet, 1986. (*) Megalepthyphantes camelus (Tanasevitch, 1990), (■) M. kronebergi (Tanasevitch, 1989), (#) M. nebulosoides (Wunderlich, 1977), (+) M. nebulosus (Sundevall, 1939), (A) M. pseudocollinus Saaristo, 1997, (O) M. nebulosoides (after Roewer, 1955, as Lepthyphantes nebulosus). (5) (*) Oedothorax meridionalis Tanasevitch, 1987, (#) Pelecopsis laptevi Tanasevitch & Fet. 1986. (+J Piniphantes pinicola (Simon, 1884), (A) Walckenaeria alticeps (Denis, 1952). (6) (#) Sengletus latus sp. n., (+) S. longiscapus Tanasevitch, 2008, (-k) Tenuiphantes perseus (Helsdingen, 1977), (A) Trichoncoides piscator (Simon, 1884). 384 A. V. TANASEVITCH Araeoncus humilis (Blackwall, 1841) Material: IRAN: 1 3 [7503], Western Azarbayjan, N of Khoy (38°37'N, 45°02'E), l.VI. 1975. - 1 3, 3 5 [7526], Kermanshah, N of Kermanshah (34°28'N, 47°00'E), 18.VI.1975. - 1 3, 8 2 [7592], Kordestan, Marivan (35°32'N, 46°09'E), 15.IX.1975. Remarks: This species is here reported for the first time for the Iranian fauna. Range: European-Ancient Mediterranean. Araeoncus mitriformis Tanasevitch, 2008 Figs 2, 7-11 Material: IRAN: 3 3, 3 9 (ZMMU), 18 3, 13 ? [7523], Hamadan, near Hamadan (34°44'N, 48°27'E), 2600 m a.s.l., 16.VI.1975 (type locality). -23 [7434], Bakhtiyari, Kuhrang (32°28'N, 50°08'E), 19.VI.1974. Remarks: The peculiar shape of the carapace in males from the type locality, Hamadan, is more or less uniform (Fig. 7), only a few of them show the distal part of the carapace bent down a little more than in the type. However, both males from Kuhrang differ in having the distal part of the carapace turned upward (Fig. 8), like in Savignia frontata Blackwall, 1833. Most probably, this is an individual variability because the conformation of the male palp in specimens from both localities is identical. This species has been described from a single male. A description of the female is given below for the first time. Description of female: Total length: 2.15. Carapace unmodified, 0.88 long, 0.70 wide, brown. Chelicerae: 0.38 long, unmodified. Legs pale reddish brown. Leg I 2.48 long (0.70+0.25+0.58+0.55+0.40), IV 2.68 long (0.75+0.25+0.70+0.60+0.38). Chaetotaxy: 2.2.1.1. Metatarsi I-III each with a trichobothrium. Tml 0.57. Abdomen 1.25 long, 0.88 wide, grey. Epigyne and vulva as in Figs 9-11. Records from Iran: Hamadan: Hamadan (Tanasevitch, 2008). Range: Iranian. Archaraeoncus alticola Tanasevitch, 2008 Figs 3, 15, 16 Material: IRAN: 1 3, 3 9 [7323 - type locality], Tehran, near Dizan (36°02'N, 51°25'E), 3800 m a.s.l., in snow, 13.VII.1973. Remarks: This species was hitherto known from two males, both from high- montane localities in Tehran: Shemshak (2600 m a.s.l.) and Dizan (3800 m a.s.l.). Now that female material of A. alticola has become available, its description is provided below for the first time. Description of female: Total length: 1.55. Carapace unmodified, 0.60 long, 0.45 wide, pale reddish brown. Chelicerae: 0.25 long, unmodified. Legs pale reddish brown. Leg I 1.39 long (0.40+0.13+0.33+0.28+0.25), IV 1.50 long (0.45+0.15 +0.35+0.30+0.25). Chaetotaxy: 2.2.1.1, tibial spines poorly visible (in male chaetotaxy is unclear or diagnosticated as 1.1.1.1). Metatarsi I-III each with a trichobothrium. Tml 0.47. Abdomen 0.90 long. 0.68 wide, grey. Epigyne as in Fig. 16. LINYPHIID SPIDERS OF IRAN 385 8 9-11 Figs 7-11 Araeoncus mitriformis Tanasevitch, 2008. (7, 8) Varieties of 3 carapace: (7) specimen from Hamadan, Iran, (8) specimen from Kuhrang, Iran. (9, 10) Epigyne, ventral and dorsal view, respectively. (11) Vulva, dorsal view. Taxonomic remarks: The epigynes of A. alticola and A. prospiciens (Thorell, 1875) are very similar to each other and can easily be confused, but the males are well distinguishable by structural details of the palp (see Tanasevitch, 2008). Variability: The shape of the carapace in all known males of A. alticola is almost identical (Fig. 15), in contrast to that of the very similar A. prospiciens (see below under Variability of A. prospiciens; Figs 12-14). Records from Iran: Tehran: Dizan and Shemshak (Tanasevitch, 2008). Range: Iranian. Archaraeoncus prospiciens (Thorell, 1875) Figs 12-14 Material: IRAN: 2 9 [7599], Western Azarbayjan, Qarazia-ed-Din (38°56'N, 45°03'E), 1 .IX. 1975. - 1 3 [7503], N of Khoy (38°37'N, 45°02'E), l.VI. 1975. - 1 o\ 1 9 [7507], Eastern Azarbayjan, Maragheh (37°24'N, 46°16'E), 4.VI.1975. - 1 3, 1 9 [7508], N of 386 A. V. TANASEVITCH Figs 12-16 Archaraeoncus prospiciens (Thorell, 1875) (12-14) and A. alticola Tanasevitch, 2008, paratypes (MHNG) (15, 16). (12) 8 carapace, lateral view, specimen from Dizgaran, Iran. (13, 14) Same, two specimens from Aqa, Iran. (15) 6 carapace, lateral view, specimen from Dizan, Iran. (16) Epigyne, ventral view, specimen from Dizan. Iran. Bonab (37°26'N, 45°57'E), 4.VI.1975. - 1 9 [7511], W of Miyaneh (37°29'N, 47°36'E), 7.VI.1975.-2 9 [7598], NW of Sowfyan(38°21'N, 45°51'E), 21. IX. 1975.-1 9 [7307], Gilan, Hashtpar (37°50'N, 48°58'E), 29.VI.1973; 2 o\ 1 9 (ZMMU) [7365], Rudbar (36°49'N, 49°25'E), 4.IX.1973. - 1 9 [7328], Tehran, Polur (35°51'N, 52°04'E), 2300 m a.s.l, meadow, 17.VII.1973.-2 d*, 2 9 [7363], Aqa (36° 19'N,49°49'E), 2.IX. 1973. - 1 3*, 4 9 [7455], same locality, 6.VII.1974. - 1 3* [7338], Avaj (35°38'N, 49°13'E), 27.VII.1973. - 1 8 [7485], Tarazan/Lowshan (36°28'N, 49°31'E), 8.VIII.1974. - 1 3, 3 9 [7364], Tarazan/Lowshan (36°28'N, 49°31'E), 3.IX.1973. - 4 9 [7536], Shahrak (36°25'N, 50°30'E), 1500 m a.s.l., 2.VII.1975. - 4 9 [7333], Mazandaran, Sari (36°34'N, 53°09'E), 22.Vn.1973. - 1 9 [7372], near Tang-e-Ram (37°25'N, 55°45'E), 28.VII.1974. - 3 o* [7546], near Kiyasar (36°16'N, 53°25'E), 10.VII.1975.- 5 9 [7571], Khorasan, Bodjnour (37°29'N, 57°26'E), 20.VIII. 1975. - 1 o\ 2 9 [7557], E of Badranlu (37°31'N, 57°08'E), 18.VII.1975. - 1 8 [7582], Lorestan, Veseyan (33°29'N, 48°04'E), 8.IX.1975. - 2 o\ 5 9 [7345], Ma'amulan (33°20'N, 47°54'E), 6.VIII.1973.-1 8 [7346],Aligudarz(33 21'N,49 48'E),7.VIIL1973.-l 8,3 9 [7439], Azna (33°28'N, 49°22'E), 23.VI.1974. - 2 3,5 9 [7440], Hatemvand (33°28'N, 48°07'E), 23.VI.1974. - 3 3, 8 9 [7444], Dizgaran (33°43'N, 47°00'E), 25.VI.1974. -23,3 9 [7581], N of Pol-e-Dokhtar (33°12'N, 47°44'E), 7.IX.1975. - 2 3, 6 9 [7582], Veseyan (33°29'N, 48°04'E), 8.IX.1975. - 2 3, 3 9 [7341], Kermanshah, Sahneh (34°28'N, 47°36'E). 2.VIII.1973. - 1 8, 4 9 [7344], Garavand/Shahabad (33°55'N, 46°47'E), 5.VIII.1973. - 2 8, 4 9 [7348], Esfahan, Eskandari (32°50"N, 50°21'E), 8.VIII.1973. -> 3 8, 15 9 [7353], Riz-e-Landjan LINYPHIID SPIDERS OF IRAN 387 (32°24'N, 51°19'E), 1600 m a.s.l., rice fields, 1 1. VIII. 1973. - 5 3, 19 9 [7358], W of Esfahan (32°34'N, 51°31'E), 23.VIII.1973. - 4 9 [7431], Bakhtiyari, E of Farsan (32°17'N, 50°38'E), 17.VI.1974. - 2 3, 2 9 [7432], road from Kuhrang (32°23'N, 50°18'E), 17.VI.1974. - 2 9 [7434], Kuhrang (32°28'N, 50°08'E), 19.VI.1974. - 13 9 [7352], Farsan (32°17'N, 50°31'E), 11.VIII.1973. - 1 9 [7451], Kermanshah, Kangavar (34°29'N, 47°55'E), 1.VII.1974. -2 3, 4 9 [7525], Behistun (= Bisitum or Bisutun) (34°23'N, 47°26'E), 17.VI.1975. - 1 J, 4 9 [7526], N of Kermanshah (34°28'N, 47°00'E), 18.VI.1975. - 1 3,1 9 [7445], Ham, Sarab Eyvan (33°45'N, 46°22'E), 26.VI.1974. - 7 3, 18 -9 [7448], Tchaharmelleh (33°57'N, 46°17'E), 28.VI.1974. - 5 o\ 3 9 [7528], Kordestan, S of Sanandaj (35°13'N, 47°00'E), 21.VI.1975. - 1 3 [7593], E of Marivan (35°32'N, 46°20'E), 16.IX.1975. - 1 3 [7596], Hoseynabad (35°33'N, 47°08'E), 17.IX.1975. - 2 3, 16 9 [7354], Fars, Izadkhast (31°31'N, 52°09'E), 16.VIII.1973. - 1 3, 2 9 [7578], Bishapoor (29°47'N, 51°53'E), 3.IX.1975. Variability: In some samples from different localities (marked with an asterisk in the Material section above), the shape of the male carapace, earlier believed to be uniform, shows a tendency towards the formation of a globular cephalic outgrowth carrying the posterior median eyes, thus making the carapace of A. prospiciens and A. alticola look similar (Fig. 14 cf. Fig. 15). This is a good example of Vavilov's law of homological series (Vavilov, 1922), postulating that closely related taxa are charac- terised by similar hereditary trends. Records from Iran: Western Azarbayjan: Qarazia-ed-Din. - Eastern Azarbayjan: Sowfyan. - Esfahan: Nowghan, Falayarjan, Pol-e-Kaleh. - Lorestan: Dizgaran, Pol-e-Dokhtar. - Khuzestan: Shush. - Kohgiluyeh, Charam, Basht, Yasudj. - Fars: Bishapoor, Dasht-e-Arjan, Serizjan, Allabad, Izadkhast. - Mazandaran: Tang- e-Rah, Shahpasand, Valiabad. - Khorasan: Chaman Bid, Quchan, Emam Qoli, Amirabad, Shandiz Valley, Bojnurd (Tanasevitch, 2008). Range: European- Ancient Mediterranean. Bathyphantes gracilis (Blackwall, 1841) Material: IRAN: 1 9 [7310], Gilan, Parehsar (37°37'N, 49°03'E), sifting debris from tree holes, 2.VII.1973. - 1 9 [7311], Galugah (37°31'N, 49°19'E), 4.VII.1973. - 1 o\ 2 9 [7520], Galugah (37°31'N, 49°19'E), 12.VI.1975. Remarks: This species is here reported for the first time for the Iranian fauna. Range: Holarctic. Bolyphantes elburzensis sp. n. Figs 3, 17-26 Material: IRAN: 3 holotype [7361], Tehran, Shemshak (36°01'N, 51°29'E), 2600 m a.s.l., 27.VIII.1973. - Paratypes: 2 3, 2 9 (ZMMU), 8 3, 11 9, from same locality, together with holotype. - 1 9 [7335], Firuz Kun (35°45'N, 52°46'E), 23.VII.1973. - 1 9 [7544], Mazandaran, W of Razan, (36°12'N, 52°08'E), 1500 m a.s.l, 8.VII.1975. - 4 3, 2 9 [7598], Eastern Azarbayjan, NW of Sowfyan (38°21'N, 45°51'E), 21.IX.1975. Etymology: The specific name, an adjective, refers to the Elburz Mts, the terra typica of the new species. Diagnosis: The species is characterised by the particular shape of the lamella characteristica in the male; a short, drop-shaped pseudoscape, as well as a totally reduced stretcher of the epigyne in the female. Description: Male. Total length 3.00 (2.70-3.15). Carapace 1.25 long, 1.05 wide, yellow to pale brown, with a grey median stripe and a darker margin present in 388 A. V. TANASEVITCH Figs 17-22 Bolyphantes elburzensis sp. n., S paratype from Shemshak, Iran. (17) Right palp, retrolateral view. (18, 19) Paracymbium, different aspects. (20) Lamella characteristica. (21, 22) Embolus, different aspects. darker specimens; unmodified, only head part slightly protruded forward and carrying several short and curved spines. Chelicerae 0.55. long, stridulatory area well deve- loped. Legs yellow to pale brown. Leg I 6.70 long (1.75+0.40+1.70+1.75+1.10), IV 5.80 long (1.60+0.35+1.45+1.50+0.90). Chaetotaxy. Fel: 0-1-0-0, II-IV: 0-0-0-0; Til: 2-1-1-1; II: 2-0-1-1, III: 2-0(l)-0-l, IV: 2-1-1-1; Mtl-IV: 1-0-0-0. Tml 0.25. Metatarsi IV without trichobothrium. Palp (Figs 17-22): Patella with a special seta typical of the genus. Cymbium with a keel-shaped posterodorsal outgrowth. Paracymbium with a wide ridge-like tooth in median part. Lamella characteristica short and wide, with a small branch on dorsal side. Embolus relatively large, slightly S-shaped, with a serrate surface near embolus proper. Abdomen 1.75 long, 1.00 wide, dorsally pale, sometimes white, with a grey median stripe flanked by paramedian spots connected to it with thin bands. Female. Total length 3.35 (2.95-3.40). Carapace 1.25 long, 1.05 wide. Chelicerae 0.50 long. Leg I 6.30 long (1.65+0.45+1.60+1.60+1.00), IV 5.85 long (1.60+0.40+1.45+1.50+0.90). Tml 0.22. Abdomen 2.00 long, 0.65 wide. Epigyne (Figs LINYPHIID SPIDERS OF IRAN 389 Figs 23-26 Bolyphantes elburzensis sp. n., ? paratype from Shemshak, Iran. (23) Epigyne, ventral view. (24) same, dorsal view. (25) Pseudoscape (back wall of epigyne and distal part of scape removed, lateral view). (26) Distal part of scape, anterior view. 23-26): Pseudoscape small, drop-shaped. Distal part of scape well developed, globular, stretcher totally reduced. Posterior median plate large, V-shaped. Body and leg coloration, and chaetotaxy as in male. Variability: A melanic female is present in the above material: its carapace is brown, the dark margin and median stripe are well-expressed, and the legs are with dark bands. Taxonomic remarks: This new species is well distinguished from all congeners by the particular shape of the lamella characteristica in the male, by a short and narrow pseudoscape, as well as a totally reduced stretcher of the epigyne in the female. Distribution: Known from the Elburz Mts, Iran (see Fig. 3). Range: Iranian. Caviphantes dobrogicus (Dumitrescu & Miller, 1962) Fig. 3 Material: IRAN: 1 S [7503], Western Azarbayjan, N of Khoy (38°37'N. 45°02'E), l.VI. 1975. - 1 9 [7556], Mazandaran, Garmab (37°43'N, 56°18'E), 18.VII.1975. - 1 fc IS LJ A. flavicollis (spécimens de référence) LJa. sylvaticus (spécimens de référence) Ha. alpicola (spécimens de référence ■ A. alpicola (spécimens MHNG) 28 28.5 29 29.5 30 30.5 31 31.5 32 32.5 33 33.5 34 34.5 35 DIA/LCB (%) Fig. 2 Rapport de la longueur du diastème (DIA) sur la longueur condylobasale (LCB) de 150 crânes de référence d' Apodemus de 3 espèces et des 143 crânes d'A. alpicola nouvellement identifiés dans les collections du Muséum d'histoire naturelle de Genève (MHNG). Les crânes des 100 individus de référence d'A. sylvaticus et A. flavicollis présentent tous un rapport DIA/LCB inférieur à 31%, tandis que tous les crânes d'A. alpicola (spécimens de référence et spécimens nouvellement identifiés) affichent des valeurs plus élevées, la plupart supérieures à 31,5%, à l'exception de 3 crânes présentant des valeurs de 30,8 ou de 30,9 %. raisons évoquées plus haut. Parmi les Mulots alpestres formellement identifiés, 153 proviennent de 21 localités différentes des Grisons et 17 de 7 localités situées dans les cantons de Berne, Vaud, du Valais et du Tessin. (Tab. 1; Fig. 3). Hormis l'intérêt lié à la découverte de nouvelles localités pour cette espèce, l'examen complet des collec- tions du Muséum a mis en évidence l'intérêt historique de celles-ci, notamment avec la découverte d'un spécimen collecté dans la région de Meiringen (Berne) par Victor Fatio (Tab. 1 ), en 1861, à une époque où ce naturaliste ne reconnaissait l'existence que d'une seule espèce en Suisse (Fatio, 1869). Concernant les données altitudinales, il est possible de rejoindre les conclusions de Reutter et al. (2003) et ainsi affirmer qu'en Suisse le Mulot alpestre s'avère, comme dans le reste de son aire de distribution géographique, une espèce limitée aux étages montagnard, subalpin et alpin inférieur, avec une altitude maximum d'au moins 2200 m en ce qui concerne deux spécimens en provenance du Val S-Charl (Grisons). Par ailleurs, il est également possible de confirmer que le Mulot alpestre et le Mulot sylvestre peuvent vivre en sympatrie. En effet, sur les 28 localités notées avec une présence d'A. alpicola, 12 accueillent également la présence d'A. sylvaticus. En revanche, aucun Mulot à collier en provenance des Grisons n'a été trouvé parmi les 258 Apodemus collectés dans ce canton. Toutefois, considérant que cet échantillonnage historique n'est guère exhaustif en matière altitudinale, il n'est pas possible de déter- miner si cette espèce est réellement rare dans les zones les plus basses de cette partie des Alpes où l'essentiel des spécimens collectés provient du Parc national et des APODEMUS ALP1C0LA 433 r- C OÜÜOS 1> OJ 13 cd CQ CQ os cd CQ CQ -o -a -a -a .2 .2 .2 .2 S 'S 'S 'S 'S '> -r- ecce cd cö cd cd OO o OÜ cy cy w -a 13 -a °6 =%j •a -a oooÇocSS w cu cu cu cu eu td < >,;>>>,>ì>i;>%>,>, >»-c >,>,"C UUUUUUIUUUK UD -a M -S Ctu c > > > >,n3 rc >i >> g g >v^î >-> QqJ g S S <<<<<<<<<0< : sou a >><* Isa cudü O S 1 en te CN C) ^-'t'*ir 1 vDMXio\ooooo-'rNtNtsm^|-invoi^ooo\0"(Nm'*ininiOt^cjo 434 J. GILLIERON APODEMUS ALPICOLA 435 régions voisines, soit à des altitudes généralement supérieures à 1200 m. Enfin, soulignons que ces nouvelles données ne permettent pas encore d'affiner la distribution du Mulot alpestre dans les Alpes valaisannes, de même que dans les Alpes tessinoises où l'espèce n'a été trouvée qu'à Airolo (Tab. 1 ; Fig. 3). Aussi, afin de savoir si l'espèce est réellement moins répandue dans ces régions des Alpes, des recherches ciblées y devraient être entreprises, soit par des campagnes de capture, soit par des recherches complémentaires dans les différentes collections museales de Suisse en utilisant la méthode simplifiée proposée ici. REMERCIEMENTS Pour ses encouragements, son aide ponctuelle et ses remarques avisées, mes remerciements s'adressent à M. Ruedi (Genève); ils s'adressent également à P. Vogel (Lausanne) qui m'a mis à disposition la collection IZEA et fait part de ses précieux commentaires relatifs au Mulot alpestre. Ma reconnaissance va aussi à B. Pavillard pour son assitance technique dans les collections de cet institut et à Brigitte Reutter (Berne) qui m'a aimablement fourni des compléments d'information relatifs au matériel examiné dans le cadre de sa thèse. RÉFÉRENCES Aeschimann, J., Bourquin, L., Engels, B., Thomas, C. & Vogel, P. 1998. Comparative winter thermoregulation and body temperature in three sympatric Apodemus species (A. alpi- cola, A. flavicollis and A. sylvaticus). Zeitschrift für Säugertierkunde 63: 273-284. Debernardi, P., Patriarca, E. & Reutter, B. A. 2003. Contribution to the knowledge of Apodemus genus in the Gran Paradiso National Park. Hystrix - Italian Journal of Mammalogy 14 (1-2): 55-75. Fatio, V 1869. Faune des vertébrés de la Suisse. Vol. I. Histoire naturelle des mammifères. Georg, Genève, 410 pp. Heinrich, G. 1951. Die deutschen Waldmäuse. Zoologische Jahrbücher. Abteilung für Syste- matik, Ökologie, Geographie und Biologie der Tiere 80: 92-122. Heinrich, G. 1952. Apodemus flavicollis alpicola. Journal of Mammalogy 33: 260. Reinwaldt, E. 1955. Zur Kenntnis der Gelbhalsmaus, Apodemus flavicollis alpicola Heinrich, 1951. Säugetierkundliche Mitteilungen 3 : 151-1 54. Reutter, B. A. 2002. The Alpine mouse Apodemus alpicola, Heinrich 1952: discrimination and distribution in comparison with sympatric congeneric species. Thèse, Lausanne, 162 pp. Reutter, B. A., Bertouille, E. & Vogel, P. 2005. The diet of the Alpine mouse Apodemus alpicola in the Swiss Alps. Mammalian Biology 70(3): 147-155. Reutter, B. A., Brünner, H. & Vogel, P. 2001. Biochemical identification of three sympatric Apodemus species by protein electrophoresis of blood samples. Mammalian Biologx 66(2): 84-89. Reutter, B. A., Hausser, J. & Vogel, P. 1999. Discriminant analysis of skull morphometric characters in Apodemus sylvaticus, A. flavicollis and A. alpicola (Mammalia; Rodentia) from the Alps. Acta Theriologica 44(3): 299-308. Reutter, B. A., Helfer, V, Hirzel, A. H. & Vogel, P. 2003. Modelling habitat-suitability using museum collections: an example with three sympatric Apodemus species from the Alps. Journal of Biogeography, 30, 581-590. Fig. 3. Localisation géographique des spécimens <¥ Apodemus alpicola présents dans les collections du Muséum d'histoire naturelle de Genève (ronds noirs) ou issus de la littérature (Vogel, Maddalena & Mabille, 1991; Reutter et al, 2002; triangles noirs). Les numéros se réfèrent aux localités citées dans le tableau 1 . 436 J - GILLIERON Reutter, B. A., Petit, E. & Vogel, P. 2002. Molecular identification of an endemic Alpine mammal, Apodemus alpicola, using a PCR-based RFLP method. Revue suisse de Zoologie 109(1): 9-16. Spitzenberger, F. 2001. Die Säugetierfauna Österreichs. Bundesministerium für Land- und Forstwirtschaft Umwelt und Wasser-wirtschafi, Graz. 895 pp. Spitzenberger, F. & Englisch, H. 1996. Die Alpen waidmaus (Apodemus alpicola Heinrich, 1952) in Österreich. Mammalia austriaca 21. Bonner zoologische Beiträge 46: 249-260. Storch, G. & Lütt, O. 1989. Artstatus der Alpenwaldmaus, Apodemus alpicola Heinrich, 1952. Zeitschrift für Säugetierkunde 54: 337-346. Vogel, P. 1995. Le Mulot alpestre - Apodemus alpicola (pp. 279-281). In: Hausser, J. (ed). Mammifères de la Suisse. Birkhäuser, Basel, XII + 501 pp. Vogel, P., Maddalena, T., Mabille, A. & Paquet, G. 1991. Confirmation biochimique du statut spécifique du mulot alpestre Apodemus alpicola Heinrich, 1952 (Mammalia, Rodentia). Bulletin de la Société vaudoise des Sciences naturelles 80 (4): 471-481. Revue suisse de Zoologie 1 16 (3-4): 437-439; décembre 2009 Magapia, nom. nov., replacing Laingia Bouillon, 1978, and Magapiidae, nom. nov., replacing Laingiidae Bouillon, 1978 [Cnidaria, Hydrozoa] Peter SCHUCHERT and f Jean BOUILLON Muséum d'histoire naturelle, CP 6434, CH-121 1 Genève 6, Switzerland. E-mail: Peter.Schuchert@ville-ge.ch Magapia, nom. nov., replacing Laingia Bouillon, 1978, and Magapiidae, nom. nov., replacing Laingiidae Bouillon, 1978 [Cnidaria, Hydrozoa]. - The genus name Laingia Bouillon 1978 is an invalid junior homonym of Laingia Theobald, 1922 (Insecta). Magapia, nom. nov., is proposed as a new replacement name for Laingia Bouillon, 1978b, and the invalid family- group name Laingiidae Bouillon, 1978b based on it is replaced by Maga- piidae, nom. nov. Keywords: Hydrozoa - Laingia - homonymy - new replacement name - Magapia. Bouillon (1978b) introduced the genus name Laingia for a remarkable hydro- medusa found near Laing Island, Papua New Guinea. The classification of this medusa (Laingia jaumotti Bouillon, 1978b) poses considerable problems in presenting a mixture of characters found in two otherwise rather distant groups of Hydrozoa, the Narcomedusae and Anthomedusae. Bouillon (1978b) thus assigned the genus to a family and a subclass of its own, Laingiidae and Laingiomedusae. Kantiella enigmatica Bouillon, 1978a, originally included in the hydrozoan family Proboscidactylidae, shares characters with Laingia jaumotti and was also assigned to Laingiidae by Bouillon (1978b). Later, Fabienna Schuchert, 1996 was also included in Laingiidae (Bouillon & Barnett, 1999). When naming Laingia, Bouillon was unaware that the name was already in use for a genus of insects {Laingia Theobald, 1922; Laingia psammae Theobald, 1922; family Aphidae). Laingia Bouillon, 1978b is thus an invalid junior homonym and must be replaced [International Code of Zoological Nomenclature, ICZN Art. 60]. No available and potentially valid synonyms for the name are known [ICZN Art. 60.3], and a new substitute name is proposed here. The genus name Laingia is also used for some marine red algae {Laingia Kylin, 1929; Rhodophyta). Jean Bouillon became aware of the homonymy discussed here in 2008, and despite his deteriorating health he planned to publish new replacement names for the genus and family he had established. In early 2009, he discussed the plans and names he had in mind with his friend Ferdinando Boero in Lecce, Italy. Prof. Bouillon un- fortunately passed away on 29 March 2009 without finalizing a manuscript draft and Prof. Boero thus asked the first author to publish the new name according to the wishes of J. Bouillon. Manuscript accepted 17.06.2009 438 P- SCHUCHERT AND J J. BOUILLON A new replacement name, Magapia, nom. nov., is proposed herewith for the preoccupied genus name Laingia Bouillon, 1978b. The type species of the genus remains Laingia jaumotti Bouillon, 1978b [ICZN, Art. 67.8], now Magapia jaumotti, comb. nov. (Bouillon, 1978b). The family-group name Laingiidae Bouillon, 1978b is also invalid and must be replaced [ICZN Art. 39]. Magapiidae, nom. nov., is proposed as a new replacement name for this hydrozoan family. The subclass name Laingiomedusae Bouillon, 1978b need not be replaced. The name Magapia was chosen in memory of Miller Magap, manager of the biological station Leopold III on Laing Island (see Bouillon et al., 1987). Prof. Bouillon spent many months at the station and discovered many remarkable new species there, including Magapia jaumotti (see Bouillon, 1978b-c, 1980-1984; Bouillon et al, 1988-1986). Miller Magap, a Papua New Guinean, offered invaluable assistance to scientists visiting the station, and he was much appreciated by all. Aged 49, he was brutally mur- dered on 24 March 1997. While travelling back to the station on a truck, he and seve- ral scientists were ambushed by local bandits intent on robbery. A stone thrown by the attackers - meant to stop the vehicle by breaking the window - hit his head, and he died within minutes. His untimely death while at work deeply touched Prof. Bouillon and other scientists. ACKNOWLEDGEMENTS I wish to thank Prof. N. Boero for bringing the homonymy problem to my attention and for his confidence by entrusting to me the task to fulfil Jean Bouillon's plans. My sincere thanks go also to Prof. Dale Calder who read and commented a first draft of this publication. As with so many other publications, his corrections consi- derably improved the style and language of this contribution. REFERENCES Bouillon, J. 1978a. Hydroméduses de l'archipel des Sechelles et du Moçambique. Revue de Zoologie Africaine 92: 117-172. Bouillon, J., 1978b. Hydroméduses de la mer de Bismarck (Papouasie, NouvelleGuinée. Partie II. Limnomedusa. Narcomedusa, Trachymedusa et Laingiomedusa (sous-classe nov.). Cahiers de Biologie Marine 19 : 473-483. Bouillon, J. 1978c. Hydroméduses de la mer de Bismarck (Papouasie, Nouvelle-Guinée). Partie 1: Anthomedusae Capitata (Hydrozoa - Cnidaria). Cahiers de Biologie Marine 19: 249-297. Bouillon, J. 1980. Hydromeduses de la Mer de Bismarck. (Papouasie Nouvelle-Guinée). Partie 3: Anthomedusae - Filifera (Hydrozoa - Cnidaria). Cahiers de Biologie Marine 21: 307-344. Bouillon, J. 1984. Hydroméduses de la mer de Bismarck (Papouasie, Nouvelle-Guinée). Partie IV: Leptomedusae (Hydrozoa-Cnidaria). Indo Malayan Zoology 1: 25-1 12. Bouillon, J.. Boero, F., & Seghers. G. 1988. Notes additionelles sur les hydroméduses de la mer de Bismarck (Hydrozoa-Cnidaria) IL Indo Malayan Zoology 5: 87-99. Bouillon, L, Boero, F., & Seghers, G. 1991. Notes additionnelles sur les méduses de Papouasie Nouvelle-Guinée (Hydrozoa. Cnidaria) 4. Cahiers de Biologie Marine 32: 387-411. NEW REPLACEMENT NAME 439 Bouillon, J., Seghers, G., & Boero, F. 1988. Notes additionnelles sur les méduses de Papouasie Nouvelle-Guinée (Hydrozoa, Cnidaria). 3. Indo-Malayan Zoology 5: 225-253. Bouillon, J., Barnett T. J. 1999. The Marine Fauna of New Zealand: Hydromedusae (Cnidaria: Hydrozoa). Niwa Biodiversity. Memoir, New Zealand 113: 1-136. Bouillon, J., Claereboudt, M., & Seghers, G. 1986. Hydroméduses de la baie de Hansa (Mer de Bismarck; Papouasie Nouvelle-Guinée). Répartition, condition climatiques et hydro- logiques. Indo Malayan Zoology 3: 105-152. Kylin, H. 1929. Die Delesseriaceen Neuseelands. Acta Universitatis Lundensis. Nova Series. Lunds Universitets Arsskrift. Ny Foljd. Andra Afdelningen. Medicin samt Matematiska ochNaturvetenskapliga Amnen 25: 1-15. Schuchert, P. 1996. The marine fauna of New Zealand: athecate hydroids and their medusae (Cnidaria: Hydrozoa). New Zealand Océanographie Institute Memoir 106: 1-159. Theobald, F. V. 1922. A new aphid genus and species found in England. Bulletin of Entomo- logical Reseach - London 12: 429-430. Revue suisse de Zoologie 1 16 (3-4): 441-507; décembre 2009 The European athecate hydroids and their medusae (Hydrozoa, Cnidaria): Filifera Part 5 Peter SCHUCHERT Muséum d'histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. E-mail: Peter.Schuchert@ville-ge.ch The European athecate hydroids and their medusae (Hydrozoa, Cnidaria): Filifera Part 5. - This study reviews all European hydroids belonging to the filiferan families Bythotiaridae, Proboscidactylidae, Magapiidae, Ptilocodiidae, Eucodoniidae, Russelliidae, Niobiidae, Pro- tiaridae, and Trichydridae. Protiara tetranema (Péron & Lesueur, 1810) is considered as an unrecognizable species. The gonozooids of Halitiara formosa and Trichydra pudica are described, both have reduced blastostyles in a gonotheca-like casing. Keywords: Cnidaria - marine - Hydrozoa - revision - taxonomy - north- eastern Atlantic - Mediterranean. INTRODUCTION This study is the sixth in a series of taxonomic revisions and reviews of the European athecate hydroids and their medusae (Order Anthoathecata = Anthomedusae = Athecata). The previous ones are: Schuchert (2004; Oceaniidae and Pachy- cordylidae), Schuchert (2006; Acaulidae, Boreohydridae, Candelabridae, Cladocory- nidae, Cladonematidae, Margelopsidae, Pennariidae, Protohydridae, Tricyclusidae), Schuchert (2007; Bougainvilliidae, Cytaeididae, Rathkeidae, and Pandeidae), Schuchert (2008a; Hydractiniidae, Rhysiidae, Stylasteridae), Schuchert (2008b: Eudendriidae). This publication concludes the Filifera with the families Bythotiaridae, Proboscidactylidae, Magapiidae, Ptilocodiidae, Eucodoniidae, Russelliidae, Niobiidae, Protiaridae, and Trichydridae. MATERIAL AND METHODS See also Schuchert (2008a), for a general introduction to the Hydrozoa see Bouillon et al. (2004,2006). Where possible, it was attempted to supplement the species descriptions by sequence information from part of the 16S mitochondrial rRNA gene. The methods to obtain DNA sequences are described in Schuchert (2005, 2007). All sequences have been submitted to the EMBL database (accession numbers FN422378-FN422379). DNA of some specimens was also given to other researches who determined 16S or other gene sequences. The origin and identity of the material used to obtain the 16S sequence data as well as the accession numbers are given for each species in the section "Material examined". Manuscript accepted 30.07.2009 442 P. SCHUCHERT Abbreviations BMNH The Natural History Museum, London, England ERMS European Register of Marine Species (Costello et al., 2001) MHNG Muséum d'histoire naturelle de Genève, Switzerland ICZN International Code of Zoological Nomenclature IRSN Institut Royal des Sciences Naturelles de Belgique, Bruxelles ZMUC Zoological Museum Copenhagen, Denmark ZMA Zoological Museum Amsterdam, The Netherlands ZMB Zoological Museum Berlin, Germany TAXONOMIC PART Family Bythotiaridae Maas, 1905b Synonyms: Calycopsidae Mayer, 1910: 104. - Bythotiaridi Mayer, 1910: 183. Diagnosis: Medusa without apical projection, marginal bulbs very indistinct or absent; four, eight or more hollow marginal tentacles, each terminating in large swelling covered by nematocysts, basal portion of tentacles usually adnate to exum- brella; with or without rudimentary or dwarf solid tentacles; four or eight radial canals, simple or branching, with or without centripetal canals growing from circular canal towards centre; without gastric peduncle, mouth with four simple or crenulated lips; gonads on manubrial wall, simple or horizontally folded, adradial or interradial. Rarely abaxial ocelli on tentacle bases. Hydroid stage only known for Bythotiara, living in prebranchial cavities of tunicates, colonial or solitary, monomorphic; without perisarc; hydrorhiza root- or plate-like, hydranths unbranched, hydranth composed of a body and a sometimes indistinctly demarcated naked pedicel; with up to five irregular whorls of filiform ten- tacles concentrated in a narrow band below hypostome; medusa buds arising from polyps at junction of pedicel and hydranth body. Cnidome includes microbasic euryteles and large desmonemes. Remarks: See Bouillon et al. (2006) for more details and a key to all genera. The subdivision of the family in genera is not well resolved and progress is hampered by the fact that many polyp stages of this family remain unknown. The genera some- times intergrade and it is sometimes difficult to draw a clear line separating them. Key to the genera of Bythotiaridae medusae of the ERMS zone: la centripetal radial canals present, blindly ending or joining base of stomach Calycopsis lb without centripetal radial canals 2 2a radial canals not bifurcated or more usually bifurcated once (some few additional branches may occur as abnormalities) Bythotiara 2b radial canals branching repeatedly at various levels Sibogita Genus Bythotiara Guenther, 1903 Synonyms: Ascidioclava Kirk, 1915, type species Ascidioclava parasitica Kirk, 1915. - Crypta Fraser, 1911 [name preoccupied]. - Endocrypta Fraser, 1912, type species Crypta hunts- mani Fraser, 1911. Type species: Bythotiara murrayi Guenther, 1903 by monotypy. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 443 Diagnosis: Bythotiaridae medusae with four simple or bifurcated radial canals, without centripetal canals; with or without secondary (rudimentary) tentacles, these entirely covered with nematocysts; gonads interradial, with transverse furrows; no ocelli. Polyp stage as given in family diagnosis. Remarks: Pages et al. (1992) provided a table with the characteristics of all known Bythotiara medusae. Recently, Raskoff & Robison (2005) described a new Bythotiara polyp from pelagic tunicates, making it necessary to modify slightly the diagnosis of the polyp stage. Only one Bythotiara species occurs in the ERMS zone. Bythotiara murrayi Guenther, 1903 Fig. 1 Bythotiara murrayi Guenther, 1903: 424, pi. 10 figs 4-5. - Mayer, 1910: 185, figs 97-98. - Vanhöffen, 1911: 213, figs 9a-c. - Kramp & Damas, 1925: 281. - Hartlaub, 1914: 355, figs 304-306. - Kramp, 1924: 12, figs 8-12. - Kramp, 1926: 97, figs 38-40. - Ranson, 1936: 98, pi. 1 fig. 12. -Russell, 1940: 515, figs 8-10. - Russell, 1953: 215, figs 113a- b, 114a-b, 115-116, pi. 13 fig. 1.- Kramp, 1959: 125, figs 1, 132. - Kramp, 1961: 118. - Kramp, 1968: 54, fig. 142. - Van der Spoel & Bleeker, 1988: 167, fig. 17. - Pages et al. 1992: 7, fig. 7. - Schuchert, 1996: 22, fig. 8. Type material: Not located (not found in BMNH). Material examined: BMNH 1985.10.21.1-4; North Atlantic; depth 821 m; 3 medusae collected 20.07.1955 to 20.06.1957. Diagnosis: Medusa about 20 mm high and wide, thick walls; stomach small; four interradial gonads with transverse furrows, no vertical subdivision. Four primary radial canals each bifurcating once (but additional branching may occur); eight or more tentacles (as many tentacles as radial canals); with small secondary and minute tertiary tentacles. Description (after Russell (1953) and own observations): Umbrella hemi- spherical or somewhat bell-shaped, about as high as or slightly higher than wide, with rounded apex, jelly thick, no gastric peduncle. Stomach small, somewhat rhomboid in shape; mouth with four short simple lips. Four gonads, one covering each interradial wall of stomach, with few transverse folds subdividing them into several masses, top mass triangular and with central depression (Fig. IB). Four primary radial canals, each bifurcating near point of origin from stomach to form eight straight, narrow, smooth canals joining narrow ring canal; each primary canal leaves stomach as short narrow funnel (= mesentery); additional branchings and irregularities of radiais canals frequent (Fig. 1C-F). No ocelli. Eight or more primary marginal tentacles (as many tentacles as radial canals), large, smooth, hollow, without bulbs or basal swellings, tentacles at base flattened oral- aborally and adnate to umbrella margin, each tentacle terminating in large hemi- spherical nematocyst cluster composed of desmonemes; one to five small hollow secondary marginal tentacles between every two primary marginal tentacles, secondary tentacles contain ?euryteles and desmonemes, otherwise similar to primary tentacles; additional tertiary dwarf tentacles may be present. Nematocysts: ? microbasic euryteles and desmonemes, discharged desmonemes with five coils. 444 P. SCHUCHERT Fig. 1. Bythotiara murrayi Guenther, 1903. (A) Mature medusa, maximal diameter about 25 mm, modified after Hartlaub (1914). (B) Manubrium with gonads, interradial view, modified after Russell (1953). (C-F) Variation of the branching pattern of the radial canals, from Kramp (1924), C-D in aboral view, E-F in lateral view, numbers and letters identify the radial canals and their branches. (G) Nematocysts: desmoneme and ? microbasic eurytele, for dimensions see text, from Russell (1940). EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 445 Colour of umbrella faint violet; gonad-covered part of stomach brick red; remainder of medusa colourless. Polyp unknown. Dimensions (Russell, 1940; 1953): Mature medusae 5-21 mm in height. ? Euryteles (17-20)x(10-ll) i um; desmonemes (13-14)x(6)/ym. Distribution: Eastern Atlantic from Norway to South Africa; Mediterranean; north-western Atlantic; tropical parts of the Indian Ocean; Indonesia; Papua New Guinea, New Zealand (Hartlaub, 1914; Kramp & Damas, 1925; Russell, 1953; Kramp, 1968; Bouillon, 1980; Winkler, 1982; Van der Spoel & Bleeker, 1988; Pages et al, 1992; Schuchert, 1996). Type locality: 52.30167°N 15.89833°W (SW of Ireland). Biology: Bythothiara murrayi is an oceanic medusa that is only rarely encoun- tered in inshore waters. It can be caught at depths of several hundred meters only. Remarks: The smallest medusa stages found by Russell (1953) measured 5 mm, but they already had mature gonads. Genus Calycopsis Fewkes, 1882b Type species: Calycopsis typa Fewkes, ] 882b. Diagnosis: Bythotiaridae medusae with four initially unbranched radial canals and with four or more centripetal canals arising from the ring canal, blind or joining the cruciform base of stomach or the perradial canals. Gonads interadial, usually with eight adradial rows of deep transverse furrows, alternatively transversely folded, or smooth, or with pits; basal portion of tentacles adnate to umbrella margin; tentacles hollow, usually tentacles all alike, rarely two types may be present, nematocysts only in terminal knobs. Without ocelli. Remarks: The genus Calycopsis and its species are discussed in Kramp (1959) and Lindsay et al. (2008). The polyps remain unknown. Below follows an identifica- tion key to the Calycopsis medusae of the Atlantic north of the Equator (adapted from Kramp, 1959). A species not yet known from the ERMS zone has been included in brackets. It is not unlikely that it will ultimately also be found in the ERMS zone. Key to the Calycopsis species of ghe ERMS zone: la umbrella with funnel-shaped apical depression; 3-4 centripetal canals in each quadrant C. typa lb umbrella without an apical depression 2 2a with only 4 perradial tentacles; one interradial centripetal canal; tentacle-base with adaxial appendage C. krampi 2b with 8 or more tentacles, no adaxial appendages on tentacle bases 3 3a umbrella margin with exumbrellar papillae; 2 centripetal canals in each quadrant; 8-12 tentacles [C. papillata] 3b marginal lobes of umbrella without papillae 4 4a gonads smooth or with few, irregular transverse folds; one centripetal canal in each quadrant; 8 tentacles; all alike C. simplex 446 P- SCHUCHERT 4b gonads with 8 adradial rows of deep, transverse furrows; more than 8 tentacles 5 5a 7 or more centripetal canals in each quadrant, most of them joining base of stomach or upper part of neighbouring canals; 16-32 tentacles . . . . C. chuni 5b 1 or 2 centripetal canals in each quadrant 6 6a one centripetal canal in each quadrant, blind; 8-16 long and several small tentacles C. bigelowi 6b 1-2 centripetal canals in each quadrant, joining base of stomach; about 40 long tentacles C. gara Calycopsis typa Fewkes, 1882 Fig. 2 Calycopsis typa Fewkes, 1882b: 304, pi. 1 fig. 34. - Mayer, 1910: 131, fig. 70. - Hartlaub, 1914: 359: figs 293-294, 307. - Kramp, 1959: 21, 127, fig. 139, revision. - Kramp, 1961: 121, synonymy. -Van Soest, 1973: 121, fig. 2a. Sibogita nuarchus Bigelow, 1909b: 206, pi. 1 figs 1-8. - Bigelow, 1913: 21, synonym of C. typa. Not Calycopsis typa. - Vanhöffen, 1911: 214, pi. 22 fig. 6. [= C. chimi Vanhöffen, 1911] Not Calycopsis typa. -Vanhöffen, 1912: 364 [?= C. papillata] Sibogita typa. - Mayer, 1910: 491. ? Calycopsis typa. - Pages et al, 1992: 7, fig. 8A-C. Type material: United States National Museum, catalogue number 9727 (not seen). Diagnosis: Calycopsis medusa with a characteristic funnel-shaped depression at apex, 3-4 blind centripetal canals in each quadrant, 16 long tentacles. Description: Medusa umbrella spherical to barrel-shaped, jelly thick, solid, with a characteristic funnel-shaped depression at apex, umbrella somewhat laterally flattened, plane in which flattening occurs is either radial or interradial. Umbrella margin lobed through embayments for the tentacles. Manubrium large and voluminous, length 1/2 to 2/3 of bell cavity, urn-shaped, about two-thirds as broad as long, aboral end cruciform in section, mouth with four short perradial lips, mouth rim smooth or crenulated. Gonads interradial, discontinuous in the perradii, with eight adradial rows of about 20 horizontal furrows, irregular, projecting lobes of gonads may be developed. Four radial canals, widening near manubrium and forming short mesenteries, 3- 4 blind centripetal canals in each quadrant, interradial centripetal canal the furthest de- veloped, reaching nearly to the base of the manubrium, maximally usually around 16 canals in total, maximum number of canals observed 21. Connected with each canal is a well-developed tentacle, thus around 16 in total, between long tentacles usually occur very small tentacles (16), long tentacles adnate to umbrella margin in furrows at the rim, long tentacles with club-like swollen ends, concentration of nematocysts at tip. Colours: gonads deep brownish-red; terminal knobs pale yellowish. Dimensions: Up to 37 mm high and 40 mm wide. Bigelow (1909b, as Sigogita nuarchus) gives a table with measurements, tentacle- and canal numbers. Biology: A deep-water species, usually occurring at depths of several hundred metres (Kramp, 1959). EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 447 Fig. 2 Calycopsis typa Fewkes, 1882; modified after Bigelow (1909b) and Hartlaub (1914), note the characteristic apical funnel (arrow). Distribution: Atlantic Ocean, off the north-eastern coast of the USA, Cape Verde Islands (Kramp, 1959), Azores (Van Soest, 1973), ? South Africa (Pages et al., 1992). Type locality: Atlantic, off the New England, coast USA. Remarks: This species is rare in the ERMS area, it is only known from one specimen caught in deep waters NE of the Azores. The South African medusae described by Pages et al. (1992) matched the description given above, but additionally they had knobs of nematocysts on the adaxial sides of the tentacle bases. Such knobs have never been mentioned for C. typa by other authors and perhaps the South African medusae belong to a new, unnamed species. Calycopsis krampi Petersen, 1957 Kramp, 1959: 126, fig. 135. Fig. 3 Kramp, 1961: Cahcopsis krampi Petersen, 1957: 31, figs. 2-3. 120. -Fraser, 1974: 13. Material examined: ZMUC; holotype, without registration number; Dana Station 9806; 50.917°N 14.000°W; net tow with 120-200 m wire out; 21 August 1955. Diagnosis: Calycopsis medusa up to 4 mm high, 4 tentacles, 4 centripetal ca- nals, at base of each tentacle an adaxial appendage pointing into bell cavity. Description: Medusa spherical, jelly thick. Manubrium large and voluminous, length 3/4 of bell cavity. Four large interradial gonads leaving only mouth-region and perradial parts of manubrium visible; each gonad with very deep, vertical interradial furrow and three to four irregular transverse folds. The perradial surfaces of the sto- mach form four prominent longitudinal ridges, continued as short mesenteries where they join the radial canals. 448 P. SCHUCHERT Fig. 3 Calycopsis krampi Petersen, 1957; modified after Petersen (1957), size 4 mm, the arrow points at one of the characteristic adaxial projections. Four complete radial canals and four narrower, blind centripetal canals reaching almost to the top of the bell. Four perradial tentacles without tentacular bulbs, proximal parts adnate to bell margin, structure of distal ends unknown. Four prominent adaxial appendages at the places where the radial canals join the ring canal, one in each perradius, projecting obliquely into bell cavity (arrow in Fig. 3). Each projection forms a direct adaxial continuation of the basal part of the tentacle attached to the exumbrella, the projection is about half the diameter of the bell opening long, nearly half as broad as long, tapering a little distally, drawn into two 'papillae' at the tip, aboral side with a longitu- dinal ridge. Dimensions: Umbrella 4 mm high and 3 mm wide. Biology: Likely restricted to depths below 100 m, Fraser (1974) recorded it from deep waters east of Rockall. Distribution: A very rare species occurring west of the British Isles (Petersen, 1957; Fraser, 1974). Type locality: Porcupine Seabight SW of Ireland, 50.917°N 14.00°W, max. 200 m depth. Remarks: This species is known from only three specimens. Its perradial pro- jections are unique and render the species quite distinct. The transversal folds of the gonads seen in the type specimen could be a fixation artifact. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 449 Calycopsis simplex Kramp & Damas, 1925 Fig. 4 Calycopsìs simplex Kramp & Damas, 1925: 282, figs 23-25. - Kramp, 1959: 126, fig. 133. - Kramp, 1961: 121. - Goy, 1973: 986, fig. 8. - Gili et al, 1998: 1 16, fig. 2. Type material: Bergen Museum, Norway (not seen). Material examined: MNHN1647; Mediterranean, France, Bay of Villefranche-sur- Mer; 16 December 1966; one medusa, material of Goy (1973). Diagnosis: Calycopsis medusa up to 9 mm, hemispherical; gonads smooth or with few, irregular transverse folds; 4 blind interradial centripetal canals; 8 tentacles all alike. Description: Medusa globular to hemi-spherical, jelly moderately thick, umbrella margin with embayments for tentacles. Manubrium conical, length 1/2 of bell cavity, cruciform cross-section, mouth small, coiners not much drawn out. Gonads cover manubrium almost entirely as eight large adradial pads, adnate interradially but separated by deep vertical furrows, separated perradially, surface either smooth or with few, irregular transverse folds. Four complete radial canals and four blind centripetal canals reaching to middle or top of the bell. Eight tentacles, four perradial, four interradial, all alike, without tentacular bulbs, proximal parts adnate to bell margin, distal ends swollen, elongated club-like. No ocelli. Dimensions: Umbrella diameter 8-9 mm. Distribution: Norway (Kramp & Damas, 1925), Mediterranean (Goy, 1973; Gili el al., 1998). Type locality: Hj0rundfjord, Norway, deeper than 400 m. Biology: Likely a deep water species, two of the three records were from more than 400 m depth. Calycopsis chuni Vanhöffen, 1911 Fig. 5 Calycopsis chuni Vanhöffen, 1911:217, pi. 22 fig. 8. - Bigelow, 1940: 290. - Kramp, 1959: 23, 127, fig. 140, revision. - Kramp, 1961: 119. - Kramp, 1968: 55, fig. 148. Calycopsis valdiviae Hartlaub, 1914: 360. - Kramp, 1959: 23, synonym. Calycopsis typa. -Vanhöffen, 1911: 214, pi. 22 fig. 6. [not Calycopsis typa Fewkes, 1882] Type material: Naturkundemuseum Berlin, Germany, syntypes ZMB Cni 14845 and ZMB Cni 14838 (not seen). Diagnosis: Calycopsis medusa around 30 mm high and wide, jelly thick, 32 or more radial canals, 16 long tentacles and up to 16 short ones; gonad in each quadrant with two vertical rows of 19-32 transverse clefts. Description (Vanhöffen, 191 1 ; Kramp, 1959): Umbrella spherical to somewhat oblong, jelly thick and rigid. Manubrium about half as long as bell cavity, mouth rim crenulated, perradial corners of mouth drawn out into four simple lips, manubrium-base cruciform. Gonad covering most of manubrium, with eight adradial rows of 19-32 parallel transverse furrows. Four perradial canals and usually 28 centripetal canals, occasionally up to 60, all communicate with stomach in fully grown specimens, usually directly joining the 450 P. SCHUCHERT Fig. 4 Calycopsis simplex Kramp & Damas, 1925; after preserved material from the Mediterranean, scale bar 2 mm. stomach and thus like radial canals, but sometimes fused with a neighbouring canal close to manubrium. In younger specimens some centripetal canals ending blindly. 16 large tentacles (max. 24) with terminal swellings and up to 16 small, hook- shaped tentacles, all tentacles in phase with radial and centripetal canals. Dimensions: Bell diameter 21-34 mm, height up to 38 mm (Kramp, 1959). Biology: Occurs in deep and intermediate water layers (Kramp, 1959). Distribution: Calycopsis chimi has been recorded in the ERMS zone off the coast of Morocco and near the Canary Islands (Kramp, 1959). It is known to occur in the Indian Ocean from East Africa to the Malayan Archipelago, along the east coast of Australia, the Gulf of Panama, the West Indies, and the Atlantic coast of West Africa (Kramp, 1968). Type locality: Gulf of Aden, 1200 to 2000 m depth. Remarks: The high numbers of radial canals (including former centripetal canals that have joined the manubrium) characterize this rare species. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 451 Fig. 5 Calycopsis chimi Vanhöffen, 1911; modified after Kramp (1959) and Vanhöffen (1911), bell size about 3 cm. Calycopsis bigelowi Vanhöffen, 1911 Fig. 6 Calycopsis bigelowi Vanhöffen, 1911: 218, fig. 12. - Kramp, 1957: 21, map fig. 4. - Kramp, 1959:127, fig. 136. -Kramp, 1961: 119. -Kramp, 1968: 56, fig. 149. - Van der Spoel & Bleeker, 1988: 167, fig. 16. - Schuchert, 1996: 26, fig. 10a-b. Type material: Holotype in Naturkundemuseum Berlin, Germany, ZMB Cni 14837 (not seen). Material examined: BMNH 1957.2.1.760-764; Discovery stations 100, 100c, 101, 983, west of Cape Good Hope; 6 mature medusae, some rather damaged, material described in Kramp (1957). - See also Schuchert (1996). Diagnosis: Calycopsis medusa 8-16 mm high and wide, jelly thick, 8-16 long tentacles and up to 40 short ones; four radial canals and four blind interradial centri- petal canals; gonad in each quadrant with two vertical rows of up to 16 deep, regular, transverse clefts. Description: Medusa umbrella ovoid to spherical, jelly thick, soft and adhesive. Velum somewhat shifted into subumbrella. 452 P. SCHUCHERT Fig. 6 Calycopsis bigelowi Vanhöffen, 1911; after Schüchtert (1996). (A) Medusa, only frontal tentacles shown; scale bar 5 mm. (B) Heteroneme, scale bar 10 um. (C) Intact and discharged desmo- nemes, same scale as B. Manubrium large and voluminous, length 2/3 of bell cavity, with cross-shaped base. Mouth with slightly undulating margin and with four small perradial lips. Four large, interradial gonads leaving only perradial parts of manubrium visible. Gonads with eight adradial rows of up to 16 horizontal furrows, occasionally some additional pits. Four complete radial canals, widening near manubrium and forming mesen- teries. Four additional centripetal canals which reach almost to the manubrium top, likely ending blindly, terminal region somewhat broadened. All radial canals are rather broad and are connected to the equally broad circular canal. No tentacular bulbs present. Up to 16 long tentacles, four perradial, four interradial, and up to eight adradial ones. Perradial and interradial tentacles usually fully developed in mature animals and longer than bell height, most of the adradial tentacles are also fully developed but some are shorter and in mid-development. In addition to these 16 long tentacles, 16-40 short EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 453 tentacles. All tentacles hollow and with bases adnate to the umbrella margin. Long tentacles end in a terminal swelling with a concentration of nematocysts. Shorter tentacles without terminal swelling, but with a concentration of nematocysts. Without ocelli. Nematocysts: rare heteronemes (Fig. 6B) and frequent, elongated desmonemes (Fig. 6C). Desmonemes on tentacle tips, discharged with five coils, thread with spiral pattern of small bristles Colours: gonads yellow-green colour (Kramp, 1957) or red (Vanhöffen, 1911). Dimensions Mature medusa 9-16 mm high and wide, umbrella wall 3 mm. Desmonemes (ll-12)x(5-6)^m, heteronemes (14-15)x(7-8),um (Schuchert, 1996). Biology: Oceanic species, prefers deep waters down to 2500 m (Kramp, 1957), usually around 400-600 m. Distribution: Gulf of Aden; South Africa; Indo-Malayan region; tropical Indian Ocean; New Zealand; Scotland, ? British Columbia (Kramp, 1965, Van der Spoel & Bleeker, 1988; Fraser, 1974; Schuchert, 1996; Brinckmann-Voss & Arai, 1998). Type locality: Gulf of Aden, depth 1 100 m. Remarks: This is a rare species and in the ERMS region it is only known from a single specimen collected in deep water west of Scotland (Fraser, 1974). A very similar species is Calycopsis gara Petersen, 1957. It differs from C. bigelowi in having more (up to 46) long tentacles and four to eight centripetal canals. As only very few specimens of C. bigelowi and C. gara are known, their morphological variation is not sufficiently known and the status of C. gara is somewhat uncertain. Calycopsis gara Petersen, 1957 Fig. 7 Calycopsis gara Petersen, 1957: 29, fig. 1. - Kramp, 1959: 127, fig. 137. - Kramp, 1961: 120. - Winkler, 1982: 37. - Bleeker & Van der Spoel, 1988: 231, fig. 17. Type material: ZMUC (not seen). Diagnosis: Calycopsis medusa 10-20 mm high, 8 radial canals, sometimes a few additional centripetal canals, gonads with 15-16 transverse clefts in eight vertical rows, 40-50 tentacles. Description (Petersen, 1957): Medusa umbrella ovoid to spherical, evenly rounded, somewhat laterally compressed along two interadii, jelly thick. Manubrium large and voluminous, length 2/3 of bell cavity, half as wide as long, with cross-shaped base. Mouth rim smooth, four small perradial lips. Four large, interradial gonads leaving only perradial parts of manubrium visible. Gonads with eight vertical, adradial rows of up to 16 horizontal furrows, some can be irregular. The perradial edges of the stomach form meridional ridges which in their uppermost parts connect with the radial canals to form short funnels (=mesenteries). Four radial canals, rather wide, usually four centripetal canals joining the base of the manubrium, sometimes additional blind centripetal canals. No tentacular bulbs present. 40-50 tentacles, some short, ending in a terminal swelling with a concentration of desmonemes. 454 P. SCHUCHERT Fig. 7 Calycopsis gara Petersen, 1957; modified after Petersen (1957), bell size 11 mm. Dimensions: Bell diameter 11-21 mm, height 10-22 mm (Winkler, 1982). Biology: All known records are from less than 200 m depth. Distribution: Central North Atlantic Ocean (Petersen, 1957; Winkler, 1982; Bleeker & Van der Spoel, 1988). Type locality: 50.883°N 34.417°W, 0-100 m. Remarks: See above under C. bigelowi. Genus Sibogita Maas, 1905b Type species: Sibogita geometrica Maas, 1905b. Diagnosis: Bythotiaridae with four primary radial canals which branch repea- tedly at various levels; no centripetal canals, gonads with eight rows of transverse furrows. No ocelli. Polyps unknown. Remarks: This is currently a monotypic genus. Bigelow (1919) assumed that the side branches of the radial canals were actually centripetal vessels that arose from the ring canal and only later in life joined the radial canals. He therefore synonymized the genus with Calycopsis. Kramp (1959) outlined the taxonomic history of the genus and contested Bigelow's view of the growth direction of the radial canals. Kramp thinks that the branching radial canals arise indeed through branching during ontogeny and he thus has an argument to maintain the genus. The diminution of the canal width towards the periphery is a good argument in favour of this view. Nevertheless, the EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 455 difference to Bytothiara is not so clear-cut and Sibogita can admittedly be seen as a sy- nonym of the former. As we neither know the polyp stage of Bythotiara murrayi nor of Sibogita geometrica, further discussions on the validity of Sibogita are premature. Sibogita geometrica Maas, 1905b Fig. 8 Sibogita geometrica Maas, 1905b: 17, pi. 3 figs 16-18. - Mayer, 1910: 186, fig. 99. - Kramp, 1965: 49. - Kramp, 1968: 57, fig. 151a-b. - Van Soest, 1973: 121, fig. 2c. - Winkler, 1982: 37, figs 9-11. - Bleeker & van der Spoel, 1988: 234, figs 18-19. - Pages et a/., 1992: 9, fig. 9. Sibogita geometrica occidentalis Kramp, 1959: 28, 129, pi. 2 figs 2-3, text-figs 4-5. Calycopsis geometrica. - Bigelow, 1919: 290, pi. 40 fig. 5-7, pi. 41 fig. 2. Type material: ZMA COEL no 3853 (not seen). Diagnosis: See genus diagnosis. Description: Medusa umbrella globe- to pumpkin-shaped, top rather flat or even slightly concave, walls straight, thick, bell can be laterally compressed, without gastric peduncle, bell-margin undulated through embayments for tentacles. Manubrium relatively short, square to cross-shaped in section, 1/4 to 1/2 of the height of the subumbrellar cavity, mouth with four more or less distinct lips, mouth rim undulated. Four large, interradial gonads with eight adradial rows of 6-19 horizontal furrows. Four primary radial canals divided irregularly at different levels giving rise to a total of 16-43 radial canals joining the ring-canal, anastomoses of radial canals rarely present, no centripetal canals. Radial canals widening somewhat where they join the ring-canal, intersection containing dark pigment granules. About half as many to as many long tentacles as there are radial canal endings, originating in line with radial canals, base adnate, highly contractile, distal end swollen, club-like. Between successive pairs of long tentacles usually a short, inward- directed tentacle. Living specimen colourless except for faintly yellow manubrium (Kramp, 1959). Dimensions: Bell height usually around 2 cm (max. 4 cm), diameter 2 cm. Winkler (1982) gives the following data for Atlantic specimens: height 15 mm, diameter 21 mm, jelly 3 mm, manubrium 8 mm long and 5 mm wide. Kramp (1965) gives a table with the correlation of the bell-size and the number of radial canals. Biology: A quite rare, oceanic species, usually found at depths of some 100 metres. Bleeker & van der Spoel (1988) found evidence for diel vertical migration as all night samples taken were above and all day samples below 200 m depth. Distribution: Malayan Archipelago and tropical Indian Ocean (Maas, 1905b; Bigelow, 1919), Bay of Biscay, Azores and Mid-Atlantic (Kramp, 1959; Van Soest, 1973; Winkler, 1982; Bleeker & van der Spoel, 1988), Benguela Current off South Africa (Pages et a/., 1992) Type locality: northeast of Sulawesi, 1.633°N 124.46°E, 0-900 m. Remarks: Kramp (1959) thought that the Atlantic population differs slightly from the Pacific one and he introduced for the former the subspecies Sibogita geo- 456 P. SCHUCHERT Fig. 8 Sibogita geometrica Maas, 1905b. (A) Medusa from Pacific in side-view, bell height 38 mm, modified after Maas (1905) and Mayer (1910). (B) Aboral view of medusa from Atlantic showing branching pattern of radial canals, from Winkler (1982), scale bar 3 mm. metrica occidentalis. The differences to the nominal Pacific species were given as a smaller bell-size, fewer gonadal folds, and radial canals dividing already close to the centre. After having seen more specimens from the Pacific, Kramp (1965) then syno- nymized the subspecies occidentalis with the nominal Pacific species. Also other authors (e. g. Winkler, 1982; Bleeker & van der Spoel, 1988) confirmed this. Both forms occur in the Atlantic and the Pacific and they are connected by intermediate forms. Family Proboscidactylidae Hand & Hendrickson, 1950 Synonyms: Willsiadae Forbes, 1848 [family name incorrectly derived from genus Will sia]. - Laridae Hincks, 1868 [preoccupied by Laridae Rafinesque, 1815, Seagulls]. - Hydrolaridae Allman, 1872 [type species Lar sabellum Gosse, 1857 by monotypy]. - Williadae Haeckel, 1879 [invalid emendation of original name]. - Williadi Mayer, 1910. - Willsiidae Stechow, 1913 [correct derivation of name]. Diagnosis (Bouillon et at., 2006): Hydroid usually on rim of sabellid poly- chaete tubes, with creeping, naked stolons; hydranths almost sessile, polymorphic; EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 457 gastrozooid with rounded hypostome, separated from body by a constriction; hypo- stome with large pad of nematocysts somewhat displaced onto one side, two filiform tentacles arising close together beneath hypostomial constriction, opposite to nema- tocyst cluster; gonozooids and dactylozooids without tentacles, mouth-less and smaller than gastrozooids; medusa buds close to gonozooid tip. Medusa umbrella mostly hemispherical; with exumbrellar nematocyst buttons or bands alternating with tentacles; radial canals branched, obliterated canals some- times present; usually instead of circular canal a solid gastrodermal marginal strand; manubrium base with four, six or more radial gastric lobes, extending along proximal portions of radial canals, lobes in some species inconspicuous; gonads surrounding manubrium and extending onto gastric lobes; tentacles hollow, with swollen hollow base connected to the lumen of radial canals. Remarks: For the taxonomic history and naming of this family see e. g. Browne (1905), Maas (1905a), Bigelow (1909a), Hartlaub (1917), Stechow (1923), Kramp (1939), and Hand & Hendrickson (1950). Hand and Hendrickson (1950) argued that the family name must be changed to Proboscidactylidae because Uchida & Okuda (1941) had previously synonymized Willsia with Proboscidactyla. But this renaming was not necessary, and the correct and valid name of this family would have been Willsiidae, as was argued by Stechow (1923). The name Willsiidae has, however, hardly been used and after 1950 many general treatises (e. g. Russell, 1953; Kramp, 1961) adopted the name Probosci- dactylidae. Later authors exclusively used this name. In the interest of nomenclatural stability, the usage of the name Proboscidactylidae should thus be continued. This is explicitly endorsed by article 40.2 of the ICZN, because Hand and Hendrickson (1950) replaced the name Willsiidae with Proboscidactylidae before 1961. Kramp ( 1939) referred the family to the Limnomedusae (subclass Trachylinae). The occurrence of desmonemes, the gonads on the manubrium, and the lack of stato- cysts argue strongly against this. It is nowadays again included in the Anthomedusae (=Anthoathecata = Athecata). Molecular phylogenies confirmed that they do not belong to the Trachylinae but to the Hydroidolinae, although the precise relationships within the latter subclass are not yet resolved (Cartwright et al., 2008). This family currently comprises only the genus Proboscidactyla (see Bouillon et al, 2006). Schuchert (1996) also included the genus Fabienna Schuchert, 1996 in this family, replacing in part Pochella Hartlaub, 1917, a genus that had originally been included in the Proboscidactylidae by Russell (1938a) (now in Trichydridae). Although Fabienna has a close affinity with the family Proboscidactylidae (Schuchert & Reiswig, 2006; Cartwright et al., 2008), it is here in the interim kept separate in the family Magapiidae (former Laingiidae) following Bouillon et al. (2006). See also the remarks for this family. The medusae of the Proboscidactylidae have reportedly a solid strand instead of a radial canal. It is not clear if is true for all species. A detailed histological of several species would be most helpful. Genus Proboscidactyla Brandt, 1835 Synonyms: Willsia Forbes, 1846 [type species Willsia stellata Forbes, 1846 by mono- r ypy]- - Lar Gosse, 1857. - Willia Agassiz, 1862 [invalid emendation]. - Dyscannota Haeckel. 458 P- SCHUCHERT 1879 [type species Dyscannota dysdipleura Haeckel = P. ornata]. - Dicranocanna Haeckel, 1879 [type species Dicranocanna furcillata Haeckel, 1879 = ? P. ornata]. - Willeta Haeckel, 1879. - Misakia Uchida, 1927 [type species Misakia typica Uchida, 1927, synonym of Proboscidactyla ornata]. - Psythia Agassiz & Mayer, 1902 [type species Psythia prolifera Agassiz & Mayer, 1902]. Type species: Proboscidactyla flavicirrala Brandt, 1835. Diagnosis: As for family. Key to the genera of the Proboscidactyla species of the ERMS zone: la medusa with 4 primary radial canals, occurring in Mediterranean ... P. ornata lb medusa with 6 primary radial canals, occurring in northern Atlantic . P. stellata Proboscidactyla ornata (McCrady, 1859) Figs 9-10 Willsia ornata McCrady, 1859: 149, pi. 9 figs 9-1 1. Willia ornata. -Agassiz, 1865: 171, figs 274a-279. - Fewkes, 1882b: 299, pi. 1 figs 22-24. Dyscannota dysdipleura Haeckel, 1879: 152. ? Dicrocanna furcillata Haeckel, 1879: 156. Willetta ornata. - Haeckel, 1879: 157. Willia gemmifera Fewkes, 1882b: 300, pi. 1 fig. 24. Proboscidactyla ornata. - Browne, 1905: 726. - Mayer, 1910: 189, fig. 100, pi. 20 figs 1-10. - Neppi & Stiasny, 1913: 35, pi. 2 fig. 24, pi. 3 fig. 25. - Hartlaub, 1917: 368, figs 316- 323. - Menon, 1932: 12, pi. 2 fig 18. - Kramp, 1957: 13, pi. 3 fig. 7. - Kramp, 1961: 235, synonymy. - Kramp, 1962: 342, figs 7-10, synonymy. - Kramp, 1959: 178, fig. 255. - Kramp, 1965: 103. - Kramp, 1968: 108, fig. 290. - Calder, 1970: 130, fig. 1. - Calder, 1971: 44, pi. 3 fig. A. pi. 7 Fig. A. - Brinckmann & Vannucci, 1965: 357, figs 2-6. - Bouillon et al., 2004: 74, fig. 43C-D. - Goy, 1973: 997. Proboscidactyla tropica Browne, 1905: 727. Proboscidactyla gemmifera. - Browne, 1905: 727. Proboscidactyla varians Browne, 1905: 728, pi. 54 figs 1-2. Proboscidactyla flavicirrata var. stolonifera Maas, 1905b: 21, pi. 4 figs 24-28. Proboscidactyla ornata var. stolonifera. - Bigelow, 1909a: 220, pi. 6 figs. 1-2: pi. 41 figs. 1-7. - Mayer, 1910: 191, fig. 10L Proboscidactyla ornata var. gemmifera. - Mayer, 1910: 192, fig. 101a, pi. 21, figs 1-3. Misakia typica Uchida, 1927: 237, fig. 15. ? Proboscidactyla conica Menon, 1932: 13, pi. 2 fig. 18. Proboscidactyla mutabilis. - Nair, 1951: 57. [not Proboscidactyla mutabilis Browne] ? Proboscidactyla ornata. - Goy et al., 1991: 1 18, fig. 46. Type material: Based on a single medusa, not located. Material examined: MHNG INVE 54619; Indonesia, Moluccas, Ambon; as Proboscidactyla flavicirrata var. stolonifera, material of Maas (1906), one medusa, umbrella shrunken. Diagnosis: Proboscidactyla medusa with square- to cross-shaped stomach base, without distinct gastric lobes, 4 primary radial canals, branching 1-2 times, 12-16 tentacles, medusa-buds may arise from corners of stomach or radial canals. Medusa buds on polyps with 4 tentacle bulbs. Description (Mayer, 1910; Brinckmann & Vannucci, 1965; Calder, 1970): Hydroid colonial, arising from creeping, apparently naked stolons, polyps usually confined to end of polychaete tube, polyps polymorphic with gastrozooids and gono- zooids (blastostyles). Gastrozooids in a single circle around the rim of the worm tube, almost sessile, body near base somewhat constricted and thus forming an indistinct EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 459 Fig. 9 Proboscidactyla ornata (McCrady, 1859); from Brinckmann & Vannucci (1965). (A) Hydroid on rim of polychaete tube, some incipient gonozooids (blastostyles) at bases of gastrozooids are visible; scale bar 0.5 mm. (B) Gonozooid with advanced medusa bud, at rear a gastrozooid; scale bar 0.25 mm. pedicel, oral end enlarged to form a head-like proboscis separated from the rest of body by a constriction (neck). Two smooth tentacles arise close together from base of neck, tentacles arise on side facing the opening of the polychaete tube and give the hydranths a bilateral symmetry. Apex of proboscis with a cap-like nematocyst cluster containing white pigment granules. Gonozooids usually smaller than gastrozooids (max 3/4), arising near base of gastrozooid on the side opposite the tentacles, body thin and tentacle-like, terminating in a small, spherical nematocyst cluster, without mouth. Gonozooids of more advanced stages can also be isolated from gastrozooid, occasionally several mm down the worm tube, but remaining in contact with the rest of the colony via the hydrorhiza. Gonozooids with up to four medusa buds in a single whorl located in upper half of polyp, buds in more advanced stages with four large tentacle bulbs. Nematocysts: des- monemes, microbasic euryteles of two size classes, and large macrobasic euryteles with indistinct swelling of shaft and thus difficult to distinguish from mastigophores. Newly liberated medusa bell-shaped, with umbilical canal, stomach cylindrical and about half the length of the subumbrellar cavity, lips simple, four unbranched radial canals, no gonads present. Alternating with the tentacles are small clusters of nematocysts, evenly spaced on the exumbrella just above the margin. Nematocysts: macrobasic euryteles and desmonemes with unusually long threads in many loops, discharged with up to 15 loops. 460 P. SCHUCHERT Mature medusa bell-shaped, slightly higher than wide; jelly thick, rigid, shallow gastric peduncle can be present, near rim of exumbrella up to 16 small nematocysts clusters, one in-between each tentacle-pair, connected to rim of umbrella by meridional line on exumbrella. Manubrium flask-shaped, height 1/2 to 1/1 of subumbrellar height, base square- to cross-shaped, four perradial recurved lips, mouth margin with some crenulations. Gonads on sides of stomach, may extend onto radial canals, surface smooth. Four primary radial canals, narrow, branching up to two times in well spaced branching points so that normally 16 canals reach the bell margin, rarely up to 20 ca- nals, some populations with only up to 12 canals. No ring canal, but instead a solid strand of gastrodermal cells. Usually as many tentacles as radial canals (around 16, range when mature 4-20), with relatively large marginal bulbs containing a brownish pigment. Some populations have medusae that form stolon-like blastostyles bearing me- dusa buds, these blastostyles may arise from the corners of the stomach (= gemmifera form) or the branching points of the radial canals (= tropica form). Colours of Eastern Atlantic form: gastrodermis ochre-yellow or greenish-yellow, bulbs brown. Nemato- cysts: not described, but likely as in young medusa. Dimensions: Gastrozooids in nature 0.35-1.3 mm high, tentacles up to 1.5 mm long (Brinckmann & Vannucci, 1965; Calder, 1970). In culture, the polyps get larger (up to 1.8 mm, Brinckmann & Vannucci, 1965). The gonozooids are smaller (max. 3/4 of gastrozooids). Polyp nematocysts (Mediterranean form, Brinckmann & Vannucci, 1965): small microbasic euryteles (5-6)x(2-2.5),um; large microbasic euryteles (7.5- 9.5)x(2.5-3.5)/ym; macrobasic euryteles (16-19)x(9-12.5)^m. For data of a Western Atlantic population see Calder (1970). The newly liberated medusa is about 1 mm in height and 0.8 mm wide (Brinckmann & Vannucci, 1965). The mature medusa can reach bell diameters of up to 5 mm (Kramp, 1968), but in the Mediterranean they remain much smaller (about 1 mm, Goy, 1973; Goy et ai, 1991; 3.4 mm Neppi & Stiasny, 1913) and they reach rarely or never the 16 tentacle stage. They can even have mature gonads at the four- tentacle stage. For nematocyst dimensions see Brinckmann & Vannucci, (1965) and Calder (1970). Biology: The polyps occur exclusively on the tubes of sabellid polychaetes. Calder (1970; east coast of the USA) found it on Sabella microphthalma Verrill living in somewhat brackish waters (salinities from 18ppt to 23ppt). In the Mediterranean, Brinckmann & Vannucci (1965) found it on the tubes of Branchiomma vesiculosum (Montagu) and other sabellids. In the Bay of Naples, the polyps are locally quite abun- dant (in depths of about 100 m), but the medusa is very rare (Brinckmann & Vannucci, 1965). It is also rare in other regions of the Mediterranean (Mediterranean (Neppi & Stiasny, 1913; Goy, 1973; Goy et al, 1991). The liberated medusa develops gonads rather rapidly (after 2-6) days and is fully grown within 3-4 weeks (Brinckmann & Vannucci, 1965; Calder, 1970). The medusa seems not to tolerate temperatures below 17-18°C (Brinckmann & Vannucci, 1965). It can be found close to the surface (Kramp, 1965). EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 461 Fig. 10 Proboscidactyla ornata (McCrady, 1859); modified after Mayer (1910), see text for dimensions. (A) Young medusa. (B) Mature female medusa. (C) Immature medusa with medusa buds on the corners of the stomach. (D-G) Sequence of the branching of the radial canals. Uchida & Sugiura (1975) examined the medusa budding. Kawamura & Kubota (2008) investigated the influence of temperature and salinity on the asexual budding of the medusa. Many biological observations made by Hand & Hendrickson (1950) on a Californian Proboscidactyla species are certainly also valid for P. ornata (see also Hand, 1954). 462 P. SCHUCHERT Distribution: The medusa has a circumglobal distribution in warm and coastal waters (Kramp, 1965). In Europe it occurs only in the Mediterranean (Bouillon et al., 2004). The polyp stage is known from the Chesapeake Bay, USA (Calder, 1970) and the Mediterranean (Brinckmann & Vannucci, 1965). Type locality: Charleston Harbor, South Carolina, USA. Remarks: The life cycle of this species has been observed by Brinckmann & Vannucci (1965) for Mediterranean animals and by Calder (1970) for hydroids from Virginia, USA. Calder tabulates also differences between the two populations, mainly concerning the size of the nematocyst capsules. He interpreted the small differences as due to intraspecific variation and different culture conditions. The medusae of P. ornata are quite variable and a number of species and sub- species have been proposed, mainly based on the presence and position of vegetative medusa buds. Kramp (1957; 1965) found good evidence that all these variants likely belong to the same species. Kramp (1962) also found a single medusa in Vietnam with a polyp growing out of the manubrium. This polyp had a dome-shaped hypostome sur- rounded by a whorl of 1 8 tentacles, thus is very much unlike all other Proboscidactyla polyps known so far. The hydroids of the Pacific P. ornata have never been observed and it remains to be shown that they are identical to the Atlantic and Mediterranean ones. It could well be that our current concept of P. ornata nevertheless comprises several species. The Mediterranean P. ornata medusae are relatively small and develop fewer tentacles and radial canals (max. 12). Gonads can even be present at the four tentacle stage and with unbranched radial canals (Neppi & Stiasny, 1913; Goy, 1973). The medusa from Lebanese waters described by Goy et al. (1991) also had these charac- teristics, in addition to medusa buds on the manubrium. This location of the medusa buds has so far never been observed in other populations. Proboscidactyla stellata (Forbes, 1846) Figs 11-12 Willsia stellata Forbes, 1846: 268. - Mayer. 1910: 193. Lar sabellarum Gosse, 1857: 113, pi. 20. -Hincks, 1868: 36, fig. 2. - Hincks, 1872: 313, pi. 19. - Browne, 1896: 468, pi. 16 figs 3-4, synonym. - Browne, 1897: 818, figs 1-9. Willia stellata. - L. Agassiz, 1862: 346. [incorrect subsequent spelling] Willia stellata. - Browne, 1905: 725, 729. - Hartlaub, 1917: 374. figs 324-328. - Ranson, 1937: 323, fig. 1. - Russell, 1938b: 154, fig. 45. - Kramp, 1939: 503, figs 1-5. Willisa cornubica Peach, 1867: 355, pi. 1 figs 1-2. - Russell, 1953: 393, synonym. Willia furcata Haeckel, 1879: 158. - Russell, 1953: 393, synonym. ? Proboscidactyla brooksi Mayer, 1910: 194, fig. 101C-F. - Russell 1953: 393, ? synonym. Proboscidactyla stellata. - Russell, 1953: 386, figs 250-256, pi. 23 figs 3-4. - Hand, 1954: 64. - Kramp. 1959: 178, fig. 256. - Kramp, 1961: 236, bibliography. - Pages et al,. 1992: 37, fig. 41. - Galea, 2007: 29. pi. 1 fig. J. - Buecher et al, 2005: 44. Type material: Not located, likely lost. Material examined: France, Roscoff ; living medusae, used for DNA extraction; 2 April 1998; plankton in 10 m depth; 16S sequence accession number AM183138. - BMNH 1962. 1 1.7.34, polyps with medusae buds; Norway, Bergen, Espegrend; 30 m depth; collected 09.08.1962; on Pseudopotamilla renifonnis (curled tube). - BMNH 1985.9.1.13 polyps with medusae buds: Great Britain, Plymouth; collected 28.06.1937; leg. & det. W. J. Rees. Diagnosis: Proboscidactyla medusa with star-shaped stomach with 6 lobes and 6 primary radial canals, 24 tentacles. Medusa buds on polyps have 6 tentacle bulbs. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 463 Fig. 1 1 Proboscidactyla stellata (Forbes, 1846); A-B, after preserved material from Norway, C-F modi- fied after Russell (1938b). (A) Gastrozooids and gonozooids without medusa buds on the rim of a polychaete tube. Note that the tentacles are directed towards the opening of the tube, while the nematocyst clusters on the heads are directed in the opposite direction, scale bar 0.2 mm. (B) Gonozooid with medusa buds of different developmental stages. (C) Discharged and intact des- moneme, scale bar 10 urn. (D) Discharged and intact large microbasic eurytele, same scale as C. (E) Discharged and intact small microbasic eurytele, same scale as C. (F) Intact macrobasic heteroneme, same scale as C. Description (Russell, 1938b, 1953; own observations): Hydroid colonial, arising from creeping, apparently naked stolons, covering region near rim of poly- chaete tube, stolons net-like, more or less parallel to worm tube and with connections at right angles. Polyps usually confined to end of polychaete tube, polyps polymorphic with gastrozooids and gonozooids (blastostyles), gonozooids without medusa buds often present, can be interpreted as dactylozooids. Gastrozooids in a single circle around the rim of the worm tube, almost sessile, body near base somewhat constricted and thus forming an indistinct pedicel, oral end enlarged to form a head-like proboscis separated from the rest of body by a constric- 464 P. SCHUCHERT tion (neck). Two smooth tentacles arise close together from base of neck, tentacles arise on side facing the opening of the polychaete tube. Proboscis with a cap-like nematocyst cluster on side opposite to the tentacles. The nematocyst cluster and the two tentacles give the hydranths a bilateral symmetry. Gonozooids usually smaller than gastrozooids (1/2), arising on stolons close to gastrozooid on the side opposite the tentacles, body thin and tentacle-like, terminating in a small, spherical nematocyst cluster, without mouth. Medusa buds in upper part of gonozooids, up to four in one whorl, buds not covered by membrane, buds in more advanced stages with six large tentacle anlagen (bulbs). Some gonozooid-like polyps without medusa buds can be present and may act as defensive dactylozooids. Nematocysts of polyp stage: large macrobasic heteronemes (mastigophores or euryteles with faint swelling of shaft end), present only on proboscis; microbasic euryteles of two size classes; desmonemes with unusually long threads in many coils, discharged with up to seven coils. Newly liberated medusa bell-shaped, with umbilical canal, velum broad, stomach cylindrical and about half the length of the subumbrellar cavity, attached to a very short gastric peduncle, gonads not yet developed, base with six short prolon- gations or lobes from which six primary radial canals start, radial canals unbranched. Six marginal tentacles, equal in size, on margin of umbrella, one opposite termination of each radial canal. Base of each marginal tentacle large and conspicuous, containing dark brown or blackish pigment. Alternating with marginal tentacles are small clusters of nematocysts, evenly spaced on exumbrella just above bell-rim. During the ensuing growth of the medusa the radial canals branch three times. Umbrella of mature medusa dome-shaped, slightly wider than high; jelly thick. Velum narrow. On exumbrella near rim small pads of nematocysts, usually one per tentacle and one in-between each tentacle-pair (around 24 in total), the two series at different levels, usually a number of pads lost, all connected to margin of bell by a thin, meridional line of thickened epidermis. Stomach short, six-sided, with six basal lobes prolonged for a short distance over the subumbrellar surface resulting in star-shaped base. Mouth with six folded lips. Very short gastric peduncle. Gonads situated at base of stomach wall and continued along the six lobes of stomach. Six primary radial canals, each giving rise to three additional branches making twenty-four in all; all branching subject to considerable variation. Radial canals narrow. No ring canal, but instead solid strand of gastrodermis. Twenty-four rather short marginal tentacles, each situated at end of a radial canal branch, filiform, hollow, with adaxial basal nematocyst cushion. Colour of tentacle bases yellowish brown, dark purple-brown or black; colour of stomach and gonads yellow or reddish yellow. Nematocysts: not described. Variation: Rarely only four primary canals are present. The population of Japan shows more variability (Uchida, 1927). Dimensions: Medusa when full grown 8 mm high and 9 mm wide. Newly liberated medusa about 1 mm high. Gastrozooids 0.6-2.5 mm high. Nematocysts of polyp stage (Russell, 1938b): large macrobasic heteronemes (24-33)x(9-13),um; euryteles (9-13)x(4-5)^/m, and (6.5-7)x(2.5-3)/jm; desmonemes 10x5^m. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 465 Fig. 12 Proboscidactyla stellata (Forbes, 1846); A-C, modified after Russell (1953); D-I, modified after Browne (1897). (A) Newly liberated medusa, bell height approx. 1 mm. (B) Mature medusa, bell diameter 4.2 mm. (C) Rim of medusa in side view showing tentacle and exumbrellar nematocyst clusters (arrow). (D-G) Development of the tentacles and the branching of the radial canals, pre = primary radial canal, 1 = first side-branch, 2 = second side branch, etc. (H-I) Irregular canal branching. Biology: The life cycle of this species was elucidated by Browne (1896). The hydroid occurs exclusively on the tubes of sedentary polychaetes like Megalomma vesiculosum (Montagu), Potamilla torelli (Malmgren), Pseudopolamilla reniformis (Müller), or Sabellaria spec. The known depth range is from subtidal to about 50 m. (Gosse, 1857; Hincks, 1872; Russell, 1957; own data). In the British Isles, the medusa may be found in any month of the year (Russell, 1953; Ballard & Myers, 2000). It occurs in the upper water layers. Distribution: In Europe from the northern part of the Bay of Biscay over the British Isles to Norway (Hartlaub, 1917; Rees, 1952; Russell, 1953; Russell, 1957; 466 P. SCHUCHERT Kramp, 1959; Teissier, 1965; Fraser, 1974; Ballard & Myers, 2000). The medusa has also been reported from Japan (Uchida, 1927), South Africa (Pages et al., 1992; Buecher et al., 2005), and the Fjords of Chile (Galea, 2007). Perhaps also present along the Atlantic coast of the USA (Kramp, 1961, as P. brooksi). Type locality: Bay of Oban, Scotland. Remarks: Russell (1938b) identified the large heteronemes of the polyp as mastigophores, although he depicted one shaft that is clearly a eurytele. Hand & Hendrickson (1950) concluded that there are likely only euryteles in P. stellata and that Russell's mastigophores were a fixation artefact. In other Proboscidactyla hydroids the large heteronemes are of the eurytele type (Hand & Hendrickson, 1950; Brinckmann & Vannucci, 1965). Some heteronemes of hydrozoans can have a faint, variable swelling of the shaft and cannot be classified unambiguously as mastigophores or euryteles. Similar observations were made by Brinckmann & Vannucci (1965). Family Magapiidae Schuchert & Bouillon, 2009 Diagnosis (modified after Bouillon et al., 2006): Medusa umbrella almost hemispherical, margin lobed through furrows for tentacles; four radial canals; four tentacles with nematocysts concentrated terminally; no typical circular canal but a solid strand of gastrodermis; tentacles solid, issuing on exumbrellar surface above bell margin; alternating with tentacles there may be narrow exumbrellar nematocyst bands or triangular ciliated fields; manubrium simple, tubular or with cruciform base, mouth opening quadrangular to circular, no lips; gonads in four masses on the manubrium or on interradial pockets of the manubrium; no sense organs; cnidome: macrobasic masti- gophores or macrobasic euryteles. Hydroid stage unknown. Remarks: This family was previously known as Laingiidae. Because the genus name Laingia is preoccupied, Schuchert & Bouillon (2009) introduced the replacement names Magapia and Magapiidae. The family Magapiidae is here placed in the order Anthoathecata and not in a subclass Laingiomedusae of its own as suggested by Bouillon (1978b). The medusae of the Magapiidae have a lobed umbrella margin and tentacles originating more or less on the exumbrellar side of the bell. They thus exhibit some characters otherwise found in the Narcomedusae. However, a similar situation is also found in Thecocodium quadratimi (Anthoathecata, family Ptilocodiidae; see Jarms, 1987). In any case, with their gonads on the manubrium and the fully developed radial canals they match more typically anthomedusae rather than Narcomedusae and they are thus here classified as Anthoathecata Filifera. Molecular analyses (Collins et ai, 2006; Cartwright et al., 2008) have shown that Fabienna, a genus placed in Laingiomedusae by Bouillon & Barnett (1999), does not belong to the Trachylinae and is closely related to some Proboscidactyla species. Admittedly, a final decision on the status of the Laingiomedusae can only be made once the position of its name-giving species, Laingia jaumotti Bouillon, 1978b, has been determined by a molecular phylo- genetic analysis. european athecate hydroids, filifera 5 467 Key to the Laingiidae genera: la with exumbrellar nematocyst bands Kantiella lb without exumbrellar nematocyst clusters 2 2a bell margin with interradial ciliated fields; marginal bulbs only slightly displaced from bell rim Fabienna 2b no ciliated fields, tentacles shifted high up to exumbrella [Magapia]* * not in ERMS zone Genus Kantiella Bouillon, 1978 Type species: Kantiella enigmatica Bouillon, 1978. Diagnosis: Magapiidae medusa with radial exumbrellar nematocyst bands; gonads on four interradial pouches of manubrium; four short marginal tentacles with terminal cluster of nematocysts, tentacle bulbs not on bell margin but on exumbellar side at some distance from margin. Remarks: The genus is currently monotypic. Kantiella enigmatica Bouillon, 1978a Fig. 13 Kantiella enigmatica Bouillon, 1978a: 158, fig. 13-14. - Bouillon, 1978b: 477, fig. 2, pi. 1. figs 1-3. -Goy et al., 1991: 116, fig. 43. Holotyope: In IRSN (Bouillon et al., 1995), not seen. Material examined: IRSN, Papua New Guinea, Laing Island; 20 medusae collected July 1978 by J. Bouillon. Diagnosis: See genus diagnosis. Description: Medusa umbrella hemispherical, mesogloea thick at apex (ca. 1/3 of total height), margin lobed through deep perradial embayments where tentacles originate, 4-8 short radial exumbrellar nematocyst bands originating from margin; short and broad gastric peduncle present. Manubrium quadrangular, reaching almost to level of velum, mouth margin quadrate or circular, without marked lips, mouth rim with nematocysts. With or without medusa buds at apex of radial canals. Gonads covering interradial sides of manubrium, bulging, separated perradially by cleft. Four simple radial canals, instead of circular canal a solid strand of gastro- dermi s . Four short, solid tentacles, bulbs large, placed on exumbrellar side at some dis- tance from margin, with short radial connection to circular strand, mesogloea of bell reduced below tentacles; nematocysts of tentacles confined to tip, initially in a termi- nal and an adaxial subterminal cluster, later both clusters fused. Nematocysts: ovoid holotrichous macrobasic mastigophores, on mouth rim, tentacle tips, tentacle bulbs, and exumbrellar bands. Polyp stage unknown. Dimensions: Umbrella 3-4 mm wide, 2-3 mm high (Bouillon et al., 2004). Macrobasic mastigophores (ll-23)x(7-18) pm (Bouillon, 1978b). Biology: A coastal species occurring in shallow depths. 468 P. SCHUCHERT Fig. 13 Kantiella enigmatica Bouillon, 1978a; after preserved material from Papua New Guinea, bell diameter 4 mm. Distribution: Seychelles (Bouillon, 1978a), Papua New Guinea (Bouillon, 1978b), South Africa (Pages et al, 1992), eastern Mediterranean (Goy et al, 1991), Taiwan Strait (Xu & Huang, 2004). Type locality: Seychelles. Remarks: This species does not occur along the European coasts, but it has been found in the eastern Mediterranean (Lebanon; Goy et al, 1991), thus within the ERMS zone. Genus Fabienna Schuchert, 1996 Type species: Fabienna sphaerica Schuchert, 1996 by original designation. Diagnosis: Medusae with slightly lobed umbrella margin, four perradial tentacles that have their origin slightly displaced away from the bell margin, without exumbrellar nematocyst bands but instead interradial ciliary fields near umbrella margin. Manubrium with cruciform base, mouth simple. Nematocysts of tentacles concentrated in tip in one terminal cluster immediately followed proximally by a second adaxial cluster, the two clusters usually fused in older animals. Radial canal replaced by solid strand of gastrodermis. Cnidome includes macrobasic euryteles, no desmonemes. Gonads four large, interradial pads on manubrium. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 469 Remarks: The genus Fabienna shares with Kantiella the peculiar structure of the tentacles and to some degree also the cnidome (macrobasic heteronemes, absence of desmonemes). Species of both genera also show a strong overall similarity. The ten- tacle structure is likely a synapomorphy and the inclusion of Fabienna in the family Magapiidae by Bouillon & Barnett (1999, as Laingiidae) is therefore correct. Only one Fabienna species occurs in the ERMS zone. Fabienna oligonema (Kramp, 1955) Pochella oligonema Kramp, 1955: 270, fig. 7, pi. 2, fig. 2. - Kramp, 1959: 179, fig. 259. - Kramp, 1961: 233. - Goy et al, 1991:118. - Daly Yahia et al, 2003: 650. Fabienna oligonema. - Schuchert, 1996: 87. - Bouillon et al., 2004: 115, fig. 60A. Material of F. oligonema examined: ZMUC, syntypes; Accra, Goldcoast, Atlantide station 77; 3 specimens examined for Schuchert (1996). The type material is lost as it dried out after it was returned to Copenhagen (observation made in 2005). Material of F. sphaerica for comparison: MHNG INVE 33453; few medusae collected in surface plankton near Narrow Neck Beach, Devonport, New Zealand; 3 1 June to 2 July 2002; DNA sample taken from one specimen, 16S sequence GenBank accession number AM183133, 18S AY920767, 28S AY920797. Diagnosis: See genus diagnosis. Description (Kramp, 1955; own data): Umbrella spherical to dome-shaped, jelly thick, especially at apex (about 1/3 of total height), with broad and shallow gastric peduncle (1/4 of manubrium height), margin somewhat lobed through shallow per- radial embayments where tentacles originate; exumbrella with scattered nematocysts, some interradial ciliated fields may be present near bell margin. Velum narrow. Manubrium pyramidal, base cruciform, mouth simple, opening cruciform to quadrangular, perradial corners of mouth with nematocysts. Gonads four large, bulging, interradial pads covering large part of manubrium, shape ovoid, the four gonads separated perradially. Four simple radial canals, ending in relatively large gastrodermal bulbs, bulbs somewhat displaced away from umbrella margin towards exumbrella, sometimes even with a short canal linking them to circular canal (strand?). Tentacles about as long as bell height, bases adnate to umbrellar margin, solid, single row of gastrodermal cells, tentacle tips sometimes hook-shaped, nematocysts of tentacles concentrated in tip, in younger animals in one terminal and one adaxial, sub- terminal cluster, later both clusters fused, some additional nematocysts distributed along rest of tentacle. Nematocysts: unknown. Hydroid: unknown. Dimensions: Umbrella 2 mm high and wide. Distribution: Occurs in shallow depths. Distribution: A very rare species, known from waters off Lebanon and Algeria (Goy et al, 1991; Daly Yahia et al., 2003) and the Gulf of Guinea (western Africa, Kramp, 1955). Type locality: Accra, Ghana. Remarks: Fabienna oligonema is a very rare medusa, only a few specimens have been reported so far. It does not occur along the European coasts, but it has been found in the ERMS zone. Its currently known distribution suggests that it could also 470 P- SCHUCHERT occur in southern Spain, Italy, or Greece. It is well possible that it has been confounded repeatedly with Hydractinia exigua (see Schuchert, 2008a). This is easily possible for specimens that have lost their tentacles, as it is often the case for plankton-net samples. This species was originally attributed to the genus Pochella Hartlaub, 1917. After the life cycle of the hydroid Tricky dra pudica Wright, 1858 had been revealed (Rees, 1941; Edwards, 1973), it became evident that Pochella polynema Hartlaub, 1917 is the medusa stage of Trichydra pudica. Because Pochella polynema is also the type species of the genus, Pochella thus became a synonym of Trichydra Wright, 1858. Based only on its superficial similarity and pending more information on its life cycle, P. oligonema was provisionally also transferred to Trichydra by Edwards (1973a). Schuchert (1996) then regarded Pochella oligonema Kramp, 1955 as sufficiently dis- tinct from Trichydra to be placed in a genus of its own, namely Fabienna. The genus Fabienna comprises currently only two species, F. sphaerica Schuchert, 1996 and F oligonema. Both are very similar and with the current state of knowledge hardly distinguishable. Schuchert (1996) lists the following differences: F oligonema has a gastric peduncle, the tentacle bulbs get narrower towards the cir- cular strand and are more displaced towards the exumbrella than in F sphaerica, the gonads are more oval in shape compared to triangular in F sphaerica. Unfortunately, a thorough comparison of the species was hampered by the suboptimal preservation of the syntypes of F oligonema. It seems however that in one of the syntypes there were ciliated fields as in F sphaerica. It was not possible to examine the nematocysts, nor could it be seen if the circular canal is solid or hollow. Since my first description of F sphaerica in 1996, 1 have seen more specimens, notably also fully mature living medusae (Fig. 14). The new observations make distinction of the two species even more difficult as fully grown F sphaerica can also have a shallow gastric-peduncle as in F oligonema (the gastric-peduncle of F oligo- nema in the type specimens is much smaller than given in the figure of Kramp (1955). Although F oligonema and F sphaerica could be conspecific, it seems better to retain both names for the time being, if only for biogeographic reasons. We do not know the polyp stage of both species (the youngest stages of F sphaerica had an apical funnel and umbilical canal, thus were very likely budded from a polyp stage) and the cnidome of F oligonema remains unknown. There is thus potential to find more and better defined differences. Evidently, we need a more detailed re-description of F oligonema based on living animals. Family Ptilocodiidae Coward, 1909 Diagnosis: Polyps arising from reticular stolons or encrusting hydrorhiza without apparent perisarc. Polyps stolonal, naked, polymorphic; gastrozooid without tentacles; dactylozooids with four or more capitate tentacles, sometimes filiform tentacles. Gonophores on gonozooids or gastro-gonozooids; developing into fixed sporosacs, eumedusoids, or free medusae. Mature medusa with more or less bell-shaped umbrella, with or without radial exumbrellar furrows, with marginal nematocyst ring from which usually arise several centripetal nematocyst bands or exumbrellar rows of réfringent spots. With four mar- ginal tentacles or tentacles absent. Manubrium with perradial nematocyst clusters, EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 471 Fig. 14 Fabienna sphaerica Schuchert, 1996; drawn after a living, mature male medusa from New Zealand; scale bar 0.5 mm. clusters may be on oral arms; gonads four interradial or eight adradial masses on manubrium; no ocelli. Remarks: For a recent revision of the family see Bouillon et al. (1997) or Bouillon et al. (2006). Key to the Ptilocodidae hydroids: la dactylozooids of two types [Hydrichthella]* lb dactylozooids of one type 2 2a hydrorhiza crust-like, not covered by visible perisarc [Ptilocodium]* 2b hydrorhiza a network of perisarc-protected tube-like stolons .... Thecocodiwn Key to the Ptilocodidae medusae: la no marginal tentacles Tregoubovia lb with marginal tentacles 2 2a four interradial gonads Thecocodiwn 2b eight adradial gonads [Hansiella] * * not represented in ERMS zone Genus Thecocodium Bouillon, 1967 Type species: Thecocodium brieni Bouillon, 1967 by monotypy. Diagnosis: Hydroid with reticulate, tubular hydrorhiza, covered by perisarc. Polyps on stolons, sessile, naked, polymorphic, usually with gastro-gonozooids and dactylozooids. Gastro-gonozooids cylindrical or club-shaped, without tentacles, hypo- 472 P- SCHUCHERT stome with nematocysts. Dactylozooids thin, solid gastrodermis, terminal group of capitate tentacles. Gonophores fixed sporosacs or free medusae developing in a single whorl on gonozooids. Cnidome of polyp includes desmonemes. Medusa with lobed bell margin, with marginal nematocyst ring from which usually arise several centripetal nematocyst bands or exumbrellar rows of réfringent spots. Four radial canals and hollow circular canal, short mesenteries. Four marginal tentacles with bases embedded in umbrellar furrows, no ocelli. Manubrium with short, perradial mouth arms ending in nematocyst clusters; gonads interradial on manubrium. Remarks: For descriptions of fully developed Thecocodium medusa see Jarms (1987) and Kubota (1993). The ring circular canal of the medusa of Thecocodium quadratimi is hollow (own unpublished observations on young medusae), unlike the Proboscidactylidae which have reportedly a solid circular strand. Key to the Thecocodium species of the ERMS zone: la dactylozooids with up to 5 capitate tentacles 2 lb dactylozooids with more than 7 capitate tentacles, gonophores liberated as medusae T penicillatum 2a gonophores fixed sporosacs, shallow water form T brieni 2b gonophores a medusa or medusoid, deep water form Thecocodium spec. Thecocodium brieni Bouillon, 1967 Fig. 15 Ptilocodium repens. -Teissier, 1965: 13. [not Ptilocodium repens Coward, 1909] Thecocodium brieni Bouillon, 1967: 1106, figs 1-11. - Brinckmann-Voss, 1970: 85, figs 100- 106. - Edwards & Harvey, 1983: 41, fig. 2. - Boero & Fresi, 1986: 139. - Bouillon et al, 2004: 76, fig. 44B. - Calder, 1998: 1849. Material examined - IRSNB IG 27.838, holotype and paratype colonies; type colony collected 1961, Naples, on barnacle (sponge covered?) growing on piece of rock, second small colony on algae; paratype colonies on sponge covered rock, fertile. Diagnosis: Thecocodium with dactylozooids having 4-5 tentacles, gastrozooids without pedicel, gonophores sessile sporosacs, males styloid, females with radial canals reduced to pouches. Description (Bouillon, 1967; Brinckmann-Voss, 1970; own observations): Colonies small, hydranths stolonal, issuing from creeping, mesh-like hydrorhiza for- med by round stolons, these covered by very thin perisarc. Polyps polymorphic, with gastro-gonozooids and dactylozooids. Gastrozooids without tentacles, bottle-shaped, sessile, with ovoid base tapering distally into neck-like hypostome, base without distinct pedicel, polyp thus sessile, gastrodermis of hypostome forming four thickened longitudinal ridges visible from outside, apical epidermis studded with nematocysts, lower part of hydranth sac-like. Dactylozooids much more frequent than gastro-gonozooids, contractile, composed of slightly tapering or isodiametric stem ending in a whorl of capitate tentacles, usually four tentacles, occasionally five, rarely three, trunks of tentacles radiate from end of stem, held either horizontally or directed upwards depending on contraction, base of stem with shallow perisarc collar, gastrodermis of stem and tentacles chordoid. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 473 Fig. 15 Thecocodium brieni Bouillon, 1967; after preserved type material; part of colony with three dactylozooids in different states of contraction and two gastrozooids, the left one bearing a gonophore at its side, scale bar approximately 0.2 mm. Gonophores develop singly near base of gastrozooids, the latter can get reduced in size while gonophores mature. Gonophores are sessile sporosacs, male ones a simple bulbous evagination of body wall (styloid type), female gonophores with vestiges of radial canals forming four pouches surrounding the spadix, occasionally female gono- phores also containing spermatogonia, several oogonia develop, but usually only two mature to eggs, eggs develop to planula in situ, animal thus larviparous. Nematocysts: microbasic euryteles, discharged shaft about as long as capsule; desmonemes with relatively long capsules. Colours: gastro-gonozooids pink with opaque white hypostomes, dactylozooids translucent colourless with white capitula, stolons pink (Edwards & Harvey, 1983). Dimensions (Bouillon, 1967; Brinckmann-Voss, 1970; own observations): Colony diameter usually smaller than 10 mm; gastrozooids 0.3-1.3 mm high, diameter up to 0.4 mm; dactylozooids up to 0.9 mm, diameter 0.05-0.1 mm (contractile), capitula diameter up to 65 /jm, tentacle length up to 0.2 mm (contractile); sporosacs ca. 0.2 mm or more. Microbasic euryteles 15.4x5.6 ^um, desmonemes 1 1.2x4.2 ^m. Other Data: The histology of the polyps and gonophores has been described by Bouillon (1967) and Brinckmann-Voss (1970). Bouillon (1967) also depicts the nematocysts. Biology: Occurs on stones, rocks, oyster shells, calcareous tubes of polychaete worms, algae, and tunics of ascidians, depth range 2-200 m (Brinckmann-Voss. 1970; 474 P. SCHUCHERT Calder, 1998). In the Mediterranean, fertile animals have been observed from April to November (Brinckmann-Voss, 1970; Boero & Fresi, 1986). The feeding behaviour was described in detail by Brinckmann-Voss (1970) and Edwards & Harvey (1983). Prey is caught by the dactylozooids, the gastrozooids then elongate towards them and swallow the prey. Distribution: Mediterranean (Naples: Bouillon, 1967; Brinckmann-Voss, 1970), northern Brittany (Bouillon, 1967); Western Scotland (Oban: Edwards & Harvey, 1983); Bermuda (Calder, 1998). Type locality: Naples (designation of type specimen by J. Bouillon). Thecocodium penicillatum Jarms, 1987 Fig. 16 Thecocodium penicillatum Jarms, 1987: 62, figs 8.5-8.6. Type material: The type material is likely lost (pers. com. P. Stiewe, Zoological Museum of the University of Hamburg). Diagnosis: Thecocodium with dactylozooids having 7-11 tentacles, gastro- zooids with long pedicel. Gonophores free medusae, stomach with large vacuolated cells, exumbrella with meridional tracks of refractive spots, four tentacles. Description (after Jarms, 1987; from in vitro culture): Colonies with stolonal hydranths issuing from creeping, mesh-like hydrorhiza formed by tubular stolons, co- vered by thin perisarc. Polyps polymorphic, with gastro-gonozooids and dactylozooids, without perisarc envelope. Gastrozooids without tentacles, club-shaped, with stalk (pedicel), pedicel thin and cylindrical, distal part bottle-shaped, tapering distally into neck-like hypostome, up to five gonophores developing in a single whorl at junction of pedicel and hydranth- body. Dactylozooids much more frequent than gastro-gonozooids, contractile, without mouth, composed of thin, slightly tapering or isodiametric stem ending in a radiating tuft of capitate tentacles, 7 to 11 (average 10) capitate tentacles with thin stalk, gastrodermis of stem and tentacles chordoid. Newly released medusa bell-shaped, distinctly higher than wide, umbrella rather thick, without gastric peduncle, margin lobed through perradial embayments, exumbrella with perradial and interradial meridional furrows, interradial ones shallow, on exumbrella four interradial lines of refractive spots, some nematocysts scattered on exumbrella. Velum broad. Manubrium conical, gastrodermis composed of remarkably large (vacuolated?) cells, mouth small and surrounded by many nematocysts, without any visible primor- dial gonads. Four radial canals, widening where entering manubrium and thus forming short mesenteries, narrow circular canal. Four short perradial tentacle stumps, slightly swollen near origin. Adult medusa unknown. Nematocysts: microbasic euryteles on polyp and medusae; desmonemes; holo- trichous macrobasic euryteles occurring only on dactylozooids. Dimensions (Jarms, 1987): Gastrozooids including pedicels 2-3 mm high, pedicels 1.6-2 mm; dactylozooids up to 1.6 mm high, capitate tentacles 0.1-0.3 mm EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 475 A B Fig. 16 Thecocodium penicülatum Jarms, 1987; from Jarms (1987). (A) Gastro-gonozooid, scale bar 0.25 mm. (B) Dactylozooid, same scale as A. (C) Newly released medusa in side view, scale bar 0.1 mm. (D) Same as C but oral view. long; gonophores before liberation 0.45 mm in diameter; newly liberated medusa 0.8- 0.95 mm high and 0.64 mm wide. Biology: Deep water hydroid found on slag (clinker?). Distribution: Only known from type locality, Canary Islands, 25.4006°N 1 6.2342° W, 810 m depth. Remarks: See Thecocodium spec, below. Thecocodium spec. Fig. 17 Material examined: MHNG INVE 62870; Norway, Jan Mayen, Trollveggen vent field, 71.298°N 05.773°W, 574 m, 25.07.2008; coll. by 2008 G.O. Sars Cruise of the University of Bergen, growing on Sertularella tenella, gonozooids and dactylozooids present, stolons covered by orange-red mineral precipitate (iron oxide?); 16S sequence accession number FN422378. 476 P- SCHUCHERT Description: Gastro-gonozooids and dactylozooids identical to T. brieni (comp. Fig. 15), but gonophores are medusoids or medusae with a manubrium composed of large vacuolated cells, no tentacles or bulbs, stolons covered by thicker perisarc, deep water occurrence. Nematocysts: elongated desmonemes in dactylozooids; large almond-shaped microbasic euryteles around mouth of gastrozooids and in capitula of dactylozooids, discharged shaft about as long as capsule and only faintly swollen; smaller ovoid microbasic heteroneme on gonophores. Dimensions: Desmonemes (18.5-19)x(5.5-6.5)/mi, large microbasic eurytele (21-23)x(6.5-8.5)^m, small microbasic heteroneme (8.5-9)x(5.5)^m. Distribution: Deep water (>500m), Jan Mayen, North Atlantic. Remarks: During the "GO Sars 2008 Expedition" of the Centre for Geobiology of the University of Bergen, some hydroids were collected in the vicinity of a deep- water hydrothermal vent area near the island of Jan Mayen. Among these hydroids was also a Thecocodium colony growing on Sertularella tenella. The colony consists of convoluted stolons bearing gono-gastrozooids and dactylozooids. The polyps of this Thecocodium colony resembles in almost all aspects Thecocodium brieni (Fig. 15). The dactylozooids have 4-5 capitate tentacles and the gono-gastrozooids are sessile with a whorl of gonophores in the lower region. However, the gonophores seem to be medusoid and not sessile sporosacs (Fig. 17). The most advanced gonophores (0.35 mm long, 0.2 mm wide) have a large spadix with vacuolated gastrodermal cells, but lack any traces of gametes. At the distal end of the manubrium/spadix there is a small zone with more dense tissue. There were no tentacles or bulbs visible. They thus resemble the medusoids of T penicillatum described by Jarms (1987) (see Fig. 16C). The gonophores were covered by ovoid heteronemes which are smaller and rounder than the normal microbasic euryteles of the polyp. This type of nematocyst capsule is unknown in T. brieni (Bouillon, 1967). All these differences, together with its unusual habitat, suggest that this colony likely belongs to an undescribed species. However, it could also be a form of T. penicillatum, a deep-water Thecocodium species that produces medusoids or medusae. According to our current knowledge, T. penicillatum has dactylozooids with 7-10 tentacles and stalked gastro-gonozooids. Unfortunately, this species is only known from a single, in vitro culture. As we have no idea of its intraspecific variability, especially in material collected from nature, it is not possible to relate the present sample to this species. As the only available sample did not permit to obtain sufficient information on the nature of the mature gonophores. the species is here not named in order to avoid the creation of further, ambiguous hydroid species. The 16S sequence will perhaps allow it to be identified in the future. Genus Tregoubovia Picard, 1958 Type species: Tregoubovia atentaculata Picard, 1958 by monotypy. Diagnosis: Medusa small, spherical, without tentacles, bell margin with nema- tocyst ring from which originate centripetal, flat, nematocyst containing bands running meridionally on exumbrella. Manubrium large, cruciform base, the four perradial EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 477 Fig. 17 Thecocodium spec. MHNG INVE 62870, preserved material from the North Atlantic. (A) Optical section of the most advanced gonophore; the gastrodermis of the manubrium is composed of very large, apparently vacuolated cells (arrow), only at the distal end there is some more dense tissue; scale bar 0.2 mm. (B) Desmoneme. (C) Large microbasic eurytele. (D) Small microbasic heteroneme. The scale bar for B-D equals to 10 urn. corners of mouth margin drawn into distinct oral tentacles with terminal nematocyst clusters. Gonads interradial on manubrium wall. Without desmonemes. Remarks: This is currently a monotypic genus. The polyp stage remains unknown. Tregoubovia atentaculata Picard, 1958 Fig. 18 Tregoubovia atentaculata Picard, 1958: 185, fig. - Goy, 1973: 979. - Bouillon et al, 2004: 77. fig. 44C. Material examined: MHNG INVE 39476, supposed holotype, collected 1955, ex. Picard collection. Diagnosis: See genus diagnosis. Description: Medusa umbrella spherical to ovoid, mesogloea moderately thick, bell margin slightly lobed, without gastric peduncle, exumbrella with scattered nema- tocysts, velum rather narrow. Manubrium large and voluminous, reaching to level of velum, base square to cross-shaped, the four perradial corners of mouth margin drawn into distinct oral arms, oral arms with chordoid gastrodermis, ends swollen and studded with nematocysts. Gonads four large, oblong interradial pads on manubrium, well separated perradially. Four relatively thick radial canals, widening where entering manubrium and thus forming short mesenteries. Diameter of ring canal and bell opening relatively narrow. Without free tentacles. Along bell margin a ring of thickened tissue with nema- tocysts from which originate up to 16 centripetal bands (4 perradial, 4 interradial, 8 adradial) running meridionally on the surface of the exumbrella towards aboral pole. 478 P. SCHUCHERT Fig. 18 Tregoubovia atentaculata Picard, 1958; redrawn from (Picard, 1958), male medusa, bell size about 1.5 mm. reaching maximally to mid umbrella; these tentacle-like structures contain nematocysts and are like flat bands pressed into the surface of the exumbrella so that they are flush with the exumbrellar epithelium, but they are not covered by the exumbrellar epi- thelium. Nematocysts: microbasic euryteles, desmonemes absent. Polyp stage unknown. Dimensions: Umbrella 1.5-3.2 mm high when mature, euryteles 8x4^m. Biology: Deep water species. Distribution: Only known from the type locality: Mediterranean, Villefranche- sur-Mer. Remarks: This is a very rare species; so far only two or three specimens have been reported in the literature. Tregoubovia atentaculata lacks tentacles but has flat centripetal nematocyst bands originating from a nematocyst band at the bell margin. Similar structures are present in Thecocodium quadratum (Werner, 1927) and Hansiella fragilis Bouillon, 1980 (Bouillon et al., 1997), which justifies its inclusion in the Ptilocodidae. Picard (1958) regarded these nematocyst bands as being derived from tentacles that developed inside the mesogloea. He claims to have seen that they are composed of a gastrodermal core enveloped by epidermal tissue with nematocysts. The presence of a gastrodermal core could not be verified with the available material. It was evident, however, that these bands are not covered by the normal exumbrellar epithelium, thus they are not really tentacles that have grown into the mesogloea. They more resemble flattened tentacles pressed into and fused to the surface of the exum- EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 479 brella. More details can only be obtained through new material and a histological examination using electron microscopic methods. Family Eucodoniidae Schuchert, 1996 Diagnosis: Anthomedusae without pointed apical projection, exumbrella without nematocyst tracks or clusters, with gastric peduncle; manubrium tubular; mouth quadrangular, with four inconspicuous nematocyst-clusters, with medusa budding on manubrium wall; gonads encircle manubrium without radial interruptions; four radial canals and circular canal present; four small perradial marginal bulbs and four tentacles; tentacles with a single terminal swelling; no ocelli; cnidome comprises microbasic eury teles and desmonemes. Remarks: This is a monotypic family, the polyps are unknown. See also the re- marks under Eucodonium brownei. Genus Eucodonium Hartlaub, 1907 Type species: Eucodonium brownei Hartlaub, 1907 by monotypy. Diagnosis: See family diagnosis. Eucodonium brownei Hartlaub, 1907 Fig. 19A-B Dipurena sp. Browne 1896: 473, pi. 16 fig. 2. Eucodonium brownei Hartlaub 1907: 71, fig. 67. - Neppi & Stiasny, 1913: 14, pi. 1 fig. 6. - Kramp, 1937: 28, fig. 8c. - Russell, 1953: 93, fig. 40. - Picard, 1955: 95. - Vannucci, 1957: 43, figs 2-3. - Kramp, 1959: 91, fig. 44. - Kramp, 1961: 36. - Brinckmann-Voss, 1970: 19, figs 16-19, pi. 2 fig. 4. - Goy, 1973: 972. - Petersen, 1990: 217. - Schuchert, 1996: 89, fig. 53a-b. - Bouillon et al., 2006: 56, fig. 33C. Type material: Not located (not found in BMHN). Material examined: MNHN1618; France, Villefranche-sur-Mer; plankton 50-100 m; 3 medusae collected October 1963 and July 1964; material of Goy (1973). - See also Schuchert (1996). Diagnosis: See family diagnosis. Description (Russell, 1953; Brinckmann-Voss, 1970; own observations): Medusa umbrella nearly hemispherical, sometimes slightly higher than wide, lateral walls thin, apex with thickened jelly, manubrium attached to a broad, well developed gastric peduncle, velum broad. Manubrium cylindrical, 1/2 as long as bell cavity, tube-like, mouth quadran- gular with four inconspicuous perradial lips, each containing a group of nematocysts. Immature animals produce secondary medusae via medusae buds on middle region of stomach. Mature animals without buds, gonads encircle the manubrium without radial interruptions. Four very narrow radial canals and circular canal present. Four small marginal bulbs containing blackish pigment granules, without apparent ocelli. Four equally developed tentacles with a conspicuous terminal swelling, swelling spherical to ovoid; tentacle gastrodermis chordoid; terminal swellings with enlarged gastrodermis and epidermis with fibrous structure. Nematocysts not only 480 P. SCHUCHERT Fig. 19 Eucodonium brownei Hartlaub, 1907; preserved material from New Zealand, modified after Schuchert (1996). (A) Medusa with medusa-buds on manubrium, scale bar 0.3 mm. (B) Undischarged nematocysts: microbasic eurytele from tentacles, heteroneme from manubrium, desmoneme, scale bar 10 urn. present in terminal swelling but also along the tentacles, nematocysts in terminal swelling not so dense. Nematocysts: microbasic euryteles on tentacles, heteronemes (microbasic euryteles?) from lips, desmonemes on tentacles. Colour of marginal tentacles bulbs blackish, stomach blackish brown, terminal knob of marginal tentacles brownish. Dimensions: Bell diameter and height usually 0.8-1 mm. For nematocyst dimensions see Schuchert (1996). Biology: The medusa can be found close to the water surface. In the Atlantic it is very rare, but Brinckmann-Voss (1970, 1987) found it in relatively high numbers near Naples from September to October. Also Daly et al. (2003) found it off Tunis in elevated numbers. Likewise, it was found in comparatively high numbers in northern EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 48 1 New Zealand during the summer months. Brinckmann-Voss (1970) observed that lowering the temperature from 20 to 13°C stopped the medusa budding irrespective of the season. The gonad maturation likely also depends on falling temperatures. Distribution: English Channel (Browne, 1896; Hartlaub, 1907; Franc, 1951; Russell, 1957), Faroe-Shetland Channel (Fraser, 1974), Denmark (Kramp, 1927), Mediterranean (Neppi & Stiasny, 1913; Picard, 1955 ; Brinkmann- Voss, 1970, 1987; Goy, 1973; Daly et al, 2003; Medel & López-Gonzalez, 1996), Brazil (Vannucci, 1957), SW Atlantic (Genzano et al, 2008), New Zealand (Schuchert, 1996; Bouillon & Barnett, 1999). Type locality: Plymouth, England. Remarks: With its swollen tentacle ends combined with the medusa buds and the gastric peduncle this is quite a characteristic medusa (Fig. 19A). The tentacle swellings in European animals seem to be larger than in those observed in New Zealand. The polyp stage of Eucodonium remains unknown, which makes it difficult to relate it to other species of Filifera. Eucodonium was initially placed among the Capitata, but the cnidome makes it obviously related to the Filifera (Picard, 1955; Schuchert, 1996). Picard (1955) thinks that is could be close to Podocorynoides minima (Trinci, 1903) (see Schuchert, 2007 for redescription). Indeed, both medusae are quite similar, except for the tentacle tips of Eucodonium and the oral tentacles of Podocorynoides, and Picard's suggestion could be correct. Family Russelliidae Kramp, 1957 Diagnosis: Medusa umbrella with apical projection. Manubrium on gastric peduncle, four unbranched oral filiform tentacles attached above mouth margin, mouth simple, lips indistinct. Marginal tentacles in eight groups, four perradial and four inter- radial, each group with one large tentacle flanked by two small tentacles; large tentacles hollow, without basal swellings, basal part adnate to umbrella and sunk into deep furrows of umbrella margin, adaxial ocellus at base of free portion of each tentacle. Gonads in eight large adradial pads on manubrium wall. Remarks: The polyp stages of the Russellidae are unknown. The family com- prises only one genus. Genus Russellia Kramp, 1957 Type species: Russellia mirabilis Kramp, 1957 by original designation. Diagnosis: As for family. Remarks: The genus is currently monotypic. Russellia mirabilis Kramp, 1957 Fig. 20A-D Russellia mirabilis Kramp, 1957: 24, pi. 4 figs 1-6, text-fig. 4. - Kramp, 1959: 30. 129, fig. 142. - Kramp, 1968: 58, fig. 154. - Pages et ai, 1999: 2431, fig. 1. - Bouillon et. al, 2004: 78, fig. 45A. Type material: BMNH (not seen). Diagnosis: See family diagnosis. 482 P. SCHUCHERT Fig. 20 Russellia mirabilis Kramp, 1957; redrawn from Kramp (1957). (A) Lateral view of medusa, bell height 13 mm. (B) Higher magnification of tentacle base in lateral view. (C) Oral view of a sector of the bell margin showing radial canal and two tentacle bases flanked by a pair of dwarf- tentacles. (D) Perradial corner of the mouth with an oral tentacle. Description (Kramp, 1957): Umbrella distinctly higher than wide, with broad, dome-shaped apical projection measuring about 1/3 of total height, lateral walls moderately thick, bell-margin hanging in lobes somewhat below the level of the ring- canal, stomach mounted on a broad, conical peduncle, extending partly into the cavity of the stomach, peduncle about as high as stomach, which spans somewhat less than one fourth the height of the bell cavity. Velum narrow. Base of stomach cross-shaped; stomach itself deeply folded inward in the inter- radii, and sometimes also in the perradii. In the preserved condition stomach slightly wider than long, the interior surface of the stomach densely wrinkled transversally and along each interradius a narrow, prominent ridge thrown into regular transverse folds, indicating that in living specimens the stomach may be extended to a greater length. Mouth quadrangular with very short and simple perradial lips, mouth rim smooth and entire, slightly thickened, without concentration of nematocysts in the rim itself, but with some scattered in the wall above it. At a short distance above the perradial corners of the mouth four oral tentacles, finger-shaped, tapering towards their distal end, without terminal cluster of nematocysts, but numerous nematocysts evenly scattered EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 483 throughout the entire length of the tentacle, not more concentrated towards the tip than in the basal part. Gonads in eight adradial pads, occupying almost the entire length of the sto- mach, separated in the perradii and interradii by narrow lines, surface smooth, without transverse folds. Tentacles in eight clusters, each consisting of one large and two small tentacles. Eight large tentacles of equal size, four perradial and four interradial, basal part (root) of each tentacle directed upward and outward and deeply sunken into a narrow cleft between two prominent lobes of the umbrella-margin; in the interradial tentacles ascending root directly adnate to the gelatinous tissue of the exumbrella, in the per- radial tentacles root fused to the terminal part of the corresponding radial canal by a triangular connection (Fig. 20B). Distal to the adnate root, tentacles becoming free and very thin; with one adaxial, red ocellus where tentacles become free; terminal tentacle- structure unknown. At the base of each of the eight large tentacle-roots a pair of much smaller tentacles, structure similar to larger ones (Fig. 20B-C). Four radial canals, their ascending part on the peduncle fairly wide, in transver- se section like an equilateral triangle; descending portion along subumbrella narrow and flat, with smooth edges. Ring-canal narrow. Polyp stage and cnidome unknown. Dimensions: Bell height up to 15 mm, diameter up to 9 mm, manubrium about 3 mm high. Biology: The medusa Russelia mirabilis lives mainly in the 0-300 m depth range (Pages et ai, 1999). Outside its main distribution (Antarctica) it occurs usually in deeper waters (200-1000 m: Kramp, 1959; Pages et al, 1999). Distribution: Antarctica; West Indies; Mediterranean (Kramp, 1959; Navas- Pereira & Vannucci, 1990; Pages et al, 1999). Type locality: 66.475°S 71.27083°W, Antarctica, West of Graham Land, 0-165 m. Remarks: The disjunct and rather surprising distribution pattern of this medusa is difficult to explain. Pages et al. (1999) suspect that transport in the ballast water of a cargo ship is the most plausible explanation for its occurrence in the Mediterranean, but oceanic currents could have brought it to the Caribbean. Family Niobiidae Petersen, 1979 Diagnosis: Anthomedusae with two opposite simple and two opposite bifur- cated radial canals, so that six canals reach the circular canal; gonads interradial on manubrium; marginal tentacular bulbs developing into secundary medusae; no ocelli, no gastric peduncle, no mesenteries. Remarks: This family currently comprises only one genus. The polyps of the Niobiidae are unknown and thus the systematic position of the family remains unsolved. Genus Niobio. Mayer, 1900 Type species: Niobia dendrotentaculata Mayer, 1900 by monotypy. 484 P. SCHUCHERT Diagnosis: As for family. Remarks: This genus is currently monotypic. Niobia dendrotentaculata Mayer, 1900 Fig 21A-E Niobia dendrotentaculata Mayer, 1900: 36, pi. 42 figs 141-144, pi. 43 fig. 144. - Mayer, 1910: 187, pi. 19 figs 1-5. - Brinckmann, 1959: 334, figs 1-2. - Kramp, 1959: 115, fig. 104. - Kramp, 1961: 110, fig. 103. - Kramp, 1968: 41, fig. 103. - Bouillon et al, 2004: 66, fig. 40a. Type material: Location not investigated. Diagnosis: As for family. Description (Mayer, 1910; Brinckman, 1959): Medusa umbrella hemispherical or slightly flatter than a hemisphere, no apical projection, no gastric peduncle, jelly uni- formly thin, relatively rigid. Manubrium of mature medusa tubular, about as long as height of bell, with four simple but well developed lips, mouth cruciform, gonads in four interradial patches in upper part of manubrium, gonads usually - but not always - mature only after medusa budding from bulbs has stopped. Four radial canals arise from the four corners of the stomach, but two opposite canals bifurcate closely to manubrium so that six radial canals reach the circular canal at the margin, each pair of radial canals about 60° apart. Up to twelve marginal tentacles or bulbs present, one for each radial canal and one intermediate between each successive pair of radial canals. The tentacles develop successively and they are arranged in a bilaterally symmetrical manner in accordance with age. The oldest and the youngest tentacles are opposite, situated at the ends of the two simple radial canals, the remaining 10 tentacles are arranged in a bilaterally symmetrical manner in accordance with their various ages, the axis being in the diameter of the unbranched radial-canals and the oldest and youngest tentacles (Fig. 2 IE). Tentacle bulbs and tentacles transform successively into small medusae which detach from the parent (Fig. 21A-B). Colours: gastrodermis of manubrium, bulbs, circular canal brownish-yellow, other parts transparent. Newly-released medusa with 3-5 tentacles, radial canals and unevenly developed marginal tentacles render the medusa bilaterally symmetric (Fig. 21C). Nematocysts: microbasic euryteles and microbasic mastigophores. Dimensions: Diameter of mature medusa 2.4-4 mm, height only about 0.6 times the width. Newly released medusa 1.2 mm wide. Microbasic euryteles 7x2. 5^m, microbasic mastigophores 5x2.5/mi (Brinckmann, 1959). Biology: The medusa occurs close to the surface. It thrives well in culture (Mayer, 1910). Distribution: Rare, circumglobal in warm and temperate waters, recorded from Florida (Mayer, 1900); New England (Bigelow, 1915); Vietnam (Dawydoff, 1936); Mediterranean (Brinckmann, 1959); India (Nair, 1951); Argentina (Goy, 1979); Papua New Guinea (Bouillon, 1980). Type locality: Tortugas, Florida, USA. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 485 Fig. 21. Niobia dendrotentaculata Mayer, 1900; modified after Mayer (1910). (A) Side view of medusa showing the development of secondary medusae from the tentacle bases. (B) Oral view of a medusa, note the characteristic six radial canals and the bulbs that develop into small me- dusae. (C) Newly released medusa, note bilateral symmetry (mirror symmetry). (D) Manubrium of mature medusa with eggs. (E) Symmetry and sequence of tentacle development of medusa shown in section A, 1 denotes the most advanced, 7 the least developed tentacle. 486 p- schuchert Family Protiaridae Haeckel, 1879 Diagnosis (Bouillon et al., 2006): Hydroid colonial, arising from creeping, tubular stolons; hydranths issued from short hydrocaulus; hydrorhiza and hydrocaulus covered by pensare, forming a hydrotheca-like tube; hydranth with one whorl of filiform tentacles, large nematocysts alternating with tentacles. Medusa with four fully developed marginal tentacles arising from large, hollow tentacular bulbs; four simple radial canals and a circular canal, mouth with four simple lips; gonads interradial, with smooth surface; with or without mesenteries; without rudimentary bulbs; margin with or without cirri-like tentacles; exceptionally with ocelli. Remarks: The medusae of the Protiaridae and the Pandeaidae (see Schuchert, 2007) appear very similar and the differences in the diagnoses are minimal. Both families have nevertheless rather dissimilar polyps. Protiaridae polyps resemble those of Tricky dra (see p. 495) and the cnidome comprises merotrichous isorhizas, capsules otherwise only found in some leptomedusae (i.e., Eirenidae, Eucheilotidae, Haleciidae, Lovenellidae and Tiaropsidae; Bouillon, 1985; Bouillon et al., 1988). The development of the medusae and the gonozooids are insufficiently known in this family. Dr. Brinckmann-Voss, however, has found and reared the polyps of Halitiara formosa (pers. com., unpublished) and she observed that the medusae are produced in gono- thecae. The family could thus be more closely related to the order Leptothecata. Protiaridae genera of the ERMS zone: la with cirri (= small free-hanging tentacles on bell margin, in addition to normal tentacles) Halitiara 1 b without cirri 2 2a without mesenteries, marginal tentacles without abaxial spurs Protiara 2b with short mesenteries, marginal tentacles with abaxial spurs Paratiara Genus Halitiara Fewkes, 1882a Type species: Halitiara formosa Fewkes, 1882a by monotypy. Diagnosis: Medusa with four straight radial canals; four perradial marginal tentacles; with marginal cirri between long tentacles; mouth simple, cruciform; with or without mesenteries; gonads interradial, smooth, sometimes extending along mesen- teries; without ocelli; cnidome where known including merotrichous isorhizae. Hydroid diagnosis as for family. Remarks: Gershwin & Zeidler (2003) provide a table with the characteristics of all known Halitiara species. Key to the Halitiara medusae of the ERMS zone: la with apical process H. formosa lb without apical process H. inflexa Halitiara formosa Fewkes, 1882a Figs 22A-B, 23A-B ? Dissonema gaussi Vanhöffen. 1912: 361, pi. 24 fig. 2. - Kramp, 1965: 27, possible synonym. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 487 Halitiara formosa Fewkes, 1882a: 267, pl. 4 fig. 2. - Kramp, 1959: 115, fig. 103. - Uchida, 1927: 203. - Menon, 1932; 7, pl. 1 fig. 4. - Kramp, 1961: 102. - Kramp, 1968: 40, fig. 102. - Brinckmann-Voss, 1970: pl. 11 fig. 1. - Goy, 1973: 983, flg. 7. - Bouillon, 1980; 332. - Goy et al., 1991 : 1 10, fig. 26. - Schuchert, 1996: 76. - Gershwin & Zeidler, 2003: table 3. - Bouillon et al, 2004: 75, fig. 43E. Protiara formosa. - Mayer, 1910: 107, pl. 6 figs 4-6, pl. 13 figs 1-2. in part Halitiara formosa. - Kramp, 1965: 27. [some = Leuckartiara simplex] Type material: Location not investigated. Material examined: ZMUC, without registration number; USA, Florida, St Andrews Bay; collected August 1959, 2 medusae, det. Kramp, leg. M. Hopkins. - MHNG INVE 63285 ; Mediterranean, France, Villefranche-sur-Mer, surface plankton; 10 April 1972; one mature, contracted medusa, one medusa without manubrium; specimens collected and identified by A. Brinckmann-Voss. Diagnosis: Medusa up to 3 mm high, with solid apical projection, manubrium about half as long as bell cavity; mouth opening cruciform; with indistinct mesenteries. Four long and 24-35 short, tightly coiled, cirrus-like tentacles; no ocelli. Description: Polyp stage similar to H. inflexa and Trichydra pudica (A. Brinckmann-Voss, pers. coram.). Gonozooids are reduced hydranths without tentacles (=blastostyles) enclosed in a filmy, soft perisarc tube like a gonotheca, budding one or two medusae at a time. Newly liberated medusa with 3-4 perradial tentacles and 1-3 cirri between each pair of successive perradial tentacles. Mature medusa with bell-shaped umbrella, slightly higher than wide, with distinct, solid apical projection of somewhat variable shape; without gastric peduncle; velum narrow. Manubrium pyriform to cruciform in section, about half as long as the depth of the bell-cavity, with simple, cruciform mouth. Gonads on interradial sides of the manubrium, large, covering most of manubrium, separated perradially, ova large and conspicuous. Four straight, narrow radial canals and a slender circular canal, radial canals with elongated opening where joining manubrium, thus forming mesenteries, but these inconspicuous, shorter than half the manubrium height, may be absent in specimens with less developed gonads. Four long perradial tentacles, with long, hollow, tapering basal bulbs, without exumbrellar spurs, without ocelli; in addition to long tentacles 24-35 short, solid, cirri- like tentacles, independent of long tentacles, often tightly coiled, gastrodermal cells chordoid. Colours: gastrodermis of the manubrium and tentacle bulbs in the female green, but in the males light-brown (Mayer, 1910). Dimensions: Bell height of mature medusa 1-3 mm. Nematocysts of preserved Mediterranean specimen: large ovoid capsules, one long side more straight than other, {\6-\l )x{9-\0)nm\ smaller elongate almond-shaped capsules (8-8.5)x(2.5-3) i um. The capsule types could not be identified, but the undischarged capsules resemble those of H. inflexa depicted in Bouillon et al. (1988). Biology: A coastal medusa occurring in shallow depths. The medusa is rare in the Mediterranean, but Mayer (1910) reported it as being very abundant at the 488 P. SCHUCHERT Fig. 22 Halitiara formosa Fewkes, 1882. (A) Lateral view of mature, female medusa, manubrium somewhat contracted, composite image drawn after several preserved specimens from Florida, height about 1.2 mm (B) Oral view of medusa, from Mayer (1910). Tortugas, Florida. In the Mediterranean, the medusae were observed from August to November (Goy, 1973; Brinckmann-Voss, 1987; Goy et al., 1991). Polyps budding medusae were observed in September (Brinckmann-Voss, 1987). They occur on oyster shells (A. Brinckmann-Voss, pers. com.). Distribution: Tortugas, Bahamas (Mayer, 1910); Mediterranean (Goy, 1973; Brinckmann-Voss, 1987; Goy et al., 1991); Argentina (Genzano et al. 2008); tropical parts of the Indo-Pacific Ocean (Menon, 1932; Kramp, 1965; Bouillon, 1980); NW Pacific (Uchida, 1927; Kramp, 1965); records from New Zealand are likely erroneous (Schuchert, 1996). Type locality: Tortugas, Florida, USA. Remarks: Descriptions or illustrations of the polyp stage of H. formosa have not yet been published. Dr Anita Brinckmann-Voss found the polyp repeatedly near Naples and she was able to rear the medusa released from these colonies. Dr Brinckmann-Voss was so kind to send me her drawings and notes and they were here used for the description and the figure of the gonotheca (Fig. 23A-B). EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 Fig. 23 (A-B) HaUtiara formosa Fewkes, 1882; lateral view of gonothecae, living material from the Mediterranean, after unpublished sketches of Dr. A. Brinckmann-Voss. Note that the images are not represented at the same scale as C. (A) Gonotheca with a developing medusa (oval mass). (B) Gonotheca after release of medusa, blastostyle drawn stippled. (C) Halitiara inflexa Bouillon, 1980; polyp stage, modified from Bouillon (1985). According to Kramp (1959), the medusa of this species has no mesenteries. Some authors (e. g. Goy, 1973) nevertheless described it as having mesenteries (but in the accompanying figure they are not apparent). The specimens examined for this study - coming from the same biogeographic region as the type material - clearly had mesenteries (Fig. 22A), although they were rather short and inconspicuous. The length is less than half of the manubrium height. The original figure given by Fewkes (1882a) also suggests slight mesenteries, but it is possible that their full development is corre- lated with the age or size of the animal and also the size of the manubrium and gonads. Some specimens from the NE Pacific identified by Kramp (1965) as H. formosa (in ZMUC) were re-examined for this study and they turned out to be Leuckartiara simplex Bouillon, 1980. The related species Halitiara inflexa Bouillon, 1980 has been reported from the eastern Mediterranean by Goy et al. (1991) (see also Bouillon et al, 2004: 76, fig. 43F- G). This species differs from H. formosa by the absence of an apical process. The other purported difference, namely the presence of mesenteries, is likely incorrect or unre- liable (see above). As also the apical process shows some variation, the Mediterranean records of H. inflexa could be just a variant of H. formosa. Halitiara inflexa is other- wise only known from Papua New Guinea and this population is likely distinct from H. formosa (see also Gershwin & Zeidler, 2003). 490 P- SCHUCHERT Halitiara inflexa Bouillon, 1980 Fig. 23C Halitiara inflexa Bouillon, 1980: 324, fig. 9. - Bouillon, 1985: 259, fig. 7. - Bouillon et al, 1988: 211, fig. 8. - Goy et al, 1991: 110, fig. 27. - Bouillon, 1995: 230, fig. 6. - Schuchert, 1996: 77, fig. 46a-b. - Bouillon et al, 2004: 43F-G. Type material: IRSN Bruxelles (not seen). Diagnosis: Medusa up to 3 mm high and 2.4 mm wide, bell conical without apical projection, manubrium about half as long as bell cavity; mouth opening cruci- form; with distinct mesenteries. Four long and 28-40 short, cirrus-like tentacles; no ocelli. Description (Bouillon, 1980; Bouillon, 1985; Schuchert 1996): Hydroid colonies arising from creeping, perisarc-covered stolons. Polyps with a very short caulus, a long, narrow cylindrical body and a short conical hypostome. Below hypo- stome one whorl of ten long, filiform tentacles with irregular clusters of nematocysts. Alternating with the tentacles are large nematocysts. Caulus and base of polyp body are covered by a perisarc cup into which the polyp can almost completely retract. Gonozooids unknown. Medusa umbrella bell-shaped, rather conical, jelly moderately thick, gradually thickening towards top to about two times the thickness of the lateral walls. Manubrium voluminous, quadrangular, length about 2/3 of bell cavity, joined to radial canals by mesenteries for 1/2 of their length. Mouth with four simple lips. Gonads large, bulging, filling interradial position completely, leaving only a small perradial band of stomach and the mouth region free. Four radial canals and circular canal, all narrow and with smooth margins. Four long perradial tentacles, with broad conical base then tapering, base not laterally compressed. Nematocysts evenly distributed on tentacles. Between each pair of long tentacles 3 to 10 short, cirri-like tentacles, without bulbs, often coiled, with chordoid gastrodermis, tips with nematocysts (haplonemes). Nematocysts: atrichous isorhizas of two size classes, merotrichous isorhizas, mastigophores. Dimensions: Bell height up to 3 mm, diameter up to 2.4 mm. For nematocyst dimensions see Bouillon (1980) and Bouillon et al. (1988). Polyps up to 1 mm, with a very short caulus (80 /urn), Distribution: Papua New Guinea, New Zealand, Mediterranean (Bouillon, 1980; Bouillon, 1995; Goy et al, 1991; Schuchert, 1996). Remarks: This species does not occur along the European coasts, but it has been found in the eastern Mediterranean (Lebanon; Goy et al., 1991), thus within the ERMS zone. For figures see Bouillon (1980, 1985) and Schuchert (1996). See also remarks under H. fonnosa. Genus Paratiara Kramp & Dumas, 1925 Type species: Paratiara digitalis Kramp & Dumas, 1925. Diagnosis: Protiaridae medusae without marginal cirri, four marginal tentacles with abaxial spurs, no ocelli, with smooth, interradial gonads, manubrium more or less twisted, four simple oral lips, with short mesenteries. Remarks: This is a monotypic genus. EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 491 Fig. 24 Paratiara digitalis Kramp & Dumas, 1925; from Kramp & Dumas (1925). (A) Medusa in side view, bell height about 1 cm. (B) Oral view of manubrium, note characteristic torsion of its perradial wings. (C) Horizontal section of manubrium of a female medusa. Paratiara digitalis Kramp & Dumas, 1925 Fig. 24A-C Paratiara digitalis Kramp & Dumas, 1925: 273, figs 18-20. - Kramp, 1926: 66, chart XL - Kramp, 1961: 1 14s. - Fraser, 1974: 12. Type material: Location not investigated. Material examined: ZMUC, DANA station 851 (NE of Bahamas), 22.3833°N 60.7667°W, 50 metres wire, 05.06.1920; one damaged medusa, material of Kramp (1959). - ZMUC, DANA station 891 (West-Indian Islands), 29.4667°N 69.4167°W, 50 metres wire, 24.07.1920; one much damaged medusa, material of Kramp (1959). Diagnosis: See genus diagnosis. Description: Medusa bell higher than wide, cylindrical shape with thin walls and rather flat top, roof of subumbrella with four interradial pockets. Manubrium flask-shaped, 2/3 of length of bell cavity, with short mesenteries (1/4 of manubrium height); four simple lips; manubrium cruciform in cross-section, sometimes twisted (Fig. 24B-C), the perradial edges all turned towards one side. Gonads smooth, interradial, completely covering the stomach walls except in the perradii. Four perradial tentacles with conical basal bulbs, each with a well developed, epidermal, abaxial spur; no ocelli. Four radial canals, smooth, straight, joining manubrium in upper fourth and forming mesenteries; ring canal thin. Polyp stage and nematocysts unknown. Dimensions: Bell height up to 10 mm, in tropical waters 4.5 mm high and 4 mm wide. 492 P- SCHUCHERT Biology: A rare medusa usually found close to the water surface, mostly caught during summer time. Distribution: Rockall (Fraser, 1974); between Shetland and Iceland (Kramp, 1926); North Cape (Kramp & Damas (1925); Sargasso Sea (Kramp, 1959). Type loca- lity: Vard0, North Cape, Norway, North Atlantic Ocean, near surface. Remarks: The presence of mesenteries is a feature that distinguishes this spe- cies and genus from Protiara tetranema. A re-examination of some specimens identi- fied by Kramp and also the figures in Kramp & Damas (1925, reproduced in Fig. 24) showed that they are rather short, spanning only about 1/4 of the manubrium height. The torsion of the manubrium is a unique feature of this species, but this seems not to be a constant feature as some examined medusae had a straight manubrium. Genus Protiara Haeckel, 1 879 Type species: Oceania tetranema Péron & Lesueur, 1810 by monotypy. Diagnosis: Protiaridae medusae without marginal cirri, four marginal tentacles, with or without ocelli, with four or eight smooth, vertical gonads in the adradial-inter- radial region, four simple oral lips, without mesenteries. Remarks: This genus is likely invalid as its type species is indeterminate. See the remarks below. Protiara tetranema (Péron & Lesueur, 1810) Fig. 25 zweyte karminrothe Berne Slabber, 1775: 64, pi. 14, Fig. 1. [nomenclature not binomial] Oceania Tetranema Péron & Lesueur, 1810: 347. in part Protiara berne Mayer, 1910: 106. Protiara tetranema. - Haeckel, 1879: 47. - Hartlaub, 1914: 250, Fig. 206. - Kramp, 1959: 1 14, diagnosis. - Kramp 1961: 1 14. - Bouillon et al, 2004:76, fig. 44A. Type material: Likely lost. Type locality: Coast of The Netherlands. Remarks: This species is based on the description of a medusa given in Slabber (1775). Slabber describes and depicts a microscopically small, carmine-red medusa of which he had collected two living specimens from the Dutch coast (Fig. 25). It is characterized by four tapering tentacles, a cylindrical bell, and a short manubrium with four perradial lips. The most conspicuous trait is the carmine-red colour of the manu- brium. Although Slabber's description and figure are quite accurate, it seems neverthe- less impossible to relate this obviously immature medusa to any species known today. Maybe Slabber had a young pandeid medusa (cf. also Vanhoffen (1891) who had similar thoughts), although the red manubrium remains problematic. Neoturris pileata can also have an intensively red manubrium and its youngest stages have four tentacles only, but it is rare in this region. A conspicuous red medusa of the region is Turritopsis polycirrha (see Schuchert, 2006; or Russell, 1953 as T. nutricula), but most young Turritopsis medusae are released with eight tentacles (although the newly liberated medusae of T. polycirrha has never been described). Slabber (1775) also describes another carmine-red medusa with 18 tentacles, which was almost certainly EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 493 Fig. 25 Protiara tetranema (Péron & Lesueur, 1810); original illustration of Slabber (1775) depicting the type specimen. Size given as "microscopic"; Slabber indicates: a = flat top of body [umbrel- la], b = inside of body [subumbrella], d = tubes which transport digested food [radial canals], c = stomach. T. polycirrha, and which he had collected only four days earlier. They were the size of sand grains, thus presumably not more than 2 mm. A similar size or more likely smaller must also be assumed for Slabber's second red medusa, now called Protiara tetranema. As Slabber's account did not use a binomial nomenclature, he named it his "second carmine-red Beroe" , Péron & Lesueur (1810) gave it the name Oceania tetra- nema. They also noted the presence of numerous wart-like protrusions along the inside of the ring-canal. These knobs were not mentioned in Slabber's description, but they can be seen in his figure (Fig. 25). Could they be remnants of lost tentacles or incipient tentacles? Haeckel (1879) then thought that he could recognize the species again in a me- dusa he collected in the English Channel. He had a relatively large medusa (4 mm high and wide), which had four tentacles with abaxial ocelli. A very unique feature of Haeckel's medusa was that it had four perradial gonads. He describes them as smooth, elongate, cylindrical bodies attached along the four perradial edges of the stomach. Unfortunately he provided no figure. Perradial gonads are rare in the Filifera, only in advanced stages of Merga species they can get fused perradially (see Bouillon, 1980; Schüchtert, 2007). Formally, Haeckels's medusa resembles thus Merga tregoubovi Picard, 1960 (see Schüchtert, 2007). In contrast to Haeckel's medusae, Merga tregou- bovi has no ocelli, but Haeckel sometimes misinterpreted dark pigments in the tentacle bulbs as ocelli. Whatsoever, it is by no means convincing that Haeckel had the same species as Slabber. 494 P. SCHUCHERT The uncertainty of the scope of Protiara tetranema is also reflected in Mayer (1910), who erroneously also associated Plotocnide borealis Wagner with this species. Hartlaub (1914) thought that Haeckel must have misinterpreted the gonads. He thinks that they were more likely the result of a perradial fusion of adradial bulges of very advanced gonads in a M erga species. The species was not seen after Haeckel except for Pell (1938) who gives an un- commented record of a single medusa in the Adriatic Sea. It remains unclear what Pell used as criteria to identify the medusa and later intensive searches (Benovic & Lucie, 1996) never found it again. Pell perhaps mistook a Merga species for Protiara tetra- nema. A second species of the genus Protiara was described by Hargitt (1902) as Protiara haeckeli, occurring in the Vineyard Sound, Massachusetts. This species has an apical process, lacks ocelli, and the manubrium is white. Hargitt remains somewhat vague concerning the position and form of the gonads. Protiara haeckeli has appa- rently also never been reported again and as its morphology remains incompletely known it is thus of limited help in elucidating the genus Protiara. A rather distinct, third Protiara species was described by Bigelow (1912) as P. tropica. Bouillon (1980) removed it from the genus Protiara to Pseudotiara (Family Bythotiaridae). Summarizing, Protiara tetranema is a doubtful, unrecognizable species. Because the only other species of the genus, P. haeckeli, is also somewhat doubtful, this undermines seriously the usefulness and validity of the genus Protiara and the family Protiaridae. Family Trichydridae Hincks, 1868 Diagnosis: Hydroid colonial, colony stolonal, stolons covered by thin perisarc; hydranths with a hydrotheca-like tube of perisarc at base into which polyp can with- draw, hydranths very small, sessile, with one amphicoronate whorl of filiform tentacles. Gonozooids are reduced hydranths without tentacles (blastostyles) enclosed in a filmy, soft perisarc-tube like a gonotheca. Medusa bell-shaped, with apical process, without gastric peduncle; manubrium with four large, folded lips; four radial canals; numerous fine, branched, anastomosing centripetal canals connecting non perradial marginal bulbs to radial canals; gonads interradial pads; marginal tentacles solid, with relatively large marginal bulbs; no ocelli. Remarks: The systematic position of this family and its sole genus is somewhat unclear. The majority of current authors (e. g. Bouillon et al., 2006) place it among the Anthoathecata based on its medusa, which is undeniably typical for this order. Werner (1984), however, classified it under Leptothecata. This was likely a conclusion he derived from his knowledge of the gonotheca of the hydroid stage (briefly mentioned in Werner, 1984 and pers. comm. Dr Anita Brinckmann-Voss). Trichydra and Halitiara share identical polyp stages and both are likely more closely related than previously thought. The cnidome of Trichydra remains imperfectly known, but the one of Halitiara is rather unique and resembles more the cnidomes found in the Leptothecata (i.e., Eirenidae, Eucheilotidae, Haleciidae. Lovenellidae and Tiaropsidae; Bouillon, 1985; Bouillon et al., 1988). The taxonomic position of Trichydra and Halitiara EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 495 remain thus unsettled and intriguing. Hopefully, molecular phylogenetic investigations will soon address the problem. Genus Tricky dra Wright, 1858 Synonym: Pochella Hartlaub, 1917 [type species Pochella polynema Hartlaub, 1917]. Type species: Trichydra pudica Wright, 1858 by monotypy. Diagnosis: See family diagnosis. Remarks: This is currently a monotypic genus. See also the remarks under Fabienna oligonema. Trichydra pudica Wright, 1 857 Fig 26-28 ? Oceania pusilla Gosse, 1853: 384, pi. 13 figs 11-14. not Eudendrium pudicum Van Beneden, 1867: 1 16, pi. 7 figs 1-2. - Rees, 1941:135. Trichydra pudica Wright, 1857: 168. -Wright, 1858: 257, pi. 15. Fig. 1. -Wright, 1863: 440, pi. 22 figs 1-6. -Hincks, 1868: 216, fig. 26. - Rees, 1941: 135, figs 4-5. - Hamond, 1957: 296. - Edwards, 1973: 87, Fig. 1A-E. - Fraser, 1974: 19. - Arai & Brinckmann-Voss, 1980: 75. fig. 42. - Werner, 1984: 187, fig. 1 16. Pochella polynema Hartlaub, 1917: 414, figs 344-346. - Russell, 1938a: 425, figs 1-3. - Russell, 1953: 394, figs 257-262. - Kramp, 1959: 179, fig. 258. - Kramp, 1961: 233. - Kramp, 1968: 109, fig. 296. Proboscidactyla polynema. - Foerster, 1923: 30, pi. 3 figs 5-7, pi. 4 fig. 1. ? Pochella polynema. - Goy et al, 1991: 118, fig. 45. Type material: Not located, likely lost. Material examined: BMNH 1954.11.13.383; as Pochella oligonema; England, Plymouth; mature medusa collected 30 April 1898; material depicted in Russell (1953: fig. 260). - BMNH 1955.11.23.1938; as Pochella oligonema; England, Plymouth; mature medusa collected 25.05.1934; material depicted in Russell (1953: fig. 258). - BMNH 1954.11.13.126, slide preparation, as Trichydra pudica; Ireland, Valentia; hydroid collected April 1900, reared colony from aquarium of M. J. Delap; donated by E. T. Browne, likely the material mentioned in Rees (1941: 335). Diagnosis: See family diagnosis. Description (Rees, 1941; Edwards 1973; own observations): Hydroid colonial, hydranths very small, connected by ramified, creeping, tubular stolons covered by thin perisarc. Hydranth with a basal hydrotheca-like tube of filmy perisarc into which the hydranth can partially or entirely contract, no operculum, the slightest disturbance makes them retract, hydranth very extensible, rather flexible, distal part can droop like a flower, hydranth body cylindrical, thin, hypostome conical, one whorl of 6-8 tentacles, tentacles thin, irregularly disposed nematocysts give spiny appearance to extended tentacles, fully extended often longer than polyp height. Reproductive polyps reduced to blastostyles producing one to two medusae at a time, contained in a perisarc tube resembling hydrotheca of the hydranths, these gonothecae without distinct operculum, blastostyle a simple rod-like process without tentacles. Newly liberated medusae very small, cup-shaped, jelly fairly thin, without umbilical canal; four short tentacles with conspicuous bulbs, four radial canals and a ring canal, radial canals without side-branches, stomach relatively large but without lips, velum broad, numerous nematocysts scattered over the exumbrellar surface; stomach and bulbs tinged with pale pinkish yellow. 496 P. SCHUCHERT Fig. 26 Trichydra pudica Wright, 1857; living polyps, modified after Rees (1941). Adult medusa with bell-shaped or hemispherical umbrella, slightly wider than high, jelly fairly thick, with broad, rounded apical process, without exumbrellar nema- tocyst tracks. Velum fairly broad. Stomach large, four-sided to cruciform in section, length about 1/2 to 2/3 the height of subumbrellar cavity; mouth cruciform through four rather large perradial lips, mouth-margin undulated. Four interradial gonads on stomach wall, cushion-like, each gonad when fully developed covering almost entirely the interradial wall of stomach. Thirty to forty or more solid marginal tentacles with large rounded basal swellings, without ocelli. Four straight smooth perradial canals fairly broad, usually also numerous fine, branched, anastomosing centripetal canals connecting non-perradial marginal bulbs to radial canals, connections of fine centri- petal canals to perradial canals occurring at varying levels. Colour of stomach and marginal tentacle bases brown. Nematocysts inadequately known, almond-shaped capsules resembling macro- basic heteronemes or merotrichous isorhizas, desmonemes most probably absent. Dimensions: Polyps hydranths up to 1 mm, tentacles when expanded may reach a length of 1.2 mm, basal perisarc collar (hydrotheca) 0.15-0.35 mm high and 0.1- 0.14 mm wide (Rees, 1941). Gonothecae 0.3-0.55 mm high and 0.15-0.2 mm wide. Newly liberated medusae from 0.15-0.2 mm in diameter and height (Edwards, 1973) to 0.3 mm wide and 0.4 mm high (Rees, 1941). Mature medusae may reach 3.85 mm in diameter and 3.47 mm in width (Edwards, 1973), mature medusae in the plankton have a diameter of 2-3 mm (Russell, 1953). EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 497 Fig. 27 Trichydra pudica Wright, 1857; medusa stage. A-C from Edwards (1973), D-E after Hartlaub (1917), F-G from Russell (1953). (A) Newly liberated medusa, height 0.15 mm. (B) Intermediate growth stage, diameter 1.2 mm. (C) Mature male medusa, bell height 3.3 mm, diameter 3.6. (D) Manubrium with mature gonads, medusa from plankton. (E) Bell rim with tentacle bases, note bulb-like swelling when the tentacles are contracted. (F-G) Schemata showing the variation of the branching pattern of the fine, centripetal canals. 498 P. SCHUCHERT Fig. 28 Trichydra pudica Wright, 1857; after material from Ireland (BMNH 1954.1 1.13.126, permanent slide-preparation); scale bar valid for all section, equals 0.1 mm. (A) Polyp with extended tentacles. (B-C) Polyps that are completely retracted into perisarc tube. (C-D) Gonothecae with blastostyles and maturing medusae. Other Data: The development of the medusae and the correlation of size and tentacle number are described by Russell (1953) and Edwards (1973). Biology: The hydroid was found on shells and stones (Wright, 1857; Hammond, 1957), clinker (=coal burning waste; Rees, 1941; Edwards, 1973), and the EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 499 ascidian Microcosmos claudicans (Savigny) (Hamond, 1957). The observations of Edwards (1973) indicate that medusae are liberated from April to June, becoming mature by July or August (Firth of Clyde). The length of life is about 2 months. Distribution: The hydroid is known from the British Isles (English Channel, North Sea, western and eastern Scotland, western Ireland; Wright, 1857; Hincks, 1868; Rees, 1941; Hamond, 1957; Russell, 1957; Edwards, 1973; this study). The medusa occurs in the British Isles (Russell, 1953; Fraser, 1974), the northern part of the North Sea (Hartlaub, 1917), and in the north-eastern Pacific along the coast of British Columbia (Foerster, 1923; Arai & Brinckmann-Voss, 1980). Goy et al. (1991) also reported it from the eastern Mediterranean, but their single medusa had only eight tentacles and the identification is not convincing. The occurrence in the Mediterranean needs reconfirmation. The remaining records of both medusae and polyp suggest a more northern Boreal distribution. Type locality: North Sea, Scotland, Firth of Forth; on shells and stones. Remarks: Most unusual for an Anthoathecata, this species has gonozooids in gonothecae, just like Halitiara formosa. The gonothecae of Tricky dra pudica were first mentioned in a brief species diagnosis given by Werner (1984), presumably based on unpublished observations. The gonothecae observed in this study were found in a sample identified as T. pudica by M. J. Delap and E. T Browne. As there is no infor- mation on the mature medusa connected to this sample, the identification remains somewhat uncertain. The polyps of Halitiara formosa are identical, but the medusa does not occur in the region. Some small Campanulinidae are also similar, though their hydrothecae usually have a pedicel and an operculum formed by perisarc flaps (comp. Cornelius, 1995a). The medusa of T. pudica was first described as Pochella polynema by Hartlaub (1917) and it was Edwards (1973) who linked it unambiguously with the hydroid T. pudica. The hypothetical linking of T. pudica with Lizzia biondina by Rees (1941) thus proved to be incorrect. The medusa is very rare and at first glance, or in damaged material from net catches, can be mistaken for a Podocoryna species (see Schüchtert, 2008a). The cruciform mouth and the absence of oral nematocyst clusters however allow them to be distinguished easily. Also the cnidome could be different, but unfortunately the cnidome of Trichydra remains inadequately known. ACKNOWLEDGEMENTS I gratefully acknowledge the help of my colleagues, especially Dr Dale Calder for advice on the ICZN, Drs C. Schandler & H. Rapp for the donation of the Thecocodium specimen, and Dr A. Brinckmann-Voss for the gift of specimens and her permission to reproduce original drawings of Halitiara formosa and illustrations of her publications. I am also deeply indebted to Dr Dhugal Lindsay who took the burden to read and comment an earlier draft of this manuscript. 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EUROPEAN ATHECATE HYDROIDS, FILIFERA 5 507 Index Ascidioclava 442 atentacidata, Tregoubovia All berne, Protiara 492 bigelowi, Calycopsis 451 brieni, Thecocodium All brooksi, Proboscidactyla 462 brownei, Eucodonium AI 9 Bythotiara 442 Bythotiaridae 442 Bythotiaridi 442 Calycopsidae 442 Calycopsis 445 chuni, Calycopsis 449 conica, Proboscidactyla 458 cornubica, Willisa 462 Crypta 442 dendrotentaculata, Niobia ASA Dicranocanna 458 digitalis, Paratiara 491 Dyscannota 457 dysdipleura, Dyscannota 458 Endocrypta 442 enigmatica, Kantiella A61 Eucodoniidae 479 Eucodonium 479 Fabienna 468 flavicirrata, Proboscidactyla 458 formosa, Halitiara 486 furcata, Willia 462 furcillata, Dicrocanna 458 gara, Calycopsis 453 gaussi, Dissonano 486 'gemmifera, Proboscidactyla 458 geometrica, Calycopsis 455 geometrica, Sibogita 455 Halitiara 486 huntsmani, Crypta 442 Hydrolaridae 456 inflexa, Halitiara 490 Kantiella A61 krampi, Calycopsis AA1 Laingia 466 Laingiidae 466 Lar 457 Laridae 457 Magapiidae 466 mirabilis, Russellia 481 Misakia 458 murrayi, Bythotiara AA3 mutabilis, Proboscidactyla 458 Mofc/a 483 Niobiidae 483 nuarchus, Sibogita 446 occidentalis, Sibogita 455 oligonema, Fabienna 469 oligonema, Pochella 469 ornata, Proboscidactyla 458 parasitica, Ascidioclava 442 Paratiara 490 penicillatum, Thecocodium AIA Pochella 470 polynema, Pochella 470 Proboscidactyla 457 Proboscidactylidae 456 prolifera, Psythia 458 Protiara 492 Protiaridae 486 iVyf/2/a 458 Ptilocodiidae 470 pudica, Trichydra 495 pudicum, Eudendrium 495 pusilla, Oceania 495 repens, Ptilocodium All Russellia 481 Russelliidae 481 sabellarum, Lar 462 Sibogita 454 simplex, Calycopsis 449 simplex, Leuckartiara 487 sphaerica, Fabienna All stellata, Proboscidactyla 462 sfeZZata, WifZZ/a 462 stolonifera, Proboscidactyla 458 tetranema, Protiara 492 Thecocodium All Tregoubovia AI 6 Trichydra 495 Trichydridae 494 tropica, Proboscidactyla 458 ry/?a, Calycopsis 446 typica, Misakia 458 valdiviae, Calycopsis 449 varions, Proboscidactyla 458 WWfeito 458 Wï«ifl 457 Williadae 456 Williadi 456 MZZsza 457 Willsiadae 456 Willsiidae 456 Revue suisse de Zoologie 1 16 (3-4): 509-633; décembre 2009 The Hilara species (Diptera, Empididae) of Switzerland, with respect to the fauna of the Alps and other central European mountains Milan CHVÄLA 1 & Bernhard MERZ 2 1 Charles University, Department of Zoology, Vinicnâ 7, CZ-128 44 Praha 2, Czech Republic. E-mail: mchvala@natur.cuni.cz 2 Muséum d'histoire naturelle, Department d'Entomologie, C. R 6434 CH-1211 Genève 6, Switzerland. E-mail: bernhard.merz@ville-ge.ch The Hilara species (Diptera, Empididae) of Switzerland, with respect to the fauna of the Alps and other central European mountains. - Within the fauna of the Alpine region 1 1 8 Hilara species are recognized including some species which are not yet known from the region but may be expected. The species are keyed, and full distributional data are given for the Alps. Eight new species are described: Hilara leukensis sp. n., H. luteihalterata sp. n., H. pianti sp. n. and H. crossleyi sp. n. from the Swiss Alps, H. bre- vistriata sp. n. from the Swiss and Austrian Alps, H. parvimaior sp. n. from the Swiss and Italian Alps, H. sirbitzmatrona sp. n. from the Austrian Alps, and H. polleti sp. n. from the Bulgarian Rhodope Mts. Several further species are fully redescribed and illustrated for the first time (H. cinereo- micans Strobl, H. sartor Becker, H. tiefii Strobl, H. tanychira Strobl, H. di- midiata Strobl, H. simplicipes Strobl, H. calinota Collin, H. pruinosa Wiedemann in Meigen, H. hystrix Strobl, H. pectinipes Strobl), and the females of H. helvetica Chvâla and H. zermattensis Chvâla are described here for the first time. A lectotype is designated for H. pilosopectinata Strobl. The following new synonymies are proposed: Hilara maior Strobl, 1910 (= Hilara tatra Niesiolowski, 1991) and Hilara tyrolensis Strobl, 1892 (= H. miriptera Straka, 1976). The synonymy of Hilara longesetosa Strobl, 1910 with H. pilosopectinata Strobl, 1892 is explained. Hilara longicornis Strobl, 1 894, described and still known only from the Hungarian lowlands, is fully redescribed and its correct classification within the genus Hilara is discussed in the section "additional species". The same applies to Hilara flavocoxa Straka, 1976, a species known only from a single locality in high- lands of central Slovakia. Keywords: Diptera - Empididae - Hilara - Central Europe - faunistics - taxonomy - key - new species - new synonymies. INTRODUCTION There are two recent basic monographs available for the genus Hilara (Diptera, Empididae) in Europe, one covering all Fennoscandian, British and partly central Manuscript accepted 13.10.2009 510 M. CHVÂLA & B. MERZ European species (Chvâla, 2005a), the second deals with the Mediterranean fauna (Chvâla, 2008a). There is, however, a distinct group of species mostly of central European distribution, mainly mountain forms, which were included neither within the north European fauna, nor in the Mediterranean study. Such species were not keyed in the above two monographs. Therefore, the authors present here a publication which enables the identification of all species in temperate central part of Europe, covering in particular the special mountain and Alpine species of the genus. In addition, some mountain species that occur also in the north (mostly species of boreomontane distri- bution), or species known also from the warm coastal lowlands, are included here as well. Furthermore, all species which were found or may be found in Switzerland are discussed and keyed. If the reader cannot identify "his" species with the present key, then he is recommended to consult the identification keys in Chvâla (2005a) and Chvâla (2008a). There is still a real possibility that some southern species are occurring in more northern regions, like the lowlands of the Alpine region. If the reader does not succeed even after that, then there is only a third possibility: he has a new, undescribed species before him. The fundamental work on the Alpine Hilara (Diptera: Empididae) is undoub- tedly the more than 100 years old monograph of Strobl (1892a), based mainly on the fauna of the Austrian Styrian Alps. Strobl recognized at that time already 62 species and several varieties or forms. Of the 27 described new species, 20 of them remain valid, and only 7 species were later regarded as junior synonyms. All the 5 new varieties or forms described by Strobl in this monograph were later raised to specific rank and are now considered to be valid species as well. The 7 taxa which were later synonymized include new descriptions of both sexes of Hilara albitarsis von Roser (unknown to Strobl), or the complex of the north European group of H. longivittata Zetterstedt. This species was described by Strobl as the new species H. bivittata, and all forms described as new by Strobl represent now well recognized species of the distinct H. andermattensis group of species, a complex of exclusively high alpine Hilara species. Strobl continued to study the Styrian Hilara species in the fundamental five- volume work "Die Dipteren von Steiermark" published in 1893 - 1910. Two further monographs dealing with the Alpine Hilara species by Keiser (1947) and Franz (1989) do not reach the high quality of the Strobl's publications and, therefore, they are not discussed here more in detail. For instance, Keiser (1947: 126) recognized only 9 species of Hilara in the Swiss National Park. All species were listed from many localities. A re-examination of them by the first author (MC) revealed that these specimens belong to complexes of several related species. The same applies to the elaboration of the genus Hilara by Franz (1989: 321) in the two- volume mono- graph on the Alpine Diptera of Austria. Franz mentioned altogether 73 Hilara species on pp. 321-330, but many of these names are unavailable, or synonyms, and most spe- cies are based mainly on misidentifications of E. O. Engel and E. Lindner, who both were not at all specialists of this genus. It was later Collin (1961) in his monograph of Hilara of the British Isles whose treatment is of a very high level. He understood the genus well but, since he did not study the Strobl collection, some of the species described by him (Collin, 1927) were later found to be younger synonyms of Strobl's taxa. HILARA FROM SWITZERLAND 5 1 1 In all 65 Hilara species were recorded in Switzerland in the first Checklist (Chvâla et al., 1998), additional 13 species of the genus were included in the first Supplement (Merz et al., 2002), and a further 8 species in the second Supplement (Merz et al., 2007). The present revision includes a further 12 species found in Switzerland for the first time but 3 species are deleted because of misidentifications. Therefore, in total 95 species are recorded with certainty from Switzerland. THE ALPINE FAUNA The species composition of the 118 Hilara species recorded in the present pu- blication is especially interesting. There is a large group of 18 clearly alpine endemics, i. e., species known from the Alps only (H. alpicola Chvâla, H. cinereomicans Strobl, H. pianti sp. n., H. crossleyi sp. n., H. tiefii Strobl, H. sartor Strobl, H. leukensis sp. n., H. brevistriata sp. n., H. luteihalt erata sp. n., H. helvetica Chvâla, H. parvimaior sp. n., H. styriaca Strobl, H. merzi Chvâla, H. zermattensis Chvâla, H. andermattensis Strobl, H. simplicipes Strobl, H. sulcitarsis Strobl and H. tyrolensis Strobl); 1 1 species are boreo-montane faunal elements (some perhaps of boreo-alpine distribution if of the last glacial period origin), though scarcely they are of arcto-alpine distribution (for instance, H. setosa Collin, H. eviana Straka, H. griseola Zetterstedt, H. intermedia (Fallen), H. bistriata Zetterstedt, H. coracina Oldenberg, H. scrobiculata Loew, H. diversipes Strobl and H. hybrida Collin). Finally, 8 species are exclusively mountain species of central temperate Europe which are known besides the Alps also from other central European mountains (H. nigrita Chvâla, H. goetzei Chvâla, H. perversa Oldenberg, H. tetragramma Loew, H. quadrifaria Strobl, H. calinota Collin, H. triseta Chvâla and H. hystrix Strobl). Four species are known in temperate central parts of Europe, which occur both in lowlands and high in mountains, including the Alps (H. pilosopectinata Strobl, H. borealis Oldenberg, H. pectinipes Strobl and H. bohemica Straka). It is difficult to classify at present the mountain Hilara species within special altitudinal zones, as it was done for instance by Ziegler (2008) for the fauna of Diptera in the Italian Stilfserjoch National Park. Collecting efforts in the long and diverse Alpine chain from western Austria to the Mediterranean coast of France and Italy are very scattered and insufficient and many regions are unstudied. Therefore the available data is too scarce to allow a classification of ecological or distributional patterns of the species. The other Hilara species occurring in the Alpine region are widely distributed in Europe, from the Fennoscandia in the north, and through central temperate Europe south at least to the Alpine region, very often right up to the Mediterranean. In central parts of Europe they are distributed mainly in lowlands and submountain regions, and absent from higher altitudes. There is, however, a small group of about 7 species representing faunal elements of south-western Europe, which are mainly widely distributed in Spain, often including the Pyrenees, and distributed north-west along the French coast right up to the western part of the Alpine region, especially south east France and the adjacent parts of Italy and Switzerland. As examples may be mentioned the following species: H. morenae Strobl, H. ponti Chvâla, H. veletica Chvâla, H. pygialis Chvâla, H. pria- 512 M. CHVÂLA & B. MERZ nosa Wiedemann, but also for instance H. pseudocornicula Strobl, a very common spe- cies in Spain and the Mediterranean which was known for a long time also from wes- tern parts of Europe along the Atlantic coast as H. subpoliinosa Collin. These species are discussed in the corresponding paragraphs on "Distribution". There are still possi- bilities that some further southwestern species of the Spanish fauna will be found in the Alpine region in the future. There are, however, a further two not clearly mountain Hilara species which are mentioned here, as they were not included in the previous two main monographs on the genus Hilara in Europe. First, it is Hilara longicornis, described by Strobl (1894) from 10 â and 3$ collected by Prof. Thalhammer on 1. v. 1893 at Hajós in Hungary. Five syntypes (3o* and 29) are in the Strobl Collection in Admont. Becker (1894) erroneously supposed it was a species of the genus Iteaphila Zetterstedt, 1838 (cited as Steleocheta Becker, 1887) of the empidid subfamily Oreogetoninae because of some morphological pecu- liarities (very long antennae, simple unmodified legs, the open rather "Empis"-\\k& radial fork). However, H. longicornis is a typical Hilara species possessing all the main generic characters. The species is keyed below within the mountain species, and it is redescribed in the section "additional species". Secondly, it is Hilara flavcocoxa, described by Straka (1976) from the locality Tur ek at Kremnické vrchy (Kremnické hills) in the lowlands of central Slovakia. This species seems to be closely related to H. merula Collin and, like the preceding H. lon- gicornis, it is also fully redescribed and discussed in the section "additional species". THE FAUNA OF SWITZERLAND Austria has a long dipterological tradition, in particular for the genus Hilara, thanks to the activity of Strobl. On the other hand, Switzerland did not receive a parti- cular attention for this genus in the past. Before 1930 only few specimens were collected by Th. Becker and L. Oldenberg during their short visits in few places in the Alps. Later, F. Keiser collected extensively in the Swiss National Park and published his identifications (Keiser, 1947). However, he did not specifically search for Hilara and, therefore, the fauna remains poorly known even for this region. Later, Ringdahl (1957) mentioned 4 further species. The main efforts with specific collecting of Hilara started only recently with G. Bächli (from 1971), J. -P. Haenni (from 1980) and the second author (BM) (from 1994) who organzied several shorter and longer field trips to various localities in the Swiss Midlands and the Alps, sometimes helped by students or visitors. Their specimens were mostly included in the numerous revisions of the first author, the checklist and the two supplements (see references). As only few dipterists have made a special effort they were not collecting at all suitable localities, and many regions in Switzerland remain unstudied. This is in particular the case for the Jura mountain chain for which almost no specimens are available and which may host further nordic species as it is for instance the case of the Chloropidae (Merz et al., 2005). NEW SYNONYMIES Three new synonymies of the Alpine Hilara species are given below. First, it is Hilara tatra Niesiolowski, 1991, described from the Polish Tatra Mts, which is conspe- HILARA FROM SWITZERLAND 513 cific with H. maior Strobl, 1910, the only widely distributed species in mountains of central Europe of the otherwise clearly Alpine Hilara andermattensis group of species. The second is Hilara longesetosa Strobl, 1910, described from the Styrian Alps, which Strobl (1892a) inadequately described already earlier as Hilara pilosopectinata from the Austrian locality Gmünd. However, Chvâla (2008b) already briefly mentioned this synonymy within a revision of the Diptera described by Gabriel Strobl. Finally, the third is Hilara miriptera Straka, 1976, described from the Slovenian Julian Alps, which is conspecific with Hilara tyrolensis Strobl, 1892. MATERIAL AND METHODS The present study is primarily based on several thousand specimens from the Styrian and Swiss Alps. The basic collection was that of Pater (Father) Gabriel Strobl in the Styrian Admont, who built up his large and very important scientific collection of Alpine Hilara between 1887 and 1910 (see also Chvâla, 2008b). Results of his field studies were first published in the monograph of Austrian Hilara species (Strobl, 1892a), followed by further data in the five volume monograph of Strobl "Die Dipteren von Steiermark" where the Hilara species were treated in three volumes in 1893, 1898 and 1910 (see references). This region of the Styrian Alps, covering the Ennstaler Alpen including the Gesäuse, and the Rottenmanner and Schladminger Tauern, was later very intensively studied again by the first author (MC), from 1995 until now. The second well studied area is Switzerland. The second author (BM) collected, often together with his collaborators and students, with priority in the Alps (Grisons, Valais, Ticino), but also in the Swiss Midlands around Zürich (1988-1998) and Geneva (from 1999 until now). G. Bächli and J.-R Haenni have kindly loaned for study further material from Switzerland. Especially interesting were specimens collected by P. Stucki at light traps, and by C. Besuchet at the window of his winter garden, both in the vicinity of Geneva. The second author (BM), and also J.-P. Haenni, A. C. Pont, M. Bartak and E. Castella & M. C. D. Speight, collected numerous specimens in the French Alps and the adjacent mountains. J.-P. Haenni and M. Bartâk helped also with specimens from the Italian Alps. Finally, A. R. Plant put at our disposal many interesting specimens from the Slovenian Julian Alps and the hilly country on the Slovenian-Croatian border. It should be noted that over 90% of the specimens were collected by sweeping with an insect net on vegetation or over the water surface. Only few specimens were obtained with traps (Malaise trap, light trap) or other methods. The specimens studied are housed in the collections cited below. It should be noted that the entire collection of the first author (MC) is now deposited in the UMO where also the large collection of J. E. Collin is preserved. A major part of the speci- mens collected by the second author (BM) before 1999, sometimes with his former students (for instance, C. Wolf and P. Riiegg) is deposited in the ETHZ. The specimens collected since 1999, as well as a reference collection of all species, are now stored in the MHNG. In the following systematic treatment only records from the Alpine region (Austria, France, Italy, Slovenia, Switzerland) are listed under "Material examined" for the common species which occur also elsewhere. Detailed collecting data for these species were published in the revisions and monographs of Chvâla (see references). 514 M. CHVÄLA & B. MERZ BSA Benediktinerstift Admont (coll. G. Strobl) (Austria) CGB Private collection G. Bächli, Dietikon (Switzerland) ETHZ Eidgenössische Technische Hochschule, Zürich (coll. J. Escher-Kündig, G. Huguenin, B. Merz part, W. Sauter) (Switzerland) (curator: A. Müller) MHNG Muséum d 'histoire naturelle Genève (coll. B. Merz, part) (Switzerland) MHNN Muséum d'histoire naturelle Neuchâtel (coll. J.-P. Haenni) (Switzerland) NHMB Naturhistorisches Museum Basel (coll. F. Keiser) (Switzerland) (curators: M. Brancucci, D. Burckhardt, F. Keiser) NML Natur Museum, Luzern (coll. L. Rezbanyai-Reser) (Switzerland) (curator: L. Rezbanyai-Reser) UMO University Museum Oxford (coll. J. E. Collin, coll. M. Chvâla) (U.K.) ZMB Zoologisches Museum der Humboldt Universität, Berlin (coll. L. Oldenberg, T. Becker) (Germany) (curators: H. Schumann, J. Ziegler) For the descriptions and diagnosis we adopt the terminology of morphological structures of Chvâla (2005a, 2008a). The following abbreviations are used for setae and legs: for setae (bristles) for legs acr - acrostichals dc - dorsocentrals h - humeral ih - intrahumeral ntpl - notopleural pa - postalar ph - posthumeral sa - supra-alar sc - scutellar bt - basitarsus (i.e., first tarsomere) ex - coxa f - femur t - tibia 1,2, 3 - indicate fore, mid and hind legs KEY TO THE ALPINE HILARA SPECIES The key is based on dry-pinned specimens. It should be noted that specimens which were collected in alcohol and dried only later, which are still stored in alcohol, or which are teneral, are often paler (in particular the legs and the haltère) or either smaller (shrivelled for teneral specimens) or larger (full of liquid for alcohol-preserved specimens) and may therefore lead to confusions. It is recommended to use the following key in conjunction with the illustrations in Collin (1961) and Chvâla (2005a, 2008a). 1 Small, slender, weakly built species with slender yellowish legs, body with a tendency to be yellowish, h (and usually also ih) bristle absent or very inconspicuous, but pronotum (unlike H. maura-group) always with a distinct bristle on each side. Body and legs devoid of distinct bristles; legs covered with only weak bristly-hairs, simple and slender, male bt\ and female f 3 unmodified, slender (group 1, H. flavip es- group) 2 HILARA FROM SWITZERLAND 5 1 5 Body and legs with distinct bristles or bristly-hairs (with the exception of the H. maura-group, but then pronotum without a distinct bristle on each side, and legs modified); if male bt\ slender and unmodified, as stout as following tarsomeres (H. tywlensis, simplicipes, perversa), then legs extensively blackish 5 2(1) 6: eyes touching on frons, upper ommatidia distinctly enlarged; no frontal bristles, and anterior four tibiae with distinct fine bristly-hairs beneath. $ : haltère yellow 3 ô : eyes narrowly separated on frons, all ommatidia equally small, frons about as deep as anterior ocellus. Long frontal bristles present in both sexes 4 3(2) Thoracic pleura blackish-grey. Ocellar bristles longer than upper occipi- tals. ô : tarsi not very long, bt\ together with 2nd tarsomere shorter than fj, and mid tarsus shorter than f 2 - 2: frons dull light grey, no distinct frontal bristles 1 . H. flavipes Meigen Thoracic pleura translucent yellowish. Ocellar bristles small, not diffe- rentiated from upper occipital bristles, â : tarsi very elongated, bt\ toge- ther with 2nd tarsomere longer than f], and mid tarsus clearly longer than t 2 . 9 : frons velvety-black, frontal bristles long . . . 2. H. flavidipes Chvâla 4(2) S : abdomen contrastingly snow-white on anterior four to five segments. 2 : haltère darkened, knobs grey to blackish-grey; h and ih bristles not differentiated 4. H. albiventris von Roser a: abdomen uniformly dark on dorsum, terga brownish-black with narrowly paler membraneous hind-margins. 9 : haltère yellow 4a 4a(4) Antennae uniformly blackish, segment 2 rarely slightly brownish; h and ih setae distinct, well visible, even if fine. Legs yellow on femora, more or less brownish on posterior four tibiae and tarsi. S : Frons rather wide, about as wide as anterior ocellus, and parallel-sided 3. H. morenae Strobl Basal antennal segments yellowish-brown; h and ph setae minute. Legs extensively yellow, only tarsi darkened towards tip. S : Frons very nar- row above, almost linear, narrower than anterior ocellus, and distinctly widening below towards antennae 3a. H. sturmii Wiedemann in Meigen 5(1) Body and legs devoid of distinct bristles; h, ih, and ph bristles absent, pronotum (except for H. media) without a bristle on each side. A distinct sensory pit below prothoracic spiracle. Legs with/3 often thickened and abruptly narrowed at tip, black (only party translucent yellowish in H. media and H. discalis), bty in S very thickened (except for//, simpli- cipes) (group 13, H. maura-gvoxxp) 6 Body and legs with distinct bristles or bristly-hairs, the strong marginal bristles on scutum only rarely absent, and pronotum always with one or more distinct bristles on each side. No sensory pit below prothoracic spiracle, and / 3 not clearly thickened (except for H. coracina and H. albitarsis) 17 6(5) / 3 more or less thickened, often abruptly narrowed at tip, and hairs on apical third beneath arising from small warts. The sensory pit below prothoracic spiracle covered with fine hairs 7 516 M. CHVÄLA & B. MERZ f 3 slender, not dilated, and the hairs beneath towards tip not arising from small black warts (except for H. diversipes 9 , but then a large species, 3.5 - 5 mm long) 13 7(6) Scutum with 3 dark stripes which, when viewed from in front, coalesce into a broad rusty brown median stripe; when viewed from behind, this stripe becomes velvety black and is sharply separated from the silvery sides. / 3 dilated in both sexes, but not abruptly narrowed at tip; acr irre- gularly 4- to 6-serial, dc 2- to 3-serial, all minute. Prothoracic spiracle almost whitish, â : lateral genital lamella with a cluster of golden-yellow hairs towards tip (as in H. diversipes). 9 : f 3 evenly dilated towards tip. Large, about 4 - 4.5 mm long 104. H. maura (Fabricius) Scutum rather uniformly shining or subshining black; if finely pollinose and striped, then stripes not coalescent in anterior and posterior views. / 3 abruptly narrowed towards tip; if in doubt (H. hybrida 9 and H. diversipes 9 ), then acr and dc less numerous, the former 3- to 4-serial, the latter almost 1 -serial, and prothoracic spiracle dark blackish-brown (whitish in H. maura) 8 8(7) Generally large species, body 3.5 - 5.0 mm long. Scutum finely polli- nose, with 3 indistinct dark stripes in anterodorsal view, and micro- scopically silvery pilose in frontal view. Palpi mostly with pale hairs beneath, only a few longer bristly hairs brownish, acr irregularly 4- serial, minute. 9 : f 3 dilated 9 Generally smaller species, body about 2.5 - 4 mm long. Scutum almost polished black, at least subshining from all points of view, unstriped. Palpi with dark to blackish hairs beneath. / 3 strongly dilated and abruptly narrowed before tip in both sexes. 9 : f 3 slender 10 9(8) â : lateral genital lamella with a cluster of pale golden-yellow hairs at tip (as in H. maura). dc almost 1 -serial anteriorly. 9: / 3 rather slender, practically not dilated; r 3 more evenly dilated towards tip; sternum 8 elongate, polished above, dusted below 105. H. diversipes Strobl a : lateral genital lamella with small dark brownish hairs, dc narrowly 2- serial anteriorly. Wings clear (brownish in the morphologically similar H. nitidula). 9 : / 3 dilated and abruptly narrowed at tip; f 3 slender in basal third, then conspicuously dilated apically; sternum 8 small, largely polished black 106. H. hybrida Collin 10(8) acr irregularly 4-serial, anteriorly often 3-serial, very small and in- conspicuous; also ocellar and frontal bristles generally smaller 11 acr irregularly 2-serial, anteriorly often 3-serial, then more distinct and hair-like, although small; if in doubt, and acr exceptionally almost 4- serial (H. femorella), then strong ocellar and frontal bristles in combi- nation with brown clouded wings 12 11(10) acr irregularly 3- to 4-serial, dc 2-serial at middle, at least 3-serial ante- riorly and posteriorly, all very minute. Antennal style about one-half length of 3rd segment, and palpi with one bristly hair in addition to the short hairs, no bristles. Ocellar bristles fine, frontal bristles often not H1LARA FROM SWITZERLAND 517 differentiated. / 3 below towards tip covered with soft yellowish hairs. S : bty large, equal to about 3/4 length of tibia; dorsal process of lateral lamella flattened, blunt-tipped. Larger, almost 4.0 mm long. Spring species 107. H. nitidula Zetterstedt acr almost regularly 4-serial, dc mostly 1 -serial, more hair-like. Antennal style longer, almost as long as 3rd segment, and palpi more densely bristled. Ocellar and frontal bristles well developed, bristle-like. / 3 with black bristly hairs ventrally towards tip. S : bt^ smaller, about 2/3 length of tibia; dorsal process of lateral lamella narrowed and sharply pointed. Generally smaller, about 3 - 3.5 mm long. Summer species . . . 108. //. nitidorella Chvâla 12(10) Abdominal pubescence blackish-brown, wings brownish. Ocellar and frontal bristles strong; acr and dc hair-like, about as long as 2nd tarso- mere of fore tarsi. 6: bt\ shorter than tibia; hypandrium small, not produced posteriorly 109. H. f emorella Zetterstedt Abdominal pubescence whitish, wings clear. Ocellar bristles small and fine, shorter than antennal style, and frontal bristles at most half as long, often invisible; acr and dc minute. S: bt\ conspicuously large, as long as tibia; hypandrium produced posteriorly, very flattened, subhyaline along hind margin 110.//. sulcitarsis Strobl 13(6) Pronotum without a bristle on each side, covered with uniform minute hairs 14 Pronotum with a distinct bristle on each side. Wings clear, acr 4-serial. S : bt\ stout but short, scarcely of 2/3 length of tibia. Rather a small spe- cies, body about 3 - 3.5 mm long (see also couplet 103) . 115.//. media Collin 14(13) Legs uniformly blackish including "knees". The sensory pit below pro- thoracic spiracle without hairs. Radial fork very acute, "oreogetonine- like"; acr narrowly 2-serial. ? : / 3 simple and slender 15 "Knees" yellowish. The sensory pit below prothoracic spiracle guarded by silvery hairs. Radial fork of the usual shape, not acute, "hilarine- like". Medium-sized species, body about 2.5 - 3.5 mm long 16 15(14) Abdominal pubescence blackish. Wings faintly brownish, veins blackish. S: bt^ very thickened. Larger species, body 3 - 3.5 mm long 111.//, clypeata Meigen Abdominal pubescence whitish. Wings milky-white, veins whitish, â: bt\ slender, unmodified. Smaller species, body about 2.5 mm long (= //. miriptera Straka, syn. n.) 112.//. tyrolensis Strobl 16(14) Frons subshining, with a small, shining ovate patch above antennae; clypeus largely polished black, acr 2-serial, in two widely separated rows, sometimes more numerous in front, dc 1 -serial. Legs translucent yellowish on cx l5 and also more or less so on all trochanters and femora. 5 : r 3 slender, unmodified 113.//. discalis Chvâla Frons dull velvety black; clypeus greyish dusted, narrowly polished on upper margin, acr almost regularly 4-serial, dc irregularly 2-serial. Legs black except for narrowly yellowish "knees". 9 : r 3 strongly dilated towards tip, slender at base 114.//. discoidalis Lundbeck 518 M. CHVÂLA & B. MERZ 17(5) Legs yellow, at least fore legs partly yellow on coxa and base of femur, or legs almost uniformly brownish-yellow to dark brown 18 Legs uniformly black, at most "knees" partly yellowish; if the colour varies, the species will be found in both sections (care must be taken with immature specimens with paler legs) 48 18(17) Yellow species, ground colour of thorax yellow 19 Black or greyish species, ground colour of thorax black, at most humeri may be translucent yellowish 20 19(18) Large species, body about 4 - 4.5 mm long. Eyes narrowly separated on frons in 6 , occiput blackish-grey. Scutum extensively yellow, with long, hair-like irregularly 2- to 3-serial acr, and longer 1 -serial de. â : bt\ long cylindrical, indistinctly stouter than tibia; 9 : f 3 slightly flattened and curved 24. H. thoracica Macquart (Here belongs also Hilara flavitarsis Straka, 1976, described from Albania/Macedonia; for details see remarks under H. thoracica) Small species, body about 3 mm long. Eyes, as usual, broadly separated on frons, occiput light greyish. Scutum finely, almost silvery-grey Polli- nose, acr irregularly 3- to 4-serial, small and numerous, dc 1 -serial and less numerous, about 10 bristles in one row. 8: bt\ long oval, stouter than tibia; 9 : f 3 slender, and abdomen silvery dusted on dorsum 38. H. temila (Fallen) 20( 1 8 ) Haltère clear yellow (knob darker grey in H. allogastra), and occiput dull grey, not blackish from any point of view 21 Haltère blackish, or at least knob greyish-black; if in doubt (for instance H. cuneata 9 with yellow haltère), species with occiput black from some points of view belong here 28 2 1(20) acr 2-serial, dc-l -serial, long and bristle-like, less numerous, about 10 in one row and all about as long as antennal style 22 acr irregularly 3- to 4-serial, at least 3-serial at middle; if almost 2-serial (H. gallica), then large species about 3.5 - 4.5 mm long 23 22(21) Larger species. 3 - 3.5 mm long. S : legs with the usual scattered hairing and bristling, hypopygium upturned with an unusually stout, shining black hypandrium. 9: f 3 distinctly thickened and compressed, contras- ting blackish including tarsus; all femora and anterior tibiae yellowish. Palpi clear yellow 13. H. canescens Zetterstedt Smaller species, 2.5 - 3 mm long. cT : tibiae and basitarsi on anterior two pairs unusually dilated and very densely, long black bristled. 9 : fcj only slightly dilated and curved, not contrasting black; legs extensively dar- kened on all tibiae and tarsi. Palpi dark brown to blackish 14. H. nigritarsis Zetterstedt 23(21) Small, about 3 - 3.5 mm long; acr and dc long and bristle-like, less numerous (about 10 in one row), almost as long as antennal style. Scutum dull brownish-grey except for lighter grey scutellum. Legs extensively yellow, tarsi darkened. Female: f 3 only slightly thickened and distinctly curved. Haltère often extensively darkened (see also couplet 35) 18.//. eviana Straka HILARA FROM SWITZERLAND 519 Large species, about 4-5 mm long (except for H. discolor); acr and dc small and numerous. Rather robust, light grey dusted species on occiput and thorax, female ? 3 simple (group 2, H. abdominal lis -group) 24 24(22) Frontal bristles absent, not differentiated from other small hairs on frons. Palpi and basal antennal segments vary in colour from dark brown to black. Shorter- and stouter-legged species, especially femora stout, legs very yellow, only tarsi darkened towards tip. Scutum light grey, with only an indication of 2 darker lines between rows of bristles 8. H. gallica (Meigen) Frontal bristles distinct, often not much shorter than ocellars. Palpi yellowish. Longer-legged species, legs usually more or less darkened on tibiae and tarsi, and more distinctly bristled 25 25(24) Scutum with 3 indefinite darker stripes along the lines of acr and dc. Pronotum with a single, usually light brown bristle on each side, pro- thoracic epistema with white hairs. Basal antennal segments and legs varying in colour from yellow to dark brown, tarsi always blackish, a : bt\ cylindrical, dark in contrast to yellow tibia, about 2/3 as long as length of tibia. Generally smaller, about 3.5-4 mm long . . 9. H. discolor Strobl Scutum with 2 narrow, more or less distinct darker lines between rows of bristles. Prothoracic epistema with black hairs, and pronotum with several small black spines in addition to the large black bristle on each side. Larger sized species 26 26(25) 6 : bty very enlarged, as long as, or longer than t x , also 2nd tarsomere on fore tarsi clearly stouter than following tarsomeres. All tibiae with dense, long black pubescence, the hairs longer than tibiae are deep. Legs yellow, with all tarsi and often also tibiae towards tip blackish. Basal antennal segments brownish. Pronotum usually with 4 small spines on each side in addition to the large bristle. 2 : Abdomen brown to yellowish-brown, more or less greyish dusted; bty long and slender, about 2/3 as long as length of ty Haltère uniformly yellow. Large, about 5 mm long 5. H. abdominalis Zetterstedt a : bt\ shorter, about 3/4 as long as length of t\ ; t 2 almost bare, covered with only minute hairs and a few short black bristles towards tip. Basal antennal segments yellowish 27 27(26) Generally smaller, about 4 - 4.5 mm long, haltère clear yellow. Pro- notum usually with 4 small black spines on each side in addition to the large bristle. S : Fore tarsi with 2nd tarsomere swollen, nearly twice as deep as the following tarsomere. Dorsum of abdomen subshining black in posterior view, otherwise greyish dusted. $ : bt\ shorter compared with H. abdominalis, about half as long as f j 1 . H. alpicola Chvâla Large species, about 5-6 mm long, haltère with knob more or less greyish. Pronotum with 6 to 8 small spines on each side in addition to the large bristle. 6: 2nd tarsomere on fore tarsi scarcely stouter than following tarsomeres. Dorsum of abdomen silvery-grey dusted in o\ glittering silvery-white in $ (see also couplet 30) . . . . 6. H. allogastra Chvâla 520 M. CHVÂLA & B. MERZ 28(20) Occiput and frons dull grey, occiput not dull black from any point of view . 29 Occiput dull black, at least from some points of view, frons usually blackish. Abdominal pubescence black; if pale (H. pseudosartrix) then legs extensively brownish and male genitalia with an unusual, long, slender hypandrium 36 29(28) Large species, body about 4-6 mm long 30 Smaller species, body 3 - 3.5 mm long 31 30(29) acr minute, closely 2-serial, almost 1 -serial anteriorly. 6: Fore tibia and tarsus with conspicuously long black bristles. 9 : Abdomen dull greyish- black, tergum 2 at sides with a tuft of long light brownish hairs (group 3, H. clavipes-gvoup, part) H.H. clavipes (Harris) acr 4-serial. S : Fore legs with only short pubescence, without distinct bristles except for a circlet of preapical tibial bristly hairs. 9 : Abdomen silvery-white glittering on dorsum, tergum 2 with short hairs at sides (group 2, H. abdominalis-group, part) (see couplet 27) . 6. H. allogastra Chvâla 31 (29) acr 4-serial 32 acr 2-serial, long and bristle-like, palpi blackish. S : bty very dilated, about 2/3 as long as fj, and with long bristly hairs towards tip; a pair of long bristles also at tip of first two tarsomeres. 9 : r 3 slender, but slightly curved 16. H. setosa Collin 32(31) Abdominal pubescence pale, at least at base of abdomen; acr and dc numerous, fine and small, equally long. Legs extensively darkened. Late summer and autumn species 33 Abdominal pubescence black; acr long and bristle-like, dc even longer .... 34 33(32) 6 : bt\ long and cylindrical, rather slender, clearly shorter and only sligh- tly stouter than fj. Scutum light grey with and indefinite brownish cen- tral stripe, abdomen with pale hairs on basal segments only. 9 : very light grey dusted, abdomen silvery-grey, scutum with 2 narrow darker lines between rows of bristles; f 3 slightly compressed and curved 32. H. liiorea (Fallen) S : bt] conspicuously large and swollen, as long as tibia. Abdomen uni- formly covered with white hairs, also hind-marginal bristles whitish. 9 : rather greyish species, abdomen not silvery; ? 3 simple and slender (Alps) (see also couplet 74) 34. H. sartor Becker 34(32) 6: bty and t\ dorsally with long black bristles, bt\ with 5-7 strong dorsal bristles longer than basitarsus is deep; wings faintly brownish-grey clouded. 9 : fj and f 3 dorsally with black bristles at least as long as tibiae are deep, r 3 long and slender, very indistinctly curved at middle. Palpi yellow to brownish-yellow, covered with black bristly hairs, 3 ntpl bristles 17. H. lasiopa Strobl â : bt ] with only short fine hairs, no distinct bristles. 9 : f j bare except for a small hair-like bristle in basal third above and a preapical bristle, / 3 with fine bristles dorsally, slightly compressed and distinctly curved. 1 strong ntpl bristle (in addition with 2-3 smaller bristly hairs) 35 35(34) Palpi yellowish to light brownish, covered with fine pale hairs; haltère with blackish-grey knob, stalk paler. Abdomen subshining black, almost H1LARA FROM SWITZERLAND 521 shining brown on dorsum in female. S : sternum 8 with a fan of unusual long black bristles posteriorly (as in H. manicata) 19. H. hyposeta Straka Palpi dark grey to greyish-brown, covered with black hairs; haltère almost uniformly brown or yellowish (see also couplet 23). Abdomen greyish dusted, rather light greyish pollinose in female. S: sternum 8 with short bristly hairs along hind margin, without distinct bristles .... 18. H. eviana Straka 36(28) acr irregularly 3- to 4-serial or more 37 acr regularly 2-serial; if almost 3-serial posteriorly (H. pseudosartrix), then abdominal pubescence pale 41 37(36) Scutum shining black, or at least subshining; acr irregularly 3- to 4- serial. bt^ in both sexes (less distinct in 9 ) with 2 anterodorsal bristles at tip. Generally smaller species, body about 3 mm long 38 Scutum dusted, dull grey to blackish-grey, not at all shining; if in doubt (H. campinosensis), then frons and face narrowed, legs extensively dark, and generally larger species, body 3-4 mm; acr 4-serial or more 39 38(37) Legs extensively yellow with dark tarsi; haltère with yellow stem and blackish knob; palpi yellowish-brown. Wings slightly smoky in male, almost clear in female; acr short and fine, diverging. S : t\ dorsally with fine hairs, bty slender, half as long as length of tibia, and only slightly wider; genitalia with an unusually long, slender hypandrium bowed above lamellae (like in H. pseudosartrix). $ : ? 3 simple and slender. Larger species, body 2.5 - 3 mm long 36. H. morata Collin Legs extensively blackish, at most cxy, base of all femora and all "knees" yellowish; haltère entirely black; palpi blackish. Wings strongly darkened in both sexes; acr very long, bristle-like, and widely spaced. cT: t\ long bristled before tip, bty short ovate, blunt-tipped, not much longer than half-length of tibia. $ : ? 3 slightly compressed and bent. Smaller species, body up to 2.5 mm long (see also couplet 102) 23. H. merula Collin 39(37) Large species, body 4 - 5.5 mm long, abdomen often yellowish at base. Palpi yellow, acr 4- to 6-serial. 6 : bt\ only slightly thickened, not much stouter than tibia at tip; 9 : r 3 simple and slender 68. H. lurida (Fallen) Smaller species, body 2.5 - 4 mm long, abdomen uniformly blackish. acr almost regularly 4-serial; â bt\ clearly thickened, stouter than tibia at tip 40 40(39) Generally smaller, body about 2.5 - 3 mm long. Palpi yellowish at tip, wings brownish. Legs extensively blackish, only cxy and base of /j yellowish, bt^ in both sexes with several distinct anteroventral bristles much longer than tarsomere is deep. Frons deep black above antennae, and contrasting with light grey face, very narrow, as wide as anterior ocellus. 6: bt\ short ovate, half as long as tibia. 9: t 3 slender but dis- tinctly curved (see also couplets 94, 102) 37. H. splendida Straka Larger species, body about 3-4 mm long. Palpi blackish, wings very faintly brownish infuscated. Legs varying in colour from almost 522 M - CHVÂLA & B. MERZ uniformly blackish to extensively yellowish on fore coxae, base of femora, and "knees"; bt 3 covered with short hairs. Frons black, widening into a triangle above, with a shining patch above antennae, face as wide as 2nd antennal segment is deep, â : bt\ not much stouter than tibia and about two-thirds as long. 9 : f 3 unmodified, slender as in male (see also couplet 105) 63. H. campino sensis Niesiolowski 41(36) Large species, body about 4 - 5.5 mm long. S: fore tibia and tarsus ciliated dorsally with long black bristles; 9 : abdomen with a tuft of long brownish bristly hairs on each side of tergum 8 (group 3, H. clavipes- group, part) 42 Smaller species, body at most 3.5 mm long. S: fore leg with short bristling; 9 : abdomen without a tuft of long hairs at sides of tergum 8 43 42(41)Prothoracic spiracle pale, yellowish-brown. Legs extensively yellowish to brownish-yellow; haltère yellowish, with knob at most brownish, and wings almost clear. S: f] and bt ] long bristled dorsally, following tarso- meres with short hairs only 10. H. cilipes Meigen Prothoracic spiracle blackish. Legs extensively blackish, except for yellowish cx\ and base of^; knob of haltère dark brown to blackish 6 : in addition to t\ and bt\, also following 2 tarsomeres on fore leg long black bristled 12. H. curtisi Collin 43(41) Palpi black; scutum subshining black, thoracic pleura dull grey, wings deep brown clouded. Legs extensively blackish, only cjq and base of/j yellowish. Small species of the H. chorica-group with short tarsi. 6: bt\ very thickened, short ovate; 9 : t% rather slender and distinctly curved. Small, body about 2.5 - 3 mm long (see also couplet 136) 98. H. aartseni Chvâla (To this couplet comes also Hilara flavocoxa Straka, a species known so far only from highlands of central Slovakia. A small-sized species of the H. canescens-group, body about 2-3 mm long, closely allied to H. menda. It differs by long, bristle-like 2-serial acr (short in H. aartseni), and by the unusually enlarged male bt h which is about as long as corresponding tibia (much shorter in H. aartseni, at most 1.5 times as long as wide); bt^ in both sexes with 2 distinct anterodorsal setae before tip. For redescription with illustration see "Additional species".) Palpi yellowish (brownish in H. cuneata and H. cinereomicans), dorsum of thorax dulled by greyish dust, not shining. Fore tarsi fairly long, tarsomeres 2-4 clearly longer than deep 44 44(43) Abdominal pubescence and hairs on cx^ pale. Small, body about 2.5 - 3.3 mm long, light grey dusted species, scutum with velvety brown patches posteriorly in â , and with 2 darker narrow lines between rows of bristles in 9 . Legs extensively dark, rather uniformly dark brown to brownish-yellow, very slender; acr with a tendency to be 3- to 4-serial posteriorly. S : bt x very large and stout, almost as long as tibia; genitalia with an unusual long, slender hypandrium bowed above lateral lamellae (as in H. morata). 9 : legs extensively blackish, f 3 simple and slender (see also couplets 89, 125) 33. H. pseudosartrix Strobl HILARA FROM SWITZERLAND 523 Abdominal pubescence and hairs on oq black; acr regularly 2-serial, long and bristle-like, male bt\ clearly shorter than tibia (except for H. cuneata) 45 45(44) Legs extensively blackish, often only cx x and base of /j yellowish. Scutum dark grey, slightly dark brownish pollinose, subshining, pleura lighter grey; acr in two close rows. S : bt 2 with a distinct thickening at base beneath. 9 : t$ slightly compressed, as deep as femur 46 Legs yellow on coxae and femora, but tibiae and tarsi more or less darkened. Thorax uniformly finely dark grey dusted on scutum, â : bt 2 simple, without the basal swelling 47 46(45) acr distinctly diverging; antennal style slightly longer than 3rd segment. 6: sternum 8 with unusual long bristles posteriorly forming a fan of black bristles on each side of genitalia and overlapping the lamellae posteriorly (as in H. hyposeta). $ : wings clear; £ 3 slightly curved at middle 20. H. manicata Meigen acr only slightly diverging; antennal style very long, 1 .5 times as long as 3rd segment. S sternum 8 with short bristles, not forming a fan of bristles at sides of hypopygium. 9 : wings brownish; r 3 slightly laterally compressed, almost straight 21. H. ponti Chvâla 47(45) acr narrowly biserial and very conspicuously diverging. S : Frons above antennae and face very narrowed, not as deep as 2nd antennal segment. Tibiae and basitarsi on anterior two pairs very densely, long black pubescent dorsally; bt\ not much shorter than fj and almost equally deep. $ : f 3 long and slender (and haltère whitish) 15. H. cuneata Loew acr in 2 widely separated rows, not clearly diverging. S : Frons and face of usual width, and anterior four tibiae and basitarsi with usual short bristling. Thorax uniformly finely dark grey dusted on scutum and pleura, the latter somewhat translucent brownish, and humeri (post- pronotum) yellowish. Genitalia large, especially hypandrium laterally flattened, keel-like (as in more northern distributed H. opta Collin). $ unknown 22. H. cinereomicans Strobl 48(17) Haltère pale, clear yellow, or at least uniformly yellowish-brown; if in doubt (for instance in H. platyura), the species will be found in both sections 49 Haltère black, at least knob blackish-grey, the stem may be paler 72 49(48) Scutum unstriped, uniformly coloured, or longitudinal stripes very indistinct (if in doubts, e. g., H. lacteipennis), species with milky-white wings, and dark median stripe, belong here) 50 Scutum with 2 to 4 distinct dark stripes on the grey background, or scutum velvety black in some lights, with stripes that change from different points of view (group 6, H. intermedia-group, part) 60 50(49) acr 4-serial 51 acr 2-serial 54 524 M. CHVÂLA & B. MERZ 51(50) Frons and occiput dull grey, occiput not black from any point of view. Smaller species, body about 2.5 - 3.5 mm long, scutum uniformly light grey dusted 52 Occiput dull black, at least in some lights; large species, body 4-6 mm long (if wings milky-white, and a smaller species, 2.5 - 3.5 mm long, see section 55) 53 52(5 1) Larger species, body 3-3.5 mm long, haltère brownish. Palpi brownish- yellow; wings very clear with indistinct veins, stigma invisible, radial fork acute. Scutum light grey dusted, unstriped. Abdomen rather sub- shining blackish-brown, covered with dense dark pubescence, and long, thin black hind-marginal bristles, â : bt\ long ovate, not much stouter than tibia, clearly longer than rest of tarsus, and tarsomeres 2-4 long, twice as long as deep; genitalia unusually large, hypandrium very convex; 9 : f 3 slightly dilated and indistinctly curved 39. H. platyura Loew Small species, body about 2.5 mm long, haltère clear yellow. Palpi black, greyish dusted; wings clear with distinct blackish veins, stigma faintly brownish, and radial fork of usual "hilarine" shape. Scutum slate- grey dusted, with 2 indistinct darker lines between small, numerous widely spaced acr and dc bristles. Abdominal pubescence rather long and fine, pale, hind-marginal bristles not differentiated. 6 : bt x stout and not much shorter than tibia, clearly longer than rest of tarsus, and tarsomeres 2-4 only slightly longer than deep; genitalia, especially hypandrium, small. 9 : f 3 slender and unmodified 40. H. leukensis sp. n. 53(51) Larger species, body 5-6 mm long. Frons dull black, narrow in S, as deep as 2nd antennal segment, face greyish. Scutum dark grey, with a broad, darker brownish median stripe on the widely spaced 4- to 6-serial acr, dc 1-serial. Palpi brownish at tip, frontal bristles reduced. 6: bt\ long cylindrical, longer than rest of tarsus; 9 : ? 3 simple and slender (in the morphologically similar H. aeronetha dc more than 1-serial, and haltère blackish, see couplet 90) 79. H. angustifrons Strobl Smaller, body about 4-5 mm long. Frons and face grey, of usual width in both sexes. Lighter grey dusted species with whitish pubescence, dc 2- to 3-serial, and legs silvery pilose. Haltère often darkened (H. bore- alis-group, part) (see also couplet 85) . . 93. H. pruinosa Wiedemann in Meigen 54(50) Wings milky-white, veins whitish, indistinct. Scutum light grey to silvery-grey with 2 indistinct narrow darker lines between rows of bristles, acr and dc very small, hair-like. Very light grey dusted species, abdominal pubescence whitish 55 Wings clear or clouded, veins always more or less darkened and distinct. Small to medium-sized species, body about 2.5 - 3.5 mm long 57 55(54) Vein Sc incomplete, not reaching costa. Scutum silvery-grey, acr 2- serial anteriorly, more numerous behind. Legs blackish, with "knees" and bt 3 pale, all femora whitish pilose beneath, / 3 slightly thickened and with dark ventral bristly hairs. Palpi dark greyish, body 2.5 - 3.5 mm long, (see also couplets 57, 86, 124) 116. H. albitarsis von Roser 9 HILARA FROM SWITZERLAND 525 Wings whitish with vein Sc complete, reaching costa. Combination of characters different. / 3 not thickened and spined below, acr regularly 2-serial 56 56(55) Scutum light bluish-grey dusted, with 2 indistinct narrow darker lines between rows of bristles. Legs quite slender, more uniformly brownish, "knees" narrowly paler; femora not silvery pilose beneath. Palpi yellowish. 9 : £ 3 simple and slender (body 2 - 2.5 mm) (H. litorea- group) 43. H. albipennis von Roser Scutum very light grey dusted, acr 2-serial, on a broad brown median stripe; dc 1 -serial, on very narrow dark lines. Haltère extensively darkened, palpi greyish-brown. 9 r 3 evenly thickened and curved (body 2.7 - 4 mm) (â unknown, H. borealis-group) ... 92. H. lacteipennis Strobl 2 57(54) Occiput dull black in some lights, frons and vertex light grey. Vein Sc incomplete. All femora silvery pilose beneath, / 3 distinctly thickened and with long, black, spine-like bristles beneath, hind tarsi often pale. Scutum light grey dusted, with 3 indefinite blackish stripes visible in some lights; acr 2-serial anteriorly, often 3- to 4-serial posteriorly. Abdominal pubescence whitish (see also couplets 55, 86, 124) 116. H. albitarsis von Roser S Frons and occiput dull grey, occiput not dull black from any point of view. Vein Sc complete, reaching costa. Legs blackish, all femora and tibiae long and slender, almost bare, and femora not silvery pilose beneath. Scutum dark grey dusted, more brownish-grey in anterior view; 2-serial acr widely spaced, dc 1 -serial, all bristle-like and few in number, about 8 to 9 setae in one row, and all as long as antennal style. Uniformly rather light grey dusted species (H. griseola-complex) 58 58(57) Abdominal pubescence, and hairs on lower part of occiput black. Scutum with 2 very narrow, fine, darker blackish-grey lines between acr and dc. Wings clear. 9 : ? 3 slightly dilated and curved 59 Abdominal pubescence and hairs on lower part of occiput whitish. Scutum in anterodorsal view with a faint, indistinct, darker grey median stripe on acr. Wings brownish clouded. 9 : £ 3 simple and slender, unmodified 29. H. crossleyi sp. n. 59(58) Palpi black; legs uniformly black in ground colour, finely greyish Pollinose. Rather a medium-sized species, body length varies from 2.5 to 3.5 mm H.H. griseola Zetterstedt Palpi yellow; legs black in ground colour, but cxy towards tip and all femora at least at base translucent brownish. Generally smaller species, body about 2.5 - 2.9 mm long 28. H. pianti sp. n. 60(49) Occiput dull velvety black, at most with a contrasting whitish-grey triangle behind ocelli (H. fuscipes) 61 Occiput dull grey, at most with an indefinite darker blackish-grey shadow above neck; if in doubt, (i.e. H. luteihalterata, a small species with short stout antennal style) the species is in both sections. Abdo- minal pubescence always pale, at least on basal segments 67 526 M- CHVÀLA & B. MERZ 61(60) Occiput with a contrasting whitish-grey triangle behind ocelli. Body varying in length from 3 to 5 mm, /j with 1 or 2 small dark bristles anteriorly at middle, a: hypandrium narrowed posteriorly, well sepa- rated from lateral lamellae; 9 : r 3 evenly thickened and slightly curved 49. H. fuscipes (Fabricius) Vertex and occiput without the contrasting light grey triangular patch above 62 62(61) acr 4-serial, abdominal pubescence light brownish. Scutum brownish- grey in anterior view, light grey between 2 narrow blackish median stripes, broad lateral stripes visible in dorsal view; all stripes coalescent at sides when viewed from above and behind, leaving a light grey central stripe, sides of scutum then broadly black. 6 : legs covered with only short fine hairs and bristles, bt x not much stouter than tibia; $ : f 3 rather slender but strongly curved. Species varying in length from 2.5 to 5.5 mm 48. H. beckeri Strobl acr 2-serial, and scutum with a different black pattern. Legs in cT, including tibiae, covered with long bristly hairs (except for H. embar- taki), and bt\ very thickened 63 63(62) Abdominal pubescence whitish, at least on basal three segments 64 Abdominal pubescence as well as all hairs and setae on legs black 66 64(63) Legs covered mostly with whitish hairs and setae; anterior four tibiae densely long pubescent posteriorly. Antennae with a short, stout style which is half as long as segment 3. Scutum with distinct black stripes, the 2 median stripes between lines of acr and dc coalescent in posterior view, forming a single central stripe lying on acr. Early spring species; smaller, body about 2.3 - 3 mm long (see also couplet 68) 54. H. lute ihalte rata sp. n. Legs covered with black hairs and setae. Antennal style longer and slender, about two-thirds length of segment 3. Scutum with 2 or 4 dis- tinct black stripes clearly visible as well when viewed from behind. Generally larger species, body about 2.8 - 3.3 mm long 65 65(64) Scutum with 2 distinct black stripes between acr and dc; about 16 acr setae in one row. cT: anterior four tibiae with only short sparse pu- bescence posteriorly; t\ dorsally with 3 to 4 long, fine single bristles. Apical circular process on lateral genital lamella spinose along margin only. Wings short, 2.9 - 3.3 mm long. A summer species in mountains (French Massif Central) 55. H. embartaki Chvâla Scutum with 4 distinct black stripes, the lateral 2 shorter and wider when viewed from behind; acr less numerous, about 10 setae in one row. 6: anterior four tibiae with a dense pubescence posteriorly at least as long as tibiae are deep; t± dorsally with about 6 pairs of long bristles. Apical circular process of left lamella distinctly spinose also ventrally. Wings larger, 4.2 - 4.5 mm long. Early spring species (Bulgarian Rhodope Mts) 56. H. polleti sp. n. HILARA FROM SWITZERLAND 527 66(63) A medium-sized species, body about 3-4 mm long. Thoracic stripes not well visible, scutum light brownish-grey in anterior view, almost uni- formly velvety black from other angles, with a trace of golden-brown stripes between lines of bristles, leaving central stripe on narrowly 2-serial acr bristles black. Labrum short, half as long as head is high. S : anterior four femora and tibiae dorsally with very long woolly pu- bescence, no distinct bristles; bt\ strongly swollen, also t 2 and bh thickened. Genitalia with hypandrium not narrowed or produced distally. $ : tibiae short pubescent, no distinct bristles, t 3 slightly com- pressed and curved, narrower than femur 52. H. pilipes Zetterstedt Small species, body about 2 - 2.8 mm long. The four subshining black stripes on scutum almost invisible when viewed from in front, in posterior view scutum extensively blackish. Labrum long, as long as head is high, â : all tibiae rather long pubescent, also f 3 dorsally, fj with 4 distinct dorsal bristles, and t 2 and bt 2 slender. Genitalia with a long, slender curved hypandrium. 9 : r 3 simple and slender . . 53. H. tanychira Strobl 67(60) Dark stripes on scutum almost invisible in anterior view, when viewed from above with 4 distinct blackish stripes, the 2 median stripes between acr and dc turning to golden-yellow in posterior view and acr then lying on a velvety black middle stripe. Scutum chestnut-brown in dorsal view, with margins, prescutellar depression and scutellum contrasting light grey; acr almost uniformly 2-serial. â : lateral genital lamella with a long pointed apical projection; 9 : t% slender and simple. Later summer and autumn species 46. H. hirta Strobl Scutum uniformly light grey to brownish-grey dusted, with 4 dark stripes clearly visible from all points of view. 9 : ? 3 more or less compressed and curved. Spring and rather early summer species 68 68(67) acr regularly 4-serial, at least at middle, almost 2-serial only in H. quadri/asciata. Large species, generally about 3-5 mm long 69 acr regularly 2-serial. Small species, body about 2.3 - 3 mm long, with anterior four tibiae in S densely long pubescent (?] posteriorly, like/j); antennal style short and stout, frons mostly black, and occiput nearly dull grey when viewed from behind (see also couplet 64) 54. H. luteihalterata sp. n. 69(68) Uniformly light grey to almost whitish-grey dusted species, the bristle on each side of pronotum pale, rarely darkened; cx 2 and cx 3 with uni- formly yellowish hairs and bristles. Frons entirely whitish-grey dusted, not darkened along ocular margins. Ocellar and frontal bristles very fine, much thinner than postvertical (upper postocular) bristles, and usually as long as antennal style, although they vary in length in some extent. Abdominal terga 1-4 with whitish hairs, without dark hind-marginal bristles 70 Darker grey dusted species, scutum more or less darker or brownish- grey pollinose, the bristle on each side of pronotum black; cx 2 and ca 3 with additional blackish bristles. Frons grey, ocular margins with a 528 M. CHVÂLA & B. MERZ velvety brown line touching above hind ocelli. Ocellar and frontal bristles as thick as those in upper postocular row, and much longer than antennal style. Abdominal terga 1-4 with dark hind-marginal bristles, only fine in female 71 70(69) Labrum short, about half as long as head is high. Thoracic stripes blackish, the 2 central stripes between acr and dc narrow and sharply contrasting with the bluish-grey scutum, fj and ? 3 dorsally with several dark bristly hairs clearly longer than the other pubescence; f 2 with a row of long dark anterior bristles. Occiput rather uniformly light grey dusted. 6 : ?2 densely long pubescent anteriorly, the hairs at least as long as tibia is deep; lateral genital lamella with a very narrow, simple, slender, apical projection; 2 : r 3 compressed and very curved 44. H. intermedia (Fallen) Labrum long, as long as head is high. Thoracic stripes broader, brownish and less contrasting; when viewed from in front the distance between the 2 central stripes as wide as the width of one stripe. Legs covered with uniform short pale pubescence, no distinct bristles, except for 1 or 2 short preapical bristles on tibiae, and/ 2 anteriorly with only short pale bristly hairs. Occiput when viewed from above with two large brownish patches above neck, leaving only a triangular light grey patch behind vertex. 6 : t 2 covered with only short hairs, lateral genital lamella with an ovate, spinose, apical projection; $ : f 3 compressed, but only slightly bent, almost straight 45. H. tetragramma Loew 71(69) Scutum clearly grey dusted, acr irregularly 4-serial; cx^ with fine yellowish-brown hairs at tip. Occiput very grey, with two darker, well separated patches above neck. S : t 2 with a few short black spine-like an- teroventral bristles in apical third, otherwise short pubescent; bt 2 slender, as deep as following tarsomeres, and less than half length of tibia; f 3 dorsally with about 4 bristles slightly longer than tibia is deep. 9 : ? 3 distinctly dilated, almost as deep as femur at middle, dorsally with 5-6 bristles longer than other pubescence 50. H. quadrula Chvâla Scutum more brownish in colour, especially between the dark central stripes, acr with a tendency to be 2-serial; cxy with stouter blackish bristles at tip. Occiput more uniformly darker grey dusted. 6: t 2 in addition to the anteroventral spine-like bristles with long bristly hairs anteriorly and posteriorly along the whole length, those in posterior row more numerous, longer and finer; bt 2 stouter and longer, at least half length of tibia; r 3 with about 8 bristles dorsally clearly longer than tibia is deep. $ : r 3 less compressed, clearly narrower than femur, and dorsally with a row of less differentiated, almost equally long hairs and bristles 51. H. quadri/asciata Chvâla 72(48) Occiput dull grey from all points of view, not at all black, also frons and vertex dull grey. Wings clear, abdominal pubescence black (brownish in H. nigrocincta) 73 Occiput dull black, at least when viewed from above, usually frons and vertex also mostly black 78 H1LARA FROM SWITZERLAND 529 73(72) acr 2-serial (at least anteriorly in H. sartor) 74 acr 4-serial; abdominal pubescence always black 75 74(73) acr fairly long, bristle-like, and less numerous. Small species, body 2 - 2.5 mm long, scutum uniformly dull bluish-grey dusted with a brownish pattern, unstriped, abdomen and male genitalia subshining brownish. Abdominal pubescence black, ô: t\ short and thick, bt] unusually large and swollen, clearly longer than tibia; 9 : ? 3 slightly dilated (species of the H. canescens-group) 30. H. tiefii Strobl acr small and numerous, more than 10 setae in one row, and posteriorly often 3-serial. Larger, body 2.5 - 3.5 mm long, thorax and abdomen uni- formly nearly light grey dusted. Abdominal pubescence whitish. S : bt\ very long and swollen, as long as the very slender t j ; 9 : t 3 simple and slender (species of the H. litorea-group) (see also couplet 33) 34. H. sartor Becker 75(73) Large, body about 4 - 4.5 mm long, Scutum dull, dark bluish-grey dusted and densely black bristled, with 3 broad reddish-brown stripes, 6-8 sc bristles; abdomen dull light grey. Antennal style short, half as long as 3rd segment, â : bt\ long ovate, but not much stouter than tibia; / 3 with about 4 strong black anteroventral bristles before tip, ? 3 with 3-4 dorsal bristles longer than tibia is deep, and with 3 strong spine-like bristles anteroventrally. 9 : ? 3 simple, slender, and the dark stripes on scutum more greyish-black (species of the H. interstincta-group) 80. H. caerulescens Oldenberg Smaller species, body at most 3.5 mm long, combination of characters different 76 76(75) Smaller, body about 2.5 mm long. Scutum uniformly brownish-grey dusted, sides narrowly and scutellum contrasting grey. 8: bt\ rather short and stout, about 2/3 as long as length of tibia, the latter short pubescent, dorsally with 3-4 short fine bristles; r3 with small hairs, except for 3-4 dorsal bristles which are about as long as tibia is deep, and similar smaller anteroventral bristes. 9 : f 3 evenly dilated and compres- sed, slightly curved (syn. H. griseifrons Collin) 42. H. ternovensis Strobl Generally larger species, about 3 - 3.5 mm long (rarely 2.6 mm), with a different combination of characters, â : bt x longer and rather cylindrical, about 3 times as long as deep 77 77(76) Slender, long-legged species, oq and femora at base often translucent yellowish; vein Sc usually abbreviated. Scutum brownish-grey, with a darker, almost blackish median stripe. $ : t { with a pair of long preapical setae and a few dorsal setae; ? 3 with 3-4 black dorsal and anteroventral setae at least as long as tibia is deep, other hairs minute. Terga uniformly blackish. Lateral genital lamella deeply cleft, upper process long, finger- like. 9 : f 3 simple and slender (August - October) (species of the H. cor- nicula-group) 65. H. nigrohirta Collin Rather robust, stouter-legged species, legs uniformly black; vein Sc complete. Scutum uniformly slate-grey, with widely spread acr on a 530 M. CHVÄLA & B. MERZ broad lighter grey median stripe, separated by narrow darker grey lines from dc, sides somewhat brownish-grey; prescutellar depression and scutellum almost silvery-grey. Palpi brownish at tip, abdominal pubes- cence brownish, somewhat paler; abdomen when viewed from behind with contrasting black hind tergal margins, â : t\ with numerous bristly hairs dorsally; f 3 anteriorly and anteroventrally with dense, rather adpressed long bristly hairs much longer than tibia is deep. $ : / 3 slender at base, strongly dilated and curved on apical two-third (May - July) (species of the H. litorea-group) 41. H. nigrocincta de Meijere 78(72) acr 4-serial or more; if irregularly 2- to 3-serial, then large species about 5-7 mm long (H. lugubris), or abdominal pubescence pale (H. pseudo- sartrix, H. longiocornis) 79 acr regularly 2-serial; if more than 2-serial (H. biseta of the H. chorica- group), then small species about 2 mm long, with short tarsi and sub- shining scutum 106 79(78) Wings milky-white, with indistinct pale venation. Scutum and abdomen bluish-grey; acr small, hair-like and numerous, black; abdominal pu- bescence blackish. Frontal bristles absent, palpi blackish to yellowish- brown. 6: bt\ long cylindrical, short pubescent and longer than rest of tarsus; lateral genital lamella with an unusually long, slender, pointed terminal process. 9 : ? 3 simple and slender 35. H. galactoptera Strobl Wings more or less clouded, or clear, but then with distinct blackish venation; wings not at all whitish 80 80(79) Scutum dull grey to greyish-black or greyish-brown, not at all sub- shining, often with more or less distinct darker stripes, or velvety black in some lights 81 Scutum more or less shining black, or subshining blackish-grey, not dull; if thinly greyish or brownish dusted in anterior view (H. angloda- nica, H. pseudocornicula), the species will be found in both sections 97 81(80)/ 2 spinose beneath, with a double row of spine-like bristles towards tip. Large, strongly black bristled species about 5-7 mm long of the H. inter- stincta-group 82 f 2 without black ventral spine-like bristles, at most anteroventrally with fine, bristly hairs 83 82(81) Smaller, body about 5 mm long, wings faintly greyish-brown infuscated, almost clear in 6. Scutum darker grey dusted, the 3 stripes rather brownish and less distinct, abdomen subshining greyish-black. Antennal style shorter, 3/4 length of 3rd segment. Labrum slightly shorter than head is high. Legs extensively blackish including "knees". 6: / 3 pubescent beneath, with 2-3 anteroventral preapical bristles only; Ç : / 3 slightly compressed an bent 72. H. interstincta (Fallen) Larger species, body 6-7 mm long, wings strongly darkened. Scutum with blackish stripes on a grey ground-colour, abdomen dull black. Antennal style long, as long as, or longer than 3rd segment. Labrum at least as long as head is high. Legs black, "knees" yellowish. S: / 3 HILARA FROM SWITZERLAND 531 strongly black bristled beneath; 9 : t^ slender and simple 73. H. lugubris (Zetterstedt) 83(81) Femora more or less microscopically silvery pilose beneath 84 No silver pile or whitish microscopic pubescence on femora beneath; if in doubt, the species will be found in both sections 89 84(83) Coxae with pale hairs, and abdominal pubescence mostly pale, at least at base of abdomen 85 Coxae with black hairs, abdominal pubescence black. Scutum dull bronze brown with 3 black stripes visible from in front, but in posterior view scutum uniformly velvety black. Legs, palpus and haltère quite black. Frontal bristles indistinct or absent; antennal style short, often only of half-length of 3rd segment, o* : bty short cylindrical, scarcely wider than tibia at tip, / 3 spindle-shaped and strongly black bristled beneath; 9 : / 3 simple and slender. Medium-sized species, body about 3-4 mm long 60. H. coracina Oldenberg 85(84) Larger, about 4-5 mm long, densely light silvery-grey dusted species. Scutum light grey in frontal view; when viewed from above with a narrow darker median line, and sides broadly blackish; acr narrowly 4- serial, very small and diverging, dc 2- to 3-serial. Antennal style short, half as long as 3rd segment. Frons and face light grey, humeral and frontal bristles absent, palpi greyish-black. Fore tibia and tarsus dense- ly, almost golden-yellow to silvery microscopically pubescent; wings brownish clouded. 9 : f 3 simple and slender (see also couplet 53) 93. H. pruinosa Wiedemann in Meigen Smaller, generally about 3 mm long (2.5 - 4 mm), darker grey species (except for H. albitarsis 9), with acr not diverging and dc 1 -serial. Antennal style longer, at least 2/3 length of 3rd segment; if shorter (H. albitarsis), then wings clear and vein Sc abbreviated 86 86(85) Wings clear with vein Sc abbreviated, acr with a tendency to be 2-serial, minute. Haltère sometimes yellowish and occiput often greyish, very variable species in many respects. 6 : bt^ short elliptical, slender, scar- cely wider than tibia at tip;/ 3 spindle-shaped, thickened, with long black spine-like bristles beneath. Legs with broadly yellow "knees" and base of tarsi (see also couplets 55, 57, 124). 9: thorax and abdomen very light grey dusted, legs simple and darker 116. H. albitarsis von Roser Wings with vein Sc complete, reaching costa (sometimes abbreviated in H. bohemica, but then wings brownish). S : / 3 not dilated; if slightly thickened (H. bohemica), then ventral bristles very short. 9 : body dark, dull brown or subshining black; if abdomen light grey (H. bohemica), then scutum with contrasting brown stripes. If / 3 dilated and black bristled ventrally (H. coracina), then all hairs and bristles black, and scutum velvety black when viewed from behind 87 87(86) acr narrowly 2- to 3-serial. Scutum silvery-grey pollinose in anterior view, with narrow reddish-brown stripes on lines of bristles, but when viewed from above subshining black; h bristle small and fine, or absent. 532 M. CHVÄLA & B. MERZ Frons dull black in some lights. Antennal style about 2/3 length of 3rd segment; wings faintly brownish. S : fy distinctly spindle-shaped, ven- trally with short black bristles (much shorter than in H. albitarsis and finer than in H. coracina); bt\ only slightly swollen, scarcely wider than tibia at tip; abdomen almost shining black. 9 : dorsum of abdomen sub- shining black, but silvery dusted in some lights; f 3 simple and slender (see also couplet 124) 117.//. bohemica Straka acr widely 4-serial, at least anteriorly, Wings very clear, iridescent, veins blackish. S :f$ not dilated and armed with only fine bristly hairs beneath, bt\ distinctly swollen, much wider than tibia towards tip 88 88(87) Scutum brownish dusted in anterior view, when viewed from above broadly very light bluish-grey at middle (with a narrow median line), in prescutellar depression, and on scutellum. Antennal style long, as long as 3rd segment; frons and face light grey. A distinct, although fine h bristle. Abdomen dull black when viewed from above in male, dull brownish in female. 9 : £3 simple, but fairly stout . 90. H. medeteriformis Collin Scutum light grey in all lights, with a brown or reddish-brown median stripe (mainly males), or with 3 stripes on lines of bristles (often females). Antennal style shorter, nearly half as long as 3rd segment; frons velvety blackish-brown, face light grey. Both h and ih bristles minute, practically absent. Wings with a tendency to be whitish. $ : f 3 simple and very slender 91 . H. calinota Collin 89(83) cjcj with pale hairs. Palpi yellow. Small species about 2.5 - 3 mm long, rather light grey dusted on thorax and abdomen, acr irregular, almost 2- serial anteriorly. Frontal bristles, and h bristle fine, small. Scutum dull light grey, in frontal view with 2 indistinct darker lines between acr and dc, in dorsal view postalar calli and sides of scutellum broadly deep velvety brown or almost dull black. Legs often uniformly brownish (see also couplets 44, 125) 33. H. pseudosartrix Strobl cxj with black bristly hairs, palpi blackish; and with a different combi- nation of characters 90 90(89) Large species, about 6 mm long; acr irregularly 6-serial, dc irregularly 1- to 2-serial. Frontal bristles very small, fine. Antennal style nearly as long as 3rd segment, frons and face in â narrowed (the similar species H. angustifrons has yellow haltère and dc 1 -serial, see couplet 53) 78. H. aeronetha Mik Generally smaller species, body at most 5 mm long; acr irregularly 4- serial, dc 1 -serial. Frontal bristles strong, about as long as ocellar brist- les (finer and smaller in H. splendida, absent in H. coracina) 91 91(90) Labrum long, as long as head is high. Wings clear, at most greyish hyaline . 92 Labium short, clearly shorter than head is high 93 92(91) Smaller species, about 2.5 - 3.5 mm long, finely bristled, thorax un- striped. Scutum brownish-black dusted, though subshining in some lights. Legs with a tendency to be uniformly dark brownish. Labrum long, slender, abdomen subshining black to blackish-brown. 6 : / 3 with HILARA FROM SWITZERLAND 533 4-5 long anteroventral bristly hairs on apical half much longer than femur is deep; lateral genital lamella deeply cleft apically, and the ter- minal process broad, hypandrium short bifurcated. 9 : ? 3 unusually dila- ted on apical third and strongly curved (syn. H. subpollinosa Collin) (see also couplet 97) 66. H. pseudocornicula Strobl Generally larger species, body length varies from 3 to 5 mm, robust, and strongly bristled. Scutum dull brownish-grey, in dorsal and posterior views with 3 velvety black shifting stripes. Labrum unusually strong, abdomen dull grey. â'.fy with the usual anteroventral preapical bristles; lateral genital lamella without any process at the blunt apex, hypandrium apically long bifurcated. 9 : ? 3 evenly distinctly dilated and compressed, only slightly curved 75. H. scrobiculata Loew 93(91) Generally smaller, body about 2.5 - 3.5 mm long; acr and dc rather long and bristle-like, acr regularly 4-serial and widely spaced 94 (Here belongs also H. longicornis Strobl, a species known so far from the Hungarian lowlands of the Danube region. A very small species, body about 2 - 2.8 mm long, with unusually long antennae, and simple unmodified legs in both sexes, even male bt\ long and slender; acr setae are 4-serial at middle only, 2-serial anteriorly and posteriorly, small and numerous. A species of the H. litorea-group, for redescription with illus- tration see "Additional species".) Generally larger species, body about 3.5 - 5 mm long 95 94(93) Frons above antennae and face very narrow, as wide as anterior ocellus; face light grey, contrasting with the blackish frons. Legs with a tendency to be yellow on cx\ and base of /j (see also couplets 40, 102), bt^ with long black anterodorsal bristles longer than tarsomere is deep. Palpi yellowish, at least at tip; antennal style longer than 3rd segment. Wings brownish, haltère with yellowish stem. Scutum dark brownish-grey dusted, almost black in posterior view. S : bt^ long ovate but small, not much longer than half-length of tibia; 9 : ? 3 slender and clearly bent 37. H. splendida Straka Frons and face wide as usual, uniformly velvety black to greyish-black. Legs uniformly blackish, bt 3 with short hairs. Palpi black; antennal style slightly shorter than 3rd segment. Wings clear (almost whitish in some lights), haltère black. Scutum dull brownish-grey in anterior and dorsal views, when viewed from behind with 3 wide blackish stripes leaving space between bristles grey; acr and dc sparse, long and bristle-like, â: bt\ long ovate, not much shorter than tibia is long; / 2 an d/3 before tip with very long anteroventral bristly hairs much longer than femora are deep; / 2 ventrally at base and ? 3 dorsally at tip with an unusual long bristle; t j with a pair of long preapical bristles, a small bristle dorsally in basal quarter, and posteriorly with longer pubescence which is present also on /j. 9: ? 3 evenly dilated and laterally compressed, strongly curved; acr long and regularly 4-serial 67. H. quadrifaria Strobl 95(93) Abdomen shining or subshining black from most points of view. Wings clear, squama with pale fringes; haltère with yellowish stem. Frons 534 M. CHVÂLA & B. MERZ uniformly dull, with a light grey patch above antennae; t 2 with 2-3 short antero- and posteroventral bristles on apical half, and with apical circlet of similar bristles; no anterior basal bristle. S : bt\ slender, only slightly wider than tibia at tip, everywhere with minute hairs, no distinct bristles. Palpi strongly bristled, pronotum with additional small spines in addi- tion to the long lateral bristle. $ : f 3 slender on basal half, then very dilated towards tip (resembling H. nigrocincta or the lowland H. ni- grina) 74. H. dimidiata Strobl Abdomen dulled by greyish dust, squama with black fringes (pale in H. scrobiculata and H. dimidiata); haltère with dark stem. Pronotum with a single bristle on each side 96 96(95) Abdomen rather silvery-grey pollinose when viewed from in front, dull blackish from other angles, scarcely subshining, with distinct hind- marginal bristles. Wings brownish. Frons with a rhomboid subshining patch above antennae; t 2 with a species-specific long anterior bristle at base, another in apical third anteroventrally, and a circlet of similar long preapical bristles. S : bt\ long cylindrical, nearly twice as wide as tibia at tip, dorsally with 1 or 2 distinct black bristles and a smaller preapical pair; tarsi long and slender, tarsomeres clearly longer than deep. Palpi often brownish or nearly yellowish at tip. Hypandrium dorsally before tip with a pair of small flattened spines which is an unique character. 9 : ?3 simple and slender (see also couplet 98) H.H. anglodanica Lundbeck Abdomen dull grey to greyish-brown, no hind-marginal bristles. Wings almost clear. Frons dull velvety black. S: bt\ very stout, long black bristled dorsally, similar long dorsal bristles also on the stout t x \ tarsi short and stout, tarsomeres 2-4 on fore leg as long as deep. Palpi blackish. 9 unknown 76. H. sirbitzmatrona sp. n. 97(80) Labrum long, as long as head is high. Rather a slender, long-legged species about 2.5 - 3 mm long. Wings clear, scutum finely brownish- black dusted, unstriped, subshining in some lights. S: bt\ cylindrical, not much stouter than tibia; $ : f 3 unusually dilated on apical third and distinctly curved (syn. H. subpollinosa Collin) (see also couplet 92) . . . 66. H. pseudocornicula Strobl Labrum short, clearly shorter than head is high 98 98(97) Thorax and abdomen greyish dusted when viewed from in front, scutum in posterior view with 3 indefinite blackish stripes. Face and a patch on frons above antennae grey. Palpi yellowish; antennal style as long as 3rd antennal segment, o" : bt\ long cylindrical, 4 times as long as deep and not much wider than tibia; r 3 with strong dorsal and anteroventral bristles in both sexes. 9 : r 3 slender and simple. Larger species, body about 3.5 - 4.5 mm long (see also couplet 96) . . 77. H. anglodanica Lundbeck Dorsum of thorax black, more or less shining, or covered with a faint greyish dusting. Frons dull black, without a grey patch below, or uniformly grey (in H. pygialis); other combination of characters different 99 H1LARA FROM SWITZERLAND 535 99(98) Abdominal pubescence whitish. Frons and face densely light grey dusted, occiput black, behind vertex with a more or less visible light grey triangular patch. Scutum metallic, finely brownish-grey dusted, subshining, wings brownish. S : genitalia unusually large, hypandrium laterally flattened, largely circular, much higher than end of abdomen. 9 : r 3 thickened, as deep as femur, and curved (Spain, France) (species of the H. cornicula-group) 64. H. pygialis Chvâla Abdominal pubescence black. Frons dull black; with a different combi- nation of characters 100 100(99) acr long, as long as dc, bristle-like, regularly 4-serial and widely spaced, not clearly separated from dc; the distance between acr rows as wide as the distance between acr and dc. Antennal style long and slender, longer than 3rd segment, or at least as long. Abdomen dull. Wings strongly darkened (lighter brownish in H. splendida) 101 acr short and closer together, well separated from dc, and often 2-serial anteriorly. Abdomen shining. Species generally about 3-4 mm long 103 101(100) Legs and haltère uniformly black; a rather small species, body about 2.5 mm long. Scutum uniformly subshining black, palpi strongly black bristled, including 1 very long bristle. t 2 with an anterodorsal bristle in basal quarter, bt^ with a long black anterior bristle before tip, clearly longer than tarsomere is deep in male, shorter in female. S : bt\ short ovate, about twice as long as deep, and nearly of half-length of tibia; $ : r 3 simple and slender 25. H. nigrita Chvâla (The lowland European H. nigrina (Fallen) is a larger species, body about 3-4 mm long, with bt x in cT long cylindrical, not much shorter than tibia, and f 3 in 9 is unusually dilated and curved on apical two- thirds, very slender at base, very much like as in H. nigrocincta. A new record for H. nigrina is listed here: France: Gard, Dourbies (855 m) 21.vii.2009, let, leg. Haenni.) Legs paler, exj, base of/j and "knees" more or less yellowish (see also couplets 17ff) 102 102(101) Frons (above antennae) and face very narrow, about as wide as anterior ocellus, face contrasting light grey. Palpi yellowish, at least at tip; haltère with base of stem yellow, bt^ with several long anteroventral bristles clearly longer than tarsomere is deep. Larger species, body about 3 mm long (see also couplets 40, 94) 37. H. splendida Straka Frons and face blackish, and as wide as usual. Palpi and haltère blackish. bt^ with 2 fine anteroventral bristles before tip. Generally smaller, body about 2.5 mm long (see also couplet 38) 23. H. merula Collin 103(100) h, ih and ph bristles absent, abdominal terga without hind-marginal bristles, pronotum with a distinct bristle on each side. Legs often with cxi, base of/} and "knees" more or less yellowish; c.Vj with pale hairs. acr 4-serial, dc 1 -serial, all minute. Medium-sized, body about 3-3.5 mm long (species of the H. maura-group) (see also couplet 13) 115.//. media Collin 536 M - CHVÂLA & B. MERZ h, ih and ph bristles distinct, although sometimes fine; abdomen with well developed hind-marginal bristles. Legs uniformly black, but if fore leg translucent yellowish at base, then cxy with black hairs 104 104(103) Wings strongly darkened, dark brown, legs entirely black; acr ante- riorly 2- to 3-serial, dc slightly longer. S: genitalia with a distinctly produced, very convex, keel-like hypandrium. 9: t\ with 1 or 2 bristles dorsally, f 3 with rather long anteroventral bristles clearly longer than tibia is deep 61 . H. cornicula Loew Wings almost clear or only faintly brownish clouded, legs with a tendency to be paler at least on cx^, acr almost regularly 4-serial and as long as dc. 6 : genitalia with hypandrium not enlarged and not conspi- cuously produced distally 105 105(104) Generally smaller, body about 2.5 - 3.5 mm long. Haltère black with brownish stem. S : wings clear; genitalia: lateral lamella simple, apical process short and blunt, dorsally with uniform rather short hairs pointing backwards (syn. H. monedula Collin) 62. H. longifurca Strobl Generally larger, body about 3-4 mm long. Haltère with a black knob and contrasting light yellow stem. 6: wings slightly brownish infus- cated, not clear; genitalia: lateral lamella with a long pointed terminal process, at base with a tubercle with a tuft of long, forwardly directed long black bristles (about 6), and dorsal bristling pointing backwards, clearly longer and stouter (see also couplet 40) 63. H. campinosensis Niesiolowski 106(78) Scutum distinctly black striped, or the velvety black stripes change in position from different points of view, or dorsum of thorax entirely dull velvety black in some lights. Male fore tarsi often long, tarsomeres 2-4 clearly longer than deep (except for H. brevivittata and some species of the H. anderniattensis-group). Abdominal pubescence blackish, legs, haltère and palpi black 107 Scutum more or less shining black, or only very finely pollinose, not striped; if in doubt, the species with legs not quite black, with shortened tarsi (fore tarsomeres 2-4 as long as deep), abdominal pubescence pale, and/or with femora microscopically pilose beneath (H. bohemica, H. albitarsis, H. borealis) are included here 119 107(106) Scutum in anterodorsal view with 2 distinct black stripes between acr and dc; when viewed from above with 4 black stripes; if median stripes indistinct or partly fused in posterior view (in H. brevivittata and H. bre- vistriata), then lateral stripes very widened and more or less coalesce, whereas in anterior view scutum almost unstriped. 9 : ? 3 more or less compressed and bent; slender in H. brevistriata (species of the H. inter- media-group) 108 Scutum with 3 wide velvety black stripes that change in position in different points of view, but most often scutum extensively dull black, with 2 narrow light brownish stripes between acr and dc when viewed from above and behind. 9 : t 3 unmodified, simple and slender (except for HILARA FROM SWITZERLAND 537 H. zermattensis and H. parvimaior) (high mountain species of group 1 0, H. andermattensis-group) Ill 108(107) Scutum distinctly striped also in anterodorsal view; in dorsal and posterior view with 4 clearly separated well visible stripes. Antennal style short, almost half-length of 3rd segment. Abdomen rather brownish (in H. bistriata greyish pollinose in dorsal view). Tarsomeres on all pairs of legs in male longer than deep. A small, fine h bristle 109 Scutum brownish and almost unstriped when viewed from in front, but with distinct 4 black stripes in anterodorsal and dorsal views, in posterior view the lateral black stripes very broadened, and the 2 median stripes coalescing into a median stripe on acr setae. Antennal style longer, 3/4 length of 3rd segment. A distinct h bristle 110 109(108) f 2 with the usual black bristles anteriorly. S: anterior four tibiae and basitarsi densely long pubescent, bt 2 slightly swollen and clothed with long hairs; t^ with several dorsal bristles much longer than tibia is deep. 9 : f 3 only slightly compressed and curved, clearly narrower than femur is deep 57. H. hirtipes Collin f 2 without the usual row of anterior bristles. S : t 2 and mid tarsus simple, not stouter or compressed, and covered with short hairs only; t\ finely short pubescent, only about 4 fine dorsal bristly hairs longer than tibia is deep;/! densely long pubescent posteriorly, the hairs at least as long as femur is deep. 2 : f 3 very compressed and distinctly curved, at least as deep as femur 58. H. bistriata Zetterstedt 110(108) Generally larger, body about 2.5 - 3.5 mm. Abdomen deep black when viewed from above, more blackish-brown in 9 . S\t\ with 4 pairs of very long dorsal bristly hairs becoming longer towards tip, apical pair more than twice as long as tibia is deep; bt\ very stout, twice as deep as corresponding tibia is wide, and tarsomeres 2-4 short, as long as deep; t 7 with long anterior bristly hairs longer than tibia is deep (as in H. inter- media). Lateral genital lamella with a circular, ventrally spinose apical process (H. brevivittata complex). 9 : £ 3 compressed, almost as deep as femur, slightly curved 59. H. brevivittata Macquart Smaller species, body generally about 2.3 - 2.6 mm. Abdomen dull grey in both sexes. Legs long, all tarsomeres distinctly longer than deep in both sexes. S : ^ with short hairs only, no distinct setae; bt\ long and slender, about as deep as corresponding tibia is wide; t 2 with short hairs. Lateral genital lamella with a simple, bare, slender apical process (H. intermedia complex). 9 : f 3 slender, slightly undulating 47. H. brevistriata sp. n. 111(107) Labrum long, almost as long as or at least not much shorter than head is high 112 Labrum short, clearly shorter than head is high, often of half-length; if in doubt (H. andermattensis), the species is included in both sections 116 112(111) f\ with short hairs posteriorly, without longer bristly hairs. S: fore tarsus short, at least tarsomeres 3-4 as long as deep 113 538 M - CHVÄLA & B. MERZ fi in both sexes, and also ti in 8, densely long pubescent posteriorly, the hairs at least as long as corresponding femora or tibiae are deep, â : fore tarsi vary in length 115 113(112) Frontal bristles as long as ocellars. 6: bt x very swollen, much wider than tibia at tip; at least tergum 6 with distinct hind-marginal bristles 114 Frontal bristles absent or very small and fine. 8: bti only slightly swollen, nearly as wide as tibia at tip; abdomen practically bare on segments 3-6. Medium-sized, body about 3.5 mm long (see also couplet 118) 87. H. andermattensis Strobl 114(113) Large species, body about 3.5 - 4 mm long. Scutum dull velvety black when viewed from above, only scutellum paler, brownish. 8: ?j and bti covered with short hairs dorsally; genitalia small (= H. tatra Niesiolowski, syn. n.) 82. H. maior Strobl Smaller, body about 3 mm long. Scutum dull black, but in posterodorsal view light cupreous-brown dusted between acr and dc and in prescu- tellar depression. cT : f j and bti densely long pubescent dorsally, t j also posteriorly, posteroventrally with 3-4 long bristly hairs more than twice as long as tibia is deep; genitalia very large, half as long as rest of abdomen 81.//. helvetica Chvâla 115(112) Larger species, body about 3 - 3.5 mm long. Scutum uniformly brownish in anterior view, when viewed from above cupreous-brown in prescutellar depression and on 2 stripes between acr and dc. 8 : tarsi short, bti ver y swollen, and tarsomeres 2-4 as long as deep. 2 : squama ash-coloured, with pale fringes; bt^ clearly longer than bt^ / 3 with several long bristly hairs much longer than tibia is deep . 84. H. styriaca Strobl Smaller, body about 2.5 - 3 mm long. Scutum with black stripes on lines of bristles in anterior view, viewed from above uniformly dull velvety black, only scutellum paler. 8 : tarsi rather long, bt j long cylindrical, 3 times as long as deep and scarcely wider than tibia at tip, tarsomeres 2-3 longer than deep. 2 : squama blackish, with black fringes; bti an< ^ bt^ equally long, and f 3 dorsally with a row of almost equal bristles shorter than tibia is deep 85. H. merzi Chvâla 116(1 11) Tarsi short, tarsomeres 2-4 on fore leg in both sexes as long as deep. Face, and a small patch on frons above antennae, light grey; acr and dc small and fine. 8: bti ver y swollen, much wider than corresponding tibia. 2 : r 3 thickened, nearly as wide as corresponding femur, and distinctly curved 117 Tarsi longer, tarsomeres 2-4 on all pairs clearly longer than deep. Frons and face dull black; acr and dc longer and bristle-like. 8 : bti slender, scarcely stouter than corresponding tibia at tip. 9 : r 3 simple and slender . . 118 117(1 16) Thoracic pleura light bluish-grey dusted, contrastingly paler than scu- tum; wings almost clear. Scutum in posterodorsal view with 2 wide, light grey stripes between acr and dc. Larger species, body 3.2 - 3.6 mm long 86. H. zermattensis Chvâla Thorax uniformly dull black, pleura with a greyish tomentum; wings brownish. Scutum uniformly dull velvety black when viewed from H1LARA FROM SWITZERLAND 539 above and behind, only scutellum brownish. Smaller species, body 2.3 - 2.8 mm long 83. H. parvimaior sp. n. 118(116) Smaller, body about 2 - 2.5 mm long. Scutum almost uniformly velvety blackish-brown, light pattern very indistinct. Frontal bristles long, as long as ocellars. Fore tarsi including basitarsus slender, unmo- dified; acr and dc very long, longer than antennal style; palpi long bristled beneath, also abdominal hind-marginal bristles prominent .... 88. 77. simplicipes Strobl Larger, body about 3.5 mm long. Scutum with 3 velvety black stripes on lines of bristles visible from all points of view. Frontal bristles not developed, or only minute. 6: bt\ scarcely stouter than tibia at tip; acr and dc shorter than antennal style, though rather coarse; palpi with a single long preapical bristle in addition to short bristly hairs, and abdominal segments 3-6 almost bare, without hind-marginal bristles (see also couplet 113) 87. H. andermattensis Strobl 1 19(106)/! very thickened, posterior four femora slender; legs entirely black, only the fore "bent knees" yellowish. Wings brownish, long and narrow, axillary lobe little developed. Scutum shining black, dc very long, bristle-like, as long as antennal style and few in number, about 8 bristles in one row; acr small, in 2 close rows and of half length of dc bristles . 31.7/. perversa Oldenberg /l not conspicuously thickened, all femora almost equally stout. Wings with well developed axillary lobe, and axillary excision distinct 120 120(1 19) Tarsi long and slender, tarsomeres 2-4 on fore leg clearly longer than deep 121 Tarsi short, tarsomeres 2-4 on fore leg very short in S , at most as long as deep, scarcely longer in 9 ; if slightly longer (77. biseta of the 77. cho- r/cfl-group), then small species up to 3 mm long, bt\ in o" with 2-4 strong black bristles dorsally, and dorsum of thorax polished black. Generally small, shining black species, body about 2-3 mm long; if larger (77. borealis), then scutum more brownish-grey with 3 indefinite blackish stripes, acr and dc very inconspicuous, and pubescence on c.Vj and base of abdomen pale to rusty-brown 128 121(120) Small, body about 2 - 2.5 mm long; black species with haltère, legs and all setae and hairs black; S : bty very swollen, twice as long as deep . . 122 Larger species, body generally about 2.5 - 4 mm long, combination of characters different 123 122(121) S: bt\ covered with short hairs, without strong bristles; 9: f 3 evenly thickened and distinctly curved. Antennal style slender; wings almost clear, only indistinctly brownish clouded; scutum silvery-grey pollinose when viewed from in front 26. 77. goetzei Chvâla Ô: bt\ dorsally with 3 strong bristles (occasionally with 2 or 4); 9: £ 3 simple and slender. Antennal style unusually thickened; wings brownish, almost dark brown on costal half; when viewed from in front scutum finely dark cupreous-brownish pollinose, with 2 narrow black stripes between lines of bristles 100. triseta Chvâla 540 M- CHVÀLA & B. MERZ 123(121) All femora densely microscopically silvery pilose beneath / 3 more or less thickened, especially in â ; acr often more than 2-serial (species of the H. albitarsis-group, see also couplets 86 and 87) 124 Femora without microscopic silver pilosity beneath 125 124(123) Legs uniformly black. Abdomen subshining black, with a silvery lustre in female. S : / 3 slightly dilated and ventrally with very short, black, spine-like bristles. Vein Sc usually complete, reaching costa (see also couplet 87) 117.//. bohemica Straka Legs with yellowish "knees", often also posterior tibiae and tarsi some- what paler. Abdomen densely light grey dusted, especially in female, a : / 3 distinctly thickened, ventrally with a row of strong black bristles nearly half as long as femur is deep. Vein Sc always incomplete, not reaching costa (see also couplets 55, 57, 86) 116.//. albitarsis von Roser 125(123) cx-y with pale hairs. Scutum and abdomen when seen from above rather light grey dusted; palpi yellowish. Long-legged species, "knees" yellowish, wings clear. 9 : acr usually nearly 4-serial, wings whitish, (see also couplets 44, 89) 33. H. pseudosartrix Strobl cxi with black hairs. Scutum somewhat metallic, finely dark brownish- grey pollinose; acr and dc long and bristle-like, and only about 10 setae in one row; palpi blackish. Legs entirely black (species of the H. lasio- chira-group) 126 126(125) Wings almost clear, very faintly brownish; antennal style very thickened, nearly one-half length of 3rd segment. Thoracic pleura dull grey below on katepisternum and meron (sterno- and hypopleurum) upper half on anepisternum and anepimeron (meso- and pteropleurum) contrasting darker, subshining black. cT : ? 3 dorsally with unusually long bristly hairs at least 3 times as long as tibia is deep, some long, bristly hairs also on t\ towards tip; hind-marginal abdominal bristles very long, sternum 8 with a fan of long bristles overlapping genitalia as in H. hypo- seta and H. manicata (H. canescens-group). Genitalia small, especially hypandrium, as wide as abdomen at tip. 9 : r 3 compressed and narrower than femur, slightly bent, dorsally with a row of 5-6 bristly hairs nearly as long as tibia is deep. Small species, body about 2.5 mm long (syn. H. longesetosa Strobl) 69. H. pilosopectinata Strobl Wings brownish clouded; antennal style slender. Thoracic pleura uni- formly grey dusted including upper half of anepisternum and meron. cT : f 3 with usual hairing and bristling, dorsal and anteroventral setae at most slightly longer than tibia is deep. Abdomen with hind-marginal bristles of the usual length, and no fan-like long setae on sternum 8 pos- teriorly. Genitalia very large, larger than the 6th abdominal segment, at least hypandrium strongly flattened. 9 : f 3 evenly dilated, laterally com- pressed, narrower than femur, and only very slightly bent at middle. Generally larger species, body about 2.5 - 3.5 mm long 127 127(126) Antennal segment 3 long, style two-thirds as long; acr and dc very long, bristle-like, about as long as antennal style, and few in number HILARA FROM SWITZERLAND 541 (about 8 pairs of dc and 5 pairs of acr), acr in 2 widely separated rows, the distance between the two rows and between acr and dc almost equal. Wings brownish. S:f\ with a row of about 5 long fine bristly hairs pos- teriorly longer than femur is deep, bt x densely long pubescent dorsally, r 3 with dorsal and anteroventral bristles slightly longer than tibia is deep (temperate Europe, Southern France, Northern Italy) 70. H. lasiochira Strobl Antennal segment 3 smaller, style slightly longer; acr and dc bristle-like, but shorter than antennal style, acr in 2 closer rows, the distance bet- ween acr and dc is clearly larger. Wings more strongly brown infuscated. S'.fy posteriorly with hairs not longer than femur is deep, ty and bty mostly short pubescent, r 3 with small, thin dorsal and antero- ventral bristles shorter than tibia is deep (Spain, east to Switzerland) . . . H.H. strakai Chvâla 128(120) Larger species, body about 3 - 3.5 mm long. Scutum brownish dus- ted in anterior view, but prescutellar depression contrasting grey; when viewed from behind blackish-grey, with 3 broad indefinite black stripes along lines of bristles. Abdomen dull brownish dusted. Wings clear; palpi, legs and haltère quite black. All hairs and bristles fine, short, h and ih bristles fine; legs without distinct bristles, femora beneath with silvery pile. Abdominal pubescence and hairs on legs pale to rusty-brown. 6 : bty egg-shaped, twice as long as deep and much wider than tibia at tip, short pubescent. 9 : ? 3 slender at extreme base, otherwise very dilated, as deep as femur, and distinctly curved 89. H. borealis Oldenberg Smaller species, about 2-3 mm long, mostly shining black, or scutum only thinly pollinose when viewed from in front. Abdomen dull black also when viewed from behind. Small, short legged species (group 12, H. chorica- group) 129 129(128) o*: t] with a row of about 8 strong black dorsal bristles at least 3 times as long as tibia is deep. 9 : ? 3 spindle-shaped, strongly dilated about middle, tips narrower (also in H. pseudochorica) 130 o* : t\ with short hairs dorsally, at most with a few longer bristly hairs before tip, at most twice as long as tibia is deep at tip. 9 : f 3 slender or evenly compressed and curved, not spindle-shaped and broader than femur; if spindle-shaped and broader than femur (H. pseudochorica), then bt 3 with 2-4 distinct anterodorsal bristles 132 130(129) Scutum, when viewed from in front, uniformly finely brownish-grey dusted. S : only ty with long dorsal bristles, bty with 2 small, fine pre- apical bristly hairs 101. H. pectinipes Strobl Scutum shining black from all points of view, and without the combi- nation of other characters 131 131(130) Ocellar and frontal bristles equally long and strong. cT: bty and the following 3 tarsomeres long bristled dorsally; t 2 simple, not swollen, and covered with short hairs 102. H. barbipes Frey 542 M. CHVÀLA & B. MERZ Frontal bristles very small and fine, at most half as long as ocellar bristles, â : entire fore tarsus with short hairs, without distinct bristles; t 2 strongly dilated towards tip and dorsally densely covered with long bristles 103. H. hystrix Strobl 132(129) Scutum, when viewed from in front, finely brownish or brownish- grey dusted. S: bt\ rather longer ovate, about twice as long as deep, covered with short hairs 133 Scutum shining black from all points of view. 6: bt\ broader and shorter; if almost twice as long as deep, then with 2-5 long bristly hairs dorsally. Antennal style always long, about as long as 3rd segment, and wings more or less brownish 135 133(132) Antennal style very short, about 1/3 length of 3rd segment. Ocellar and frontal bristles equally long and fine. Scutum when viewed from in front uniformly finely dusted also anteriorly. 9 : r 3 simple and slender. Early spring species 94. H. brevistyla Collin Antennal style longer, at least 2/3 length of 3rd segment. 9 : f 3 more or less thickened, compressed and curved. Late spring and summer species . . 134 134(133) Scutum subshining black; when viewed from in front finely brownish Pollinose posteriorly, the brownish dusting clearly visible as two wide stripes also anteriorly between lines of bristles. Frontal bristles fine and very small, or completely absent. Palpi with a tendency to be light brownish at tip. 6 : t\ (except for preapical bristly hairs) and tarsus finely short pubescent, tarsomeres 2-4 simple, not produced dorsally; bt^ with short hairs; hypandrium small. 9 : / 3 slightly and evenly dilated, not deeper than femur, but distinctly curved 95. H. longivittata Zetterstedt Scutum in anterodorsal view with more or less darker, rather subshining black stripes between lines of bristles. Ocellar and frontal bristles almost equally long and strong. Palpi uniformly black. S: t\ with long bristly hairs dorsally, bt x rather densely long pubescent above and 2nd tarso- mere produced dorsally; bt^ with 4-5 long bristly hairs anteriorly; hy- pandrium very convex and produced distally. 9 : f 3 strongly dilated at middle, spindle-shaped, much wider than femur ... 96. H. pseudochorica Strobl 135(132) â : bt ± very stout and short, at most 1 .5 times as long as deep, dorsally with short hairs, at most with 1 or 2 fine bristly hairs dorsally before tip. 9 : tarsi shortened, tarsomeres 2-4 on fore leg as long as deep 136 â : bti longer, egg-shaped, about twice as long as deep, dorsally with 2 (occasionally up to 4) long black bristles; if two are present, then the upper one inserted at middle of basitarsus. 9 : tarsi longer, all tarsomeres on fore leg longer than deep; t 3 spindle-shaped, as wide as femur at middle, acr shorter than dc, and with a tendency to be 3- to 4-serial pos- teriorly; generally smaller species, body about 2 mm long . . 99. H. biseta Collin (The morphologically similar lowland species H. quadriseta Collin is larger-sized, up to 3 mm long, with acr regularly 2-serial, bristle-like, as long as dc; bt^ in both sexes with 2 small anterodorsal bristles before tip, bt j in S with 4-5 strong dorsal bristles, and ? 3 in 9 evenly dilated towards tip, but in apical third scarcely as wide as femur.) HILARA FROM SWITZERLAND 543 136(135) Legs quite black. Wings faintly brownish; acr in two uniform rows, o* : tarsomeres 2-4 on fore leg dorsally produced; hind trochanter simple. 9 : t\ dorsally at tip with a long bristle which is as long as bt^ is long 97. H. chorica (Fallen) Legs translucent yellowish at least on cx^ and base of/j (see couplet 43). Wings darker brown; acr strongly diverging on posterior half of scutum. S : only tarsomere 2 on fore leg produced dorsally; hind trochanter with a spur-like projection anteriorly, shorter 9 : dorsal bristle at tip of t\ much ] H. aartseni Chvâla LIST OF GROUPS AND SPECIES 1. Hilara flavipes group 1. H. flavipes Meigen. 1822 2. H.flavidipes Chvâla, 1998 3. H. morenae Strobl, 1899 3a H. sturmii Wiedemann in Meigen, 1822 4. H. albiventris von Roser, 1840 2. Hilara abdominalis group 5. H. abdominalis Zetterdtedt, 1838 6. H. allogastra Chvâla, 2001 7. H. alpicola Chvâla, 2001 8. H. gallica (Meigen, 1804) 9. H discolor Strobl, 1892 3. Hilara clavipes group 10. H. cilipes Meigen, 1822 11. H. clavipes (Harris, 1776) 12. H. curtisi Collin, 1927 4. Hilara canescens group 13. H. canescens Zetterstedt, 1849 14. H. nigritarsis Zetterstedt, 1838 15. H. cuneata Loew, 1873 16. H setosa Collin, 1927 17. H. lasiopa Strobl, 1892 18. H. eviana Straka, 1976 19. H. hyposeta Straka, 1976 20. H. manicata Meigen, 1 822 21. H ponti Chvâla, 1981 22. H. cinereomicans Strobl, 1892 23. H. merula Collin, 1927 24. H. thoracica Macquart, 1827 25. H. nigrita Chvâla, 2005 26. H. goetzei Chvâla, 2005 27. H. griseola Zetterstedt, 1838 28. H. pianti sp. n. 29. H. crossleyi sp. n. 30. Htiefii Strobl, 1892 31. H. perversa Oldenberg, 1916 5. Hilara litorea group 32. H. litorea (Fallen, 1816) 33. H. pseudosartrix Strobl, 1892 34. H. sartor Becker, 1888 35. H. galactoptera Strobl, 1910 36. H. morata Collin, 1927 37. H. splendida Straka, 1976 38. H. temila (Fallen, 1816) 39. H platyura Loew, 1873 40. H. leukensis sp. n. 41. H. nigrocincta de Meijere, 1935 42. H. ternovensis Strobl, 1898 (= H. griseifrons Collin, 1927) 43. H. albipennis von Roser, 1840 6. Hilara intermedia group 44. H. intermedia (Fallen, 1816) 45. H. tetragramma Loew, 1873 46. H. hirta Strobl, 1892 47. H. brevistriata sp. n. 48. H. beckeri Strobl, 1892 49. H.fuscipes (Fabricius, 1794) 50. H. quadrilla Chvâla, 2002 51. H. quadri/asciata Chvâla, 2002 52. H. pilipes Zetterstedt, 1838 53. H. tanychira Strobl, 1892 54. H. luteihalterata sp. n. 55. H. embartaki Chvâla, 2008 56. H. polleti sp. n. 57. H. hirtipes Collin, 1927 58. H. bistriata Zetterstedt, 1842 59. H. brevivittata Macquart, 1827 60. H. coracina Oldenberg, 1916 7. Hilara cornicula group 61. H. cornicula Loew, 1873 62. H.longifurca Strobl, 1892 (= H. monedula Collin, 1927) 63. H. campinosensis Niesiolowski, 1986 64. H.pygialis Chvâla, 2008 65. H. nigrohirta Collin, 1927 66. H. pseudocornicula Strobl, 1909 (= H. subpollinosa Collin, 1927) 67. H. quadrifaria Strobl, 1892 68. H lurida (Fallen, 1816) 544 M. CHVÂLA & B. MERZ 8. Hilara lasiochira group 69. H. pilosopectinata Strobl, 1892 (= H. longesetosa Strobl, 1910) 70. H. lasiochira Strobl, 1892 71. H strakai Chv âla, 1981 9. Hilara interstincta group 72. //. interstincta (Fallen. 1816) 73. //. lugubris (Zetterstedt, 1819) 74. H. dimidiata Strobl, 1892 75. H. scrobiculata Loew, 1873 76. //. sirbitzmatrona sp. n. 77. //. anglodanica Lundbeck. 1913 78. H. aeronetha Mik, 1892 79. H. angustifrons Strobl, 1892 80. H. caerulescens Oldenberg, 1916 10. Hilara andermattensis group 81. H. helvetica Chvâla, 1999 82. H maior Strobl, 1910 (= //. /arra Niesiolowski, 1991. syn. n.) 83. H. parvimaior sp. n. 84. H. snriaca Strobl, 1893 85. H. merzi Chvâla, 1999 86. H. zemiattensis Chvâla, 1999 87. H. andermattensis Strobl, 1892 88. H. simplicipes Strobl, 1892 1 1 . Hilara borealis group 89. H. borealis Oldenberg, 1916 90. H. medeterifonnis Collin, 1961 91. H. calinota Collin, 1969 92. H. lacteipennis Strobl, 1892 93. H. pruinosa Wiedemann in Meisen. 1822 12. Hilara chorica group 94. H. brevistyla Collin, 1927 95. H. longivittata Zetterstedt, 1842 96. H. pseudochorica Strobl, 1892 97. H. chorica (Fallen, 1816) 98. H aartseni Chvâla, 1997 99. H biseta Collin, 1927 100. H triseta Chvâla, 2005 101. H. pectinipes Strobl, 1 892 102. H. barbipes Frey, 1908 103. H hystrix Strobl, 1892 13. Hilara mauro, group 104. H. inaura (Fabricius, 1776) 105. H. diversipes Strobl, 1892 106. H. hybrida Collin, 1961 107. H. nitidula Zetterdtedt, 1838 108. H. nitidorella Chvâla, 1997 109. H. f emorella Zetterstedt, 1842 110. H. sulcitarsis Strobl, 1892 111. H. chpeata Meigen, 1 822 112. H. tyrolensis Strobl, 1892 (= H. miriptera, Straka, 1976, syn. n.) 113. H. discalis Chvâla, 1997 1 14. H. discoidalis Lundbeck, 1910 115. H. media Collin, 1927 14. Hilara albitarsis group 116. H. albitarsis von Roser, 1840 117. H. bohemica Straka, 1976 Additional species Hilara longicomis Strobl, 1894 Hilara flavocoxa Straka, 1976 Group 1 - H. flavipes-group For a characterization of this group, containing the original types of the genus, and for a revision of all known European species, see Chvâla (1998). 1. Hilara flavipes Meigen, 1822 References: Strobl (1892a: 160), Collin (1961: 679), Chvâla (2005a: 46) - description and illustration of male fore leg, hind femur, and postabdomen with genitalia, Chvâla (1998: 468) - synonymy. Distribution: A species of cold and temperate Europe, widely distributed from southern Scandinavia and the British Isles along the Atlantic coast south to France and Spain, in central parts of Europe south to Austria and Hungary only. In temperate and warmer parts of Europe preferring higher altitudes. It is a fairly common species for instance in the Czech Sumava Mts (Böhmerwald), found at many sites from 800 to 1 100m (see Chvâla, 1998), also in the Moravian Beskydy Mts, but there are only scarce records from the Alps. A late summer and autumn species, in mountain biotopes mainly from August to October. HÌLARA FROM SWITZERLAND 545 Material examined: Czech Republic: Sumava Mts: Antygl (1 100 m) 2.viii.l975 1 9; Prâsily, River Kremelna ( 800 m) 29. viii. 1994 1 9 ; Spâlenec (800 m) 15.viii.1994 16 29;Stary Brunst (960 m) 21.viii.1989 IS; Smauzy (960 m) 20.viii.1989 19 (all Bartâk); Jezernf slat' (1050 m) 4.-6.X.2005 16 (Spitzer). Beskydy Mts: Kosariska (550 m) 24.viii.1987 1(5; Homi Lomnâ (520 m) 10. viii. 1986 16 (both Bartâk). Austria: Carinthia: Villach 1 6 (Tief). 2. Hilara flavidipes Chvâla, 1998 References: Chvâla (1998: 466) - description and illustration of male post- abdomen. Distribution: Hilara flavidipes, a species closely related to the preceding H. flavipes, was described only recently from the Romanian Transylvanian Alps, and from western slopes of the Moravian Beskydy Mts (T inec). Its occurrence in other mountain ranges in central parts of Europe including the Alps is possible and, there- fore, it is included here as well. Material examined: Romania: Transylvanian Alps, Voineasa, 23. vii. 1987 1 6 (Bartâk). 3. Hilara morenae Strobl, 1899 References: Chvâla (1971: 334, 2008a: 32) - description, and illustration of male fore leg, postabdomen and genitalia. Distribution: This species, closely related to Hilara sturmii Wiedemann in Meigen, was described from the Spanish Cardenas, but now it is known also from Germany, Switzerland and Corsica. Recently, it was discovered among specimens collected by V. Straka in Sovakia on the River Danube at Prfbence near Samorin, 1 6 14.vii.1970 (UMO). Mainly a spring species, in May and June, in Switzerland at low altitudes at about 200 m only. Material examined: Switzerland: TI: Gordola, Bolle (220 m) 16.-19.vi.1995 9<5 8 9 (Merz & Bächli), Gordola, Verzasca (205 m) 15.-19.V.2006 26 6 9 , Gordola, Bolle di Magadino (200 m) 19.V.2006 26 29; Tenero, Lido 205 m, 15.V.2006 196 129 (all Merz), Bolle di Magadino, marais (196 m) 30.iv.1996 26 29 (Pollini). 3a. Hilara sturmii Wiedemann in Meigen, 1822 References: Chvâla (2005a: 50) - description and illustration of male post- abdomen. Distribution: Widely distributed in Europe from the British Isles and southern parts of Fennoscandia to the Mediterranean. Uncommon in central temperate Europe east to Romania; in southern Europe only in the western parts. A long flight period from May to September, but mainly in spring and early summer. Material examined: Switzerland. TI: Gordola, Bolle di Magadino (205 m) 6. vii. 2001 36 (Merz). 4. Hilara albiventris von Roser, 1 840 References: Collin (1961: 674), Chvâla (2005a: 51) - description and illustra- tion of male postabdomen and genitalia. Distribution: A species of temperate Europe, very rare in the south of Scandinavia (Norway), and from the British Isles (including Scotland) south to France 546 M - CHVÀLA & B. MERZ (Pyrenees), northern Italy and Romania. In central parts of Europe a typical summer species, occurring both in lowlands and at higher altitudes. Material examined: Czech Republic: Sumava Mts: Kasperské hory, Losenice (600 m) l.vii.1995 le? (Bartâk). Switzerland: GE: Chancy, La Laire (350 m) 25.vii.2004 19. ZH: Embrach, Haumiili (400 m) 10. vii. 1997 là (all Merz). France: Pyrenees, Ariège, Ax-les- Thermes (750 m) 8.VÜ.1990 1 S (Bartâk); Alpes-de-Haute-Provence, Montagne d. Lure (500 m) 1 S {Schacht). Romania: Transylvanian Alps, Voineasa 23. vii. 1987 AS (Bartâk). Group 2 - H. abdominalis-group For a characterization of this group, and a revision of all known Palaearctic species, see Chvâla (2001). 5. Hilara abdominalis Zetterstedt, 1838 References: Collin (1961: 618), Chvâla (2001: 205, 2005a: 55) - description, with illustration of male and female fore leg, and of male genitalia. Distribution: Throughout Fennoscandia, but much commoner in the north, especially in Lapland, also in Russian Karelia, and on the British Isles, especially in Scotland. Very probably a boreoalpine species, in temperate Europe known only at high altitudes of the Czech Krkonose Mts (Riesengebirge, or Giant Mts) and Sumava Mts (Böhmerwald). Unknown south of the Czech mountains, all other records from the Alps refer to other related species. The record from Switzerland of Schiegg et al. (1999) refers to Hilara alpicola (re-examination of A. Stark in 2009), and this species is therefore deleted from the list of Swiss Diptera. Material examined: Czech Republic: Krkonose Mts: Lyseciny (1000 m) vii. 1965 lo* (Macek). Sumava Mts: Trojmeznâ (1200 m) 12.-24.vi.2003 MT là; Boubin (1250-1350 m) vi.2004 PT 1 9 , MT 1 9 (all Farkac). 6. Hilara allogastra Chvâla, 2001 References: Strobl (1892a: 172, as H. heterogastra), Chvâla (2001: 208) - synonymy, homonymy, new name for Hilara heterogastra Nowicki, 1868, and description with illustration of male fore leg and genitalia. Distribution: A widely distributed species in mountains of central temperate Europe, south to Bulgaria (Pirin Mts), Romania (Transylvanian Alps) and east to the Caucasus (Georgia). Absent in northern Europe and the British Isles. The northern border of its distribution lies in the northern Czech mountains (Krkonose, Orlické hory, Jeseniky, Beskydy) and the Slovak West Carpathians (Velkâ Fatra, Vysoké Tatry, and the Bukovecké hory Mts on the Ukrainian border). For a long period from 19 May to 17 August, for detailed distributional data with the exception of the Alps, see Chvâla (2001). The species is often misidentified in the literature as H. abdominalis. Strobl (1893: 91; 1898: 206; 1910: 71) recorded it as H. heterogastra from many localities in the Styrian Alps, from the Ennstaler Alpen, and from the Rottenmanner and Schladminger Tauern. Material examined: Austria: Lower Austria (Schneeberg), Oberösterreich (Hinterstoder), Styria (Ennstaler Alpen, Gesäuse, Rottenmanner Tauern, Schladminger Tauern), Salzburgland (Sulzgau), Tyrol (St Ulrich, Grödnerthal, Bad Ratzes); common from 12 June to HILARA FROM SWITZERLAND 547 10 August, at altitudes from 700 to 1300 m. Switzerland: GR (St Moritz, Zernez, Valbella, Poschiavo), TI (Piora, Bedretto), VS (Längtal, Oberwald, Aletsch); common from 2 July to 17. August, at altitudes up to 1935 m. Italy: Landro, Trafoi, Monte Rosa, Macugnaga, S. Martino; common from 8 July to 15 August, at altitudes up to 1900 m. Slovenia: Julian Alps, Triglav. Aljazev dorn (1100 m) 3.VÜ.1973 4c? (Chvâla). 7. Hilara alpicola Chvâla, 2001 References: Chvâla (2001: 212) - description and illustration of male fore leg and genitalia. Distribution: Exclusively an Alpine species, known so far from the Austrian, Swiss and French Alps, mostly at high altitudes above 1000 m. A typical summer species, mainly in July and August, on dates ranging from 2 June to 14 August. Material examined: Austria: Styria: Ennstaler Alpen, Oberlaussa Polzanbach (850 m) 18.vi.2000 \S (Chvâla). Switzerland: VS: Visperterminen, Giw (1900 m) 16.vii.1997 lc? (holotype) (Merz); Riederalp 31.vii,-8.viii.l976 3 c? 69 (Bächli); Leuk, Pfynwald (600 m) 25.viii.2001 lc? (Merz & Landry). GR: Ftan 12.-14.viii.1978 2c? 29; Alp Flix 4.-8.viii.l975 2c?49 (all Bächli); Samedan Wald (1770 m) 16.vii.1985 le?, 18.vii.1989 4c?, (1840 m) 21.vii.1988 19 (ail Sauter); S. Vittore, Rebberg (290 m) 2.vi.l997 le? (Merz); Zernez, Crastatscha (1450 m) 16.vii.1980 19 (Haennï). France: Gard (Aigoual), Arphy, Cascade d'Orgon (1253 m) 17.viii.2002 le? (Haennï); Alpes Maritimes, Isola (2000 m) 30.vii.1993 le? (Stark). 8. Hilara gallica (Meigen, 1804) References: Collin (1961: 620), Chvâla (2005a: 53) - description, and illus- tration of male fore leg and genitalia. Distribution: Widely distributed in northern Europe as far north as the central parts of Fennoscandia, rare on the British Isles, where it occurs as in the Netherlands mostly on sandy heathlands; in temperate central parts of Europe south to France, Switzerland, northern Italy, Croatia and Hungary, preferring mainly dry steppe biotopes. A spring and early summer species. In Alpine regions already from the end of April to mid of July. Material examined: Switzerland: GR: S.Vittore, Rebberg (290 m) 2.vi.l997 lc?; Valbella, Casoja (1530 m) 14.vii.1998 19 (all Merz); Strada (1070 m) 13.vi.1986 le?; Ardez (1360-1380 m) 13.vi.1986 1 e? 1 9 ; Sent, Tuffarolas (1540-1580 m) 13.vi.1986 1 c? 1 9 (all Haennï). VS: Zermatt, Haveten (1760 m) 29.vi.1959 lc? (Reiser); Leuk, Platten (630 m) 22.iv.1998 lc? (Merz & Botta); Leuk, Brentjong (920 m) 15.V.2000 lc? (Merz & Ulrich). France: Hautes-Alpes, Montgenevre (1800 m) 12. vii. 1990 29 (Bartók). Italy: AO: Aosta, St Vincent Salirod (mosaique steppe) 14.V.2006 5 c? 2 9 (Haennï). 9. Hilara discolor Strobl, 1892 References: Chvâla (2001: 220) - description and illustration of male fore leg and genitalia. Distribution: A species of temperate continental Europe, in the north from Germany and Poland south to the Mediterranean (Spain, Corsica, Greece). In the Alps rather a submountain late summer species, mainly in August and September. Strobl (1892a: 165) described it as "Hilara discolor Kowarz i.litt." from specimens collected by Prof. Tief in the Jeseniky Mts (Czech Republic), and by T. Becker in German and 548 M. CHVÂLA & B. MERZ Polish Silesia; for details and lectotype designation, see Chvâla (2004: 131). In the south, in Spain and in the Mediterranean, high in mountains above 1000 m; not yet found in Austria. Material examined: Switzerland: SZ: Gersau-Oberholz 14.viii.1980 IS (Rezbanyai). TI: Piotta 19.-22.viii.1981 \S 19 (Badili). VS: Leuk, Rotafen (620 m) 10.viii.1997 IS 15 (Merz); Leuk, Finges (550 m) 9.viii.l997 IS (Haenni); Leuk (635 m) 10.viii.1997 1 5 (Haenni & Merz); Zermatt 5.viii. 1 9 (Becker, coll. Strobl); Sierre 9.x. 1890 1 ? (Huguenin). GR: Laax and Sasa vii.-viii.1892 IS 3 9 (Escher-Kündig); Ftan 12.-14.viii.1978 \S (Bächli). France: Vosges, Queleux 30.vii.1990 \S (Pont); Gard, Dourbies, 1.6 km SW Rouviere (840-880 m) ll.viii.1995 lo" 39; Gard (Aigoual), Arphy, Cascade d'Orgon (1253 m) 7.viii.2002 3 9 (all Haenni). Group 3 - H. clavipes-group For a characterization of this group, and a revision of the European species, see Chvâla (2005a: 57 and 2008a: 39). 10. Hilara cilipes Meigen, 1822 References: Strobl (1892a: 171), Chvâla (2005a: 60) - description and illus- tration of male fore leg, postabdomen and genitalia. Distribution: Not found in Fennoscandia and in the British Isles, in continental temperate Europe often misidentified as H. clavipes. From the Netherlands and central parts of Europe south to France, Switzerland, northern Italy and Romania. A spring and early summer species, everywhere rare. In the Alps also at altitudes about 1100 m. Material examined: Austria: Carinthia: Villach, Tschinowitsch, 23. v. IS (Tief). Styria: Gesäuse, Kaiserau (1100 m) 11. vi. 1997 IS (Chvâla). Switzerland: GE: Cartigny, Vers Vaux (335 m) l.vi.2002 le?; Bernex, Chante-Merle (415 m) 16.V.1999 29. SG: Wattwil (610 m) ll.vii.1997 1Ó\ZH: Zürich Katzensee (450 m) 23.v. 1995 1 S, (440 m) 25. v. 1996 le? (dWMerz). Slovenia: Spodnja Bilpa, River Kolpa (300 m) 15.vi.2006 IS 49 (Plant). 11. Hilara clavipes (Harris, 1776) References: Strobl (1892a: 169, as H. spinimana var. spinigera), Collin (1961: 655, as H. matrona), Chvâla (2005a: 58) - description and illustration of male fore leg, postabdomen and genitalia. Distribution: A widely distributed species in Europe, including northern parts of Fennoscandia, but especially common in temperate Europe, often in large swarms both in lowlands and in mountains. For a long period from June to September. A common species in mountains of central Europe, as for instance in the Krkonose Mts (Giant Mts) and Jeseniky Mts, in swarms above streams at altitudes above 1300 m. Strobl (1893: 91) recorded this species from the Austrian Alps (as a new variety spini- gera, his "//. matrona" was in fact Hilara lasiopa) from the Styrian Haller Mauern (Natterriegel), the Rottenmanner Tauern (Bösenstein), at about 1300-1700 m, also from Stelzing (1410 m) in Carinthia, and from Gastein in Salzburgland. In the French Alps also above 2000 m. Material examined: Austria: Styria: Haller Mauern, Natterriegel (about 1800 m) 22.VÌU. 19; Scheiplsee (= Scheibelsee) 30. vii. 1891 Id (all Strobl). Rottenmanner Tauern, HILARA FROM SWITZERLAND 549 Scheibelsee (1750 m) 6.vii.2002 1 o\ 15.ix.2005 39; Schladminger Tauern, Innere Grosssölkertal (1200 m) 24.viii.2001 swarms (all Chvâla). Salzburgland: Gastein 5. ix. 1$ {Becker). Switzerland: GR: Sur. Alp Flix (1965 m) 4.-8.viii. 1975 1 ö\ l.viii.2004 26 15 {Bächli); Ausserferrera (1320 m) 4.viii.l999 1(5; Valbella See (1500 m) 17.vii.1996 10c?, Valbella Casoja (1490-1500 m) 1 1 .-24. vii. 1 997- 1 998 11c? 8 9 (all Merz); Zernez, Gondas (1480 m) 4.viii.l996 1 , and a few slightly longer, fine anteroventral setae on/ 3 before tip; t\ (Fig. 17) covered with only short hairs, similarly 574 M. CHVÄLA & B. MERZ Figs 17-19 Hilara brevistriata sp. n. S (CH, TI: Magadino-Quartino, paratype). (17) Fore leg in posterior view. (18) Lateral genital lamella. (19) Hypandrium with postgonite. Scale: 0.2 mm. like the unusually long and slender bt\ and following tarsomeres on fore leg; posterior four tibiae slender and covered with only short indistinct hairs, but ? 3 with 7-8 fine dorsal setae at most as long as tibia is deep. Abdomen uniformly dull blackish-grey, rather densely short black pubescent, hind-marginal bristles long. Genitalia (Figs 18-19) fairly long, dull greyish-black like abdomen, but the long pointed apical half of hypandrium and apically very narrowed postgonites (often in a form of small spikes protruding up) distinctly shining black. The simple, and only short pubescent terminal process of lateral lamella clearly shows a close relationship of this species with the autumn species of H. hirta complex. Length: body 2.3 - 2.9 mm, wing 3.5 - 4.2 mm, holotype body 2.5 mm, wing 3.8 mm. Female. Very much like male, even the long bristling on head and thorax as in male, and wings practically clear as well. Legs long and slender, f^ posteriorly with only an indication of the dense posterior bristling; ? 3 long and only indistinctly thickened, much narrower than the corresponding femora, very slightly curved, rather undulating at middle, and the dorsal setae smaller than in male. Length: body 2.2 - 3.0 mm, wing 3.0 - 3.6 mm. HILARA FROM SWITZERLAND 575 Differential diagnosis: Superficially, this species resembles the early spring species H. brevivittata and H. bistriata, which both have a shorter and stouter antennal style, shorter setae on scutum, wing squama with pale fringes, different S genitalia with the circular and ventrally spinose terminal process on lateral lamella, and f 3 in 9 is thickened and curved, as wide as corresponding femur; H. brevivittata has the simi- larly shifting black pattern on scutum, but it has much shorter and long pubescent legs, whereas H. bistriata has similar long slender legs with long tarsi, but the 4 black stripes on scutum are well visible even in posterior view. The same black pattern on scutum with partly coalescing black stripes when viewed from behind is present also in a French coastal, still undescribed species (differing by the yellow haltère and long pubescent and bristled short stouter legs, Chvâla, in prep.), and in the Alpine H. lutei- halterata sp. n. (but legs and abdomen are whitish pubescent). It is evident, however, that although this early spring species shares many common characters with the two above discussed spring species of the H. brevivittata complex, based on the structure of male genitalia (the long simple terminal process of lateral lamella), H. brevistriata belongs to the H. intermedia complex of species sensu Chvâla (2005a). It is closely allied to the two late autumn species, H. hirta Strobl and H. hirtella Collin, 1927 (see Chvâla, 2005a: 1 14) having the same thoracic pattern with shifting black stripes, simple long slender legs devoid of distinct pubescence and bristles, and especially the structure of male genitalia is very similar. In addition to the very late summer occurrence, the larger size, dull grey occiput, yellow haltère, and the whitish pubescent abdomen may easily differentiate both these species from H. bre- vistriata. Holotype â : Switzerland: TI: Magadino-Quartino 205 m, 2.iv.2007 / 2, leg. B. Merz (MHNG). Paratypes: 13c?, 109 with the same data (MHNG, 40 and 2 9 UMO). Austria: Styria: Admont, Gesäuse Eingang, Enns River 620 m, 18.V.2003 19; Admont, Eichelau-Hall, Enns River 640 m, 22.V.2003 2 9 , all leg. M.Chvâla (UMO). Derivatio nominis: The species epithet is a combination of names of two closely similar spring Alpine species, Hilara brevi-vittata Macquart and H. bistriata Zetterstedt. Distribution: A species of lower parts of southern Switzerland and the Austrian Alps, at altitudes about 200 - 650 m, in April and May. 48. Hilara becked Strobl, 1892 References: Collin (1961: 592), Chvâla (2005a: 117) - description and illus- tration of male fore leg, female hind tibia, and male genitalia. Distribution: Practically throughout Europe, from southern Fennoscandia to the Mediterranean. In temperate Europe a common species in lowlands. It occurs also at higher altitudes, in the Alps up to above 1000 m, but not at least a "mountain species". A typical summer species, for a long period from May to the beginning of September, but common especially in July and August, though in lowlands of Switzerland (GE: Bernex) already at the end of April. 576 M. CHVÂLA & B. MERZ Material examined: Austria: Oberösterreich: St. Pankraz, River Steyer (490 m) 26.vi.2005 19. Styria: Admont (640 m) 8.vi.l996 29, 4.-7.VÜ.2006 59, ll.vii.2008 19 (all Chvâla); Gesäuse, an Ennsufer Fels (620 m) 14.viii. \S (lectotype), l.viii.1881 19, 28.v. 19; Kalbling 14.viii. 19 (all Strobl); Gesäuse, Kaiserau (1100 m) 7.viii.l998 19 {Chvâla). Switzerland: BE: Jura Schächental 2.vi.l963 IS {Reiser); Rubigen 26.V.1997 26 1 9 {Keller). ZH: Zürich, Hönggerberg (530 m) 17.vi.1997; 1 9 ; Zürich, Ahmend (430 m) 6.V.1998 IS (all Merz); Einbrach, Haumüli (400 m) 2.-19.vi.l998 2 9 {Wolf & Merz). TI: Biasca, Loderio (350 m) 17.vi.1997 1 S {Merz & Bächli); 2.vi.l997 IS, 17.vi.1995 1 S; Gordola, Verzasca-Ufer (205 m) 15.V.2006 1(5 (all Merz). GE: Chancy, Vers Vaux (335 m) l.vi.2002 le?; Bernex, Chante-Merle (415 m) 16.V.1999 1 â; Bernex, Signal (500 m) 30.iv.2007 1 9 ; Cartigny, Moulin de Vert (350 m) 5.vi.2006 1 9 (all Merz). Slovenia: Planinska Jama (800 m) 20.vi.2006 29; Gasparci, River Kolpa (310 m) 15.vi.2006 là 12; River Cabarca (280-330 m) 15.vi.2006 20 29 ; Spodnja Bilpa, River Kolpa (300 m) 15.vi.2006 7c? 49 (ail Plant). France: Gard, Dourbies, La Rouvière, Crouzoulous (970 m) 15. vii. 2005 2 9 (Haenni). 49. Hilara fuscipes (Fabricius, 1794) References: Strobl (1892a: 144 as H. carinthiaca Strobl), Collin (1961: 590), Chvâla (2005a: 1 15) - description and illustration of male fore leg, female hind tibia, male genitalia and postabdomen. Distribution: Widely distributed in Europe, from the British Isles and southern Scandinavia (Denmark and south Sweden) to the Mediterranean (France, Bosnia and Herzegovina). In central temperate Europe locally common. It occurs for a long period from May to August, both in lowlands and at lower altitudes in mountains, for instance in the Czech Sumava Mts up to 600 m. Strobl (1892a) described it as Hilara carin- thiaca, but later recorded it (Strobl 1893: 91; 1898: 206) under the correct name H. fuscipes from the Styrian Gesäuse and Admont. It is surprisingly rather a rare species in the Alpine region. Material examined: Austria: Lower Austria: Seitenstetten 1 9 {Strobl). Oberösterreich: Purgstall 14.vi.1971 19 (Ressi). Styria: Gesäuse viii. la 19 (Strobl). Switzerland: LU: Altbüron (530 m) 24.vi.1994 1 S (Merz). ZH: Embrach, Haumüli (400 m) 11. vi. 1997 IS 19 (Merz & Ziilliger), 19.vi.1998 1 9 (Merz). France: Pyrenees, Ariège, Ax-les-Thermes (750 m) 8.VÜ.1990 2 9 (Bartâk); Gard, Dourbies, 2 km SSE, rive du Crouzoulous (855 m) 21.vii.2009 IS (Haenni). Slovenia: Julian Alps, Planinska jama (800 m) 20. vi. 2006 19; Spodnja Bilpa, River Kolpa (300 m) 15.vi.2006 là; River Cabranca (330 m) 15.vi.2006 12c? 1 9 (ail Plant). 50. Hilara quadrula Chvâla, 2002 References: Chvâla (2005a: 119) - description and illustration of male mid tibia, female hind leg, and male genitalia. Distribution: A continental species known from Denmark and southern Sweden south to France; a lowland spring species, mainly in May and June. In warm temperate Europe absent in mountains, although in the south, for instance in southern France, preferring higher altitudes and where it may be found already in April. For a long time not distinguished from the following species, H. quadrif asciata, and gene- rally the two species were misquoted under the common name H. quadrivittata Meigen, a younger synonym of H. fuscipes (Fabricius). Common in lowlands of the Czech Republic, though less common towards the south, for instance not yet found in Austria, but not rare in Swiss and French lowlands. HILARA FROM SWITZERLAND 577 Material examined: Switzerland: VS: Sierre 14.V.1888 1 9 (Huguenin); Baltschieder, Rotten-Ufer (650 m) 17.V.1996 1 S {Merz). ZH: Zürich, Katzensee (440 m) 3.V.1997 16 1 9. TI: Biasca, Loderio (350 m) 2.vi. 16, 8.VÜ.1997 16 (all Merz). GR: S.Vittore, Rebberg (290 m) 2.-8.VÜ.1997 16 49 (Merz). France: Gard, Dourbies (1000-1040 m) 15.iv.1995 1 9 (Haenni). 51. Hilara quadri/asciata Chvâla, 2002 References: Collin (1961: 588 as H. quadrivittata Meigen, 1822), Chvâla (2005a: 120) - description and illustration of frons, male fore leg and mid tibia, female hind leg, and male genitalia. Distribution: A species of cold and temperate Europe, in comparison with the preceding H. quadrilla widely distributed throughout Fennoscandia and on the British Isles. In temperate Europe common also at higher altitudes, for instance in the Czech Sumava Mts frequently at about 800 m, but clearly of later occurrence, mainly in June and July. Towards the south only to the Alpine region, unknown in the Mediterranean. In the Austrian Alps fairly common as well above 1000 m, but not yet recorded from Switzerland and France. Material examined: Austria: Styria: Admont (640 m) ll.v.2007 16, 8. vi. 1996 swarms; Gesäuse Eingang, River Enns (620 m) 25.V.2009 16; Frauenberg, Ennstal (680 m) 14.vi.1997 19; Ennstaler Alpen, Unterlaussa (700 m) 14.vi.1997 19; Rottenmanner Tauern, Hohentauern, Weber Teich (1200 m) 5. vii. 2001 swarm (all Chvâla). Salzburgland: Schladminger Tauern, Obertauern. Tweng (1200 m) 17. vi. 2000 26 (Chvâla). 52. Hilara pilipes Zetterstedt, 1838 References: Chvâla (2005a: 128) - description and illustration of male fore and mid leg, and genitalia. Distribution: Northern species, common in the north of Fennoscandia in- cluding Lapland and Russian Kola Peninsula, south to the Baltic coast. In the continent only at scattered localities and very rare (Netherlands, Poland), but surprisingly not uncommon in the region of Berlin (as H. anomala Loew, 1840). Recently, it has been found in central Europe, in the Czech Krkonose Mts (Riesengebirge), and it may have a boreomontane distribution and will be found also at other sites, for instance the Alps. So far, all published records of H. pilipes from the Austrian Alps refer to other, morphologically similar species. A spring and early summer species, from the end of April to July. Material examined: Czech Republic: Krkonose Mts, Labskâ louka (1350 m) 23.- 26.V.2001 PT26 (Vanëk). 53. Hilara tanychira Strobl, 1892 References: Chvâla (2004: 1 17, 2008b: 242) - type specimens, synonymy. Diagnosis: A small species, body only about 2.5 mm long, with body and legs black, scutum with faint blackish stripes and 2-serial acr, haltère yellow. Legs long pubescent and bristled, also hind tibia in S with long bristly hairs dorsally (as in H. pilosopectinata). Antennae short, with very short style, and labrum long, as long as head is high. 578 m- chvâla & b. merz Redescription: Male. Frons and occiput dull black, face grey. Antennae with short 3rd segment, and also style very short and stout (!), half length of 3rd segment; labrum long, as long as head is high, all main characters very much like as in H. cantabrica Strobl (syn. H. lundbecki Frey; see Chvâla, 2005a: 169 and 2008a: 113). Thorax black, scutum blackish-grey in anterior view, dark grey in lateral view, with 4 indistinct, almost polished broad black stripes. The four stripes (the 2 indistinct blackish narrow stripes between 2-serial acr and 1 -serial dc, and the lateral 2 broader still less distinct stripes) very indistinct in anterior view, in posterior view scutum extensively blackish; in anterior view scutum usually almost unstriped. Mesonotal bristles long but rather hair-like. Wings clear, haltère clear yellowish. Legs quite black, fj only slightly shorter and deeper than posterior four tibiae, bti twice as stout as tibia, 2.5 times as long as deep, and shorter than t^, t\ and bt^ long pubescent, f j with 4 dorsal bristles twice as long as tibia is deep, also t 2 and f 3 densely pubescent; ? 3 dorsally with long hairs, very much as in H. pilosopectinata (syn. H. longesetosa Strobl). Genitalia large, not higher than abdomen, but produced below. Lateral lamellae long and slender, pointing up backwards; hypandrium very slender, long and strip-like, very much as in H. morata or H. cuneata. Length: body 2.3 - 2.8 mm. Female. Still smaller than the male, t\ and f 3 in addition to the pubescence with 5 short bristles. Differential diagnosis: The Scandinavian H. pilipes Zetterstedt, found also in cold temperate continental Europe and in mountains of central Europe (Krkonose Mts in the Czech Republic), is a larger species, body 3-4 mm long, with legs in $ covered with very long, fine bristly hairs, and with t 2 and bt 2 swollen and densely long pubescent (see Chvâla, 2005a: 128, Figs 199-203). The smaller-sized Alpine Hilara tiefii Strobl has the head including occiput dull grey, and blackish haltère. Distribution: The species was described by Strobl (1892a) from Melk in Lower Austria as a species "in Menge schwebend Ende April 1885 iiber feinem Ufersande einer Donauau bei Melk". Four syntypes, 3 â and 1 9 , collected on the banks of the River Danube in Austria were studied by Chvâla (2004: 1 17) in the Strobl Collection in Admont, one â was designated as lectotype. Further material is not avai- lable, and the 3$ collected on 12 May by Prof. Tief at Paternion in Carinthia, also mentioned by Strobl (1892a) in the original description, were not found. Material examined: Austria: Lower Austria: Melk, River Donau 13.iv.1885 3<5 1 9 (syntypes, Strobl). 54. Hilara luteihalterata sp. n. Figs 20-23 Diagnosis: A small, dull grey species of the H. intermedia-group (body about 2.5 - 3 mm long), with yellow haltère, black legs, whitish pubescent legs and abdo- men, and scutum dull grey, with distinct black altering stripes. Frons dull velvety black as in H. flavohalterata Strobl (known only from Croatia), and also occiput black when viewed from above, but extensively greyish when viewed from behind. HILARA FROM SWITZERLAND 579 Figs 20-23 Hilara luteihalterata sp. n. 6 (CH, GE: Chancy, Vers Vaux, paratype). (20) Fore leg in posterior view. (21) Postabdomen (macerated). (22) Lateral genital lamella. (23) Hypandrium with post- gonite. Scale: 0.2 mm. Description: Male. Frons and face very broad, face almost square-shaped and dull grey, frons velvety black, with a greyish triangular spot above antennae; vertex and occiput dull velvety black, although occiput, when viewed from behind, extensively greyish with only blackish shades, and on lower half below neck covered with whitish bristly hairs. Otherwise all setae on head black, vertical setae about as long as antennal segment 3, but the 2 pairs of ocellar and frontal setae very long and fine, much longer than upper setae on postocular margin. Antennae black, segment 3 short triangular, about twice as long as wide at base, style short and stout, only about half as long as segment 3. Palpi black, greyish pollinose, beneath with several long fine black setae. Proboscis short, labrum at most half as long as head is high. Thorax dull grey on pleura, more brownish-grey dusted on scutum; when vie- wed from anteriorly scutum almost unstriped, in anterodorsal view with 2 dull black stripes between rows of bristles, in dorsal view scutum with 4 distinct dull black stripes, the lateral stripes broader and longer, and prescutellar depression clearly 580 M. CHVÂLA & B. MERZ brownish; when viewed from behind, the 2 slender median stripes turn to a single broad median stripe on the acr setae. All setae on thorax black, only notopleural depression with several whitish hairs, similar to those on prothoracic epistema; acr narrowly 2-serial, fine and small, about 15 in a row and as long as the short antennal style; dc 1 -serial, clearly longer, but equally numerous; marginal setae in full number, 1 fine h seta, and last 2 pairs of prescutellar dc setae very long, as long as the inner pair of 4 sc setae. Wings clear, nearly whitish in some lights, with contrasting dark veins and a distinct blackish stigma, radial fork of usual hilarine-shape; anal vein practically absent, a costal bristle scarcely as long as the outer pair of sc setae. Squama dark grey to brownish-grey, fringes whitish; haltère yellow, with extreme base of stalk darkened. Legs uniformly black, almost subshining, finely greyish pollinose, and covered with mostly whitish hairs and also long setae are pale; only coxae dull grey like thoracic pleura, and covered with small whitish hairs; f x with a dense long whitish bristling posteriorly, the bristly hairs are much longer than femur is deep when viewed from above, also f 2 anteriorly with a row of long whitish setae (replacing the usual black anterior bristling), and/ 3 covered mostly with short pale hairs, longer setae only anteroventrally before tip; t] (Fig. 20) rather stout, posteroventrally with similar long pale bristling as on femur, dorsally with several long blackish setae; bt\ very stout but short, almost half as long a tibia, and covered with short hairs only; t 2 densely, rather brownish, long pubescent both anteriorly and posteriorly, the bristling is not much longer than tibia is deep (resembling in this way H. intermedia very much), but bt 2 slightly stouter than following tarsomeres, and densely, though shorter, bristled both antero- and posteroventrally (resembling in this way more H. pilipes); r 3 long and slender, only dorsally with about 7 to 8 fine blackish bristles about as long as tibia is deep; all tarsi rather short, but tarsomeres 2-4 on all pairs clearly longer than deep, although rather short on fore leg. Abdomen almost uniformly rather light grey dusted, covered with fairly long whitish hairs, hind-marginal bristles long, black. Genitalia (Figs 21-23) small, pointing backwards, lamellae covered with dark setae; lateral lamella with the characteristic circular, ventrally spinose terminal process of the H. brevivittata complex of species sensu Chvâla (2005a: 125). Length: body 2.3 - 3.0 mm, wing 3.3 - 3.6 mm. Female. Very much like male, head and thorax as in male, but wings with a ten- dency to be faintly clouded, not quite clear or whitish as in male. In contrast to the male the uniformly black legs are long and slender, covered (except for coxae) with only short dark hairs, long setae blackish and confined to the usual anterodorsal row onf 2 , and to about 4 long anteroventral preapical setae on/ 3 . Anterior four tibiae without dis- tinct setae, and/j posteroventrally (very densely long whitish bristled in S) with a row of only much shorter dark hairs, scarcely as long as femur is deep when viewed from above; f 3 distinctly thickened, rather laterally flattened, and as deep as femur, and distinctly curved when viewed from behind; dorsally with about 6-8 very fine dark setae much shorter than tibia is deep. All tarsi long and slender, even tarsomeres 2-4 on fore leg about twice as long as deep. Abdomen uniformly dull grey, more brownish- grey on dorsum, everywhere covered with short pale hairs, also the hind-marginal bristles fine and small, and clearly light brownish. HILARA FROM SWITZERLAND 581 Length: body 2.2 - 3.0 mm, wing 3.0 - 3.6 mm. Differential diagnosis: Resembling H. primula Collin, 1927 (see Chvâla, 2005a: 123) and H. flavohalterata Strobl, 1898 (see Chvâla, 2008a: 75). H. primula has less distinct blackish stripes on scutum, clearly grey dusted occiput, longer antennal style (although thickened), t 2 in S with short hairs, notopleura and humeri covered with black hairs, etc. The Mediterranean coastal H. flavohalterata seems to be more closely related, with similarly long pubescent S t 2 , but it is a larger species, body 3 - 4 mm long, with antennal style also short but thin, wings are faintly brownish infus- cated, and the pubescence on legs is blackish. Females of H. luteihalterata clearly differ from both above mentioned species by the f 3 which is thickened, about as wide as the corresponding femur, and distinctly curved. There is another similar species, H. embartaki Chvâla, which is easily separated by a long slender antennal style, scutum with only 2 blackish central stripes visible from all points of view, the wing squama is very pale, the legs in 6 are differently shaped, covered with black setae, and t-, in â bears short spine-like setae. It is not "woolly pubescent" as in H. luteihalterata, H. intermedia or H. brevivittata; also â genitalia are different (see Chvâla, 2008a: 74, Figs 67-70). Holotype S: Switzerland: GE: Chancy, Vers Vaux (335 m) 9.iv.2007 (10), leg. B. Merz (MHNG). Paratypes: 28 and 99 with the same data as holotype (MHNG, Id and 3 9 UMO). Derivatio nominis: lutei - halterata, indicates the yellow colour of haltère, but also its similarity with the Croatian H. flavohalterata Strobl. Distribution: Only known from the most southwestern locality in Switzerland, at the French border, on the border of the Rhône river. A very early spring lowland species, in the beginning of April. 55. Hilara embartaki Chvâla, 2008 References. Chvâla (2008a: 73) - description and illustration of male fore leg, postabdomen and genitalia. Distribution: Morphologically similar with H. luteihalterata, but also with the next new species of the Bulgarian Rhodope Mts, H. polleti. It has been described only very recently from the Gulf of Lions south of the French Massif Central, and now it is recorded also as a real mountain species of the French Massif Central, at an altitude above 1500 m. Its occurrence in western Alps is very probable. Material examined: France: Hérault, Golfe du Lion, Col du Cabaretous (940 m), NW of Beziers, 22.V.2006 3c? 19 (Bartàk); Lozère, Sommet de Finiels, 1 km S (Valat de la Mouline) (1570m) ll.vii.1998 1 S (Haenni). 56. Hilara polleti sp. n. Figs 24-26 Diagnosis: Medium-sized dark dull grey species (body slightly more then 3 mm long), with occiput and frons dull black, scutum dull brownish-grey, with 2-serial acr, and 4 blackish longitudinal stripes; legs densely, mostly black pubescent, wings faintly brownish and haltère yellow, abdomen whitish pubescent on basal segments. 582 m. chväla & b. merz Description: Male. The square-shaped face light grey, frons equally wide but dull black like vertex and occiput, only above antennae with a slight greyish shadow; lower part of occiput below neck greyish dusted and covered with whitish hairs. Other hairs and bristles on head black, ocellar and frontal bristles equally very long, longer than antennal segment 3 without style, postvertical bristles smaller. Antennae black, segment 3 long, style rather slender and about two-thirds length of segment 3. Palpi greyish, short black pubescent ventrally, and with a very long preapical bristle before tip. Labrum short, much shorter than head is high. Thorax rather light grey dusted on pleura, scutum including scutellum uni- formly contrasting dull brownish-grey, when viewed from in front scutum quite un- striped, in all other views with 4 dull black stripes, the 2 narrower blackish stripes between acr and dc bristles visible in anterior half of scutum only, absent in prescu- tellar region, laterally of dc 2 broader and longer similarly black stripes; when viewed from behind, the lateral stripes less distinct; compared with all related species of the H. brevivittata complex, the scutum is almost uniformly dull brownish-grey, not velvety black as usually. All thoracic setae black, acr narrowly 2-serial, dc 1 -serial, all shorter than antennal style; acr less numerous, only about 10 setae in a row, dc more numerous; large marginal setae in full number, h and ih setae rather small and fine, other setae strong, the inner pair of 4 sc the longest. A small, but stout bristle on each side of pronotum, otherwise prothoracic epistema and prosternum with minute whitish hairs; prothoracic spiracle contrasting black. Wings faintly brownish clouded, not quite clear, with contrasting dark veins and a distinct dark brown stigma; a long radial fork, anal vein indistinct, and costal seta about as long aspa seta. Squama light greyish with a very dark margin, fringes whitish. Haltère clear yellow on knob, stalk blackish-brown. Legs uniformly black, including "knees", finely greyish pollinose, and covered with dense black hairs and setae, only cjcj with white hairs; /j posteriorly with long dense bristling about as long as femur is deep, f~> with usual long anterior setae, but otherwise with short fine pubescence, as in/ 3 , with the exception of 3 to 4 long antero- ventral setae before tip of the latter; t\ (Fig. 24) stout, dorsally with 5 pairs of very long black bristles directed dorsally and posterodorsally, and posteriorly with dense bristling at least as long as tibia is deep; t 2 (Fig. 25) with similar long dense bristling as in H. intermedia, anterior bristles longer than tibia is deep, posterior bristling denser, but shorter; r 3 ventrally with about 7 to 8 setae not longer than tibia is deep, dorsally with the same number of longer setae; bty (Fig. 24) very thickened, clearly shorter than corresponding tibia, but as long as rest of tarsus, and even on dorsum covered with only short hairs; tarsi short, tarsomeres on fore leg rather stout, only slightly longer than deep, those on posterior four tarsi longer, and all covered with short hairs only. Abdomen densely dull grey dusted, more brownish-grey on dorsum, when viewed from behind with distinctly paler, almost silvery grey, narrow hind margins; pubescence whitish on basal two segments, especially long on the narrow basal segment, following segments covered with black hairs, the long hind-marginal bristles on all segments also black. Genitalia (Fig. 26) small and rather narrow, pointing back- wards, very much like as in the more northern H. brevivittata Macquart; hypandrium HILARA FROM SWITZERLAND 583 Figs 24-26 Hilara polleti sp. n. S (BG, Rhodope Mts, paratype). (24) Fore leg in posterior view. (25) Mid leg with basitarsus in dorsal view. (26) Postabdomen and genitalia. Scale: 0.2 mm. only slightly produced posteriorly, very narrowly pointed apically; lateral lamella covered with long blackish setae, and apically with the usual circular ventrally spinose apical process of the H. brevivittata complex of species. Length: body 3.2 - 3.3 mm, wing 4.2 - 4.5 mm. Female unknown. Differential diagnosis: This species of the Bulgarian Rhodope Mts is most closely related to H. embartaki, known so far only from mountains of the French Massif Central. H. embartaki is in general a smaller-sized species (body 2.8 - 3.2 mm, wing 2.9 - 3.3 mm), especially wings are shorter, there are only 2 dark stripes on scutum between lines of bristles (lateral stripes absent), the median region on acr setae is clearly greyish, and the 2 blackish median stripes are much more distinct in posterior view; acr and dc similar but more numerous, with about 16 acr setae in one row (only 10 setae in H. polleti), wings are almost clear in H. embartaki, iridescent, and haltère entirely very pale yellow including the stalk. Legs are in H. embartaki similarly uni- formly black, although only finely pollinose, much more subshining; bristling on femora is similar, but t\ bears only a few, usually 4 dorsal bristles in one row, and t 2 bears only 3 short spine-like anteroventral bristles about as long as tibia is deep. The long dense bristling of H. polleti on tibiae is absent. Genitalia of H. embartaki differ from those of H. polleti by only a sparse long bristling of lateral lamella, and the apical circular process is spinose along margins only, ventrally it is bare. 584 M. CHVÄLA & B. MERZ Holotype â: Bulgaria (Rhodope Mts): Blagoevgrad, 15.iv. 1973, sipei. kol., leg. V. Beschovsky (UMO). Paratypes: 1 6 with the same data as holotype, pinned with the holotype on the same pin (UMO). Derivatio nominis: The species is named after the Belgian dolichopodid spe- cialist Dr Marc Pollet. Distribution: Bulgaria; in the western part of the Rhodope Mts close to Macedonia, its distribution along the Balkan mountain ranges west to the Alps may be anticipated. 57. Hilara hirtipes Collin, 1 927 References: Collin (1961: 599), Chvâla (2005a: 132) - description and illus- tration of male fore and mid leg, and genitalia. Distribution: A species with a still problematic and unclear distribution in Europe. Described from Great Britain, and with certainty known so far along the Baltic Coast, from the Netherlands, Belgium and Germany. Further, it was recorded from Central Europe from Switzerland by Schiegg et al. (1999). Other records from moun- tains of central temperate Europe and from the Alps need revision because of their pos- sible confusion with morphologically similar species. The only other verified identifi- cation from Central Europe is a male in the Admont Strobl Collection from Vorarlberg, identified by Strobl as H. bisiriata Zetterstedt. Mainly a spring species, from April to the beginning of July. Material examined: Austria: Vorarlberg 13. v. \6 (coll. Strobl). Switzerland: ZH: Sihlwald 24.v.-19.vi.l996 3ô (Schiegg). 58. Hilara bistriata Zetterstedt, 1 842 References: Collin (1961: 603), Chvâla (2005a: 135) - description and illus- tration of male fore leg, postabdomen and genitalia. Distribution: A boreomontane species, common throughout Fennoscandia in- cluding Lapland and Kola Peninsula, on the British Isles mainly in Scotland, and then in mountains of central Europe including the Alps. For full data on its distribution in Europe, see Chvâla (2002b: 238). A species of peat-bogs in the Czech Sumava Mts (Böhmerwald) at altitudes between 700 and 800 m; at similar altitudes as well in the Styrian Alps, mainly along streams on flowering willows. An early spring species, in the Alps mainly in April and May. Strobl (1893: 90, and 1898: 206) recorded it also as a common species in May and June in Carinthia, and later from Admont and Gesäuse. Material examined: Austria: Styria: Admont 2.vi. 1 6 , Krumau 2.v. 1 3 ; Gstatterboden 28.v. 1 S (all Strobl); Ennstal near Admont (640 m) 28.iv. 1