Tijdschrift voor Entomologie 156 (2013) 103–112 brill.com/tve

Lydella slavonica, a new species from the western Palaearctic with notes on the subgenus Lydelloxenis (Diptera, Tachinidae) Theo Zeegers

Lydella (Lydelloxenis) slavonica sp. n. is described from Montenegro and Poland. The definition of the subgenus Lydelloxenis Mesnil, 1956 is discussed. A key to western Palaearctic subgenera of Lydella Robineau-Desvoidy, 1830 and species of Lydelloxenis is provided. A phylogenetic analysis for the genus Lydella is provided. The species from the tropics and subtropics are found to form a monophyletic group, called the Metoposisyrops-clade. The species of this clade combined with those of Lydelloxenis are found to form a monophyletic group as well. The results are weakly supported. For practical reasons and nomenclatural stability, Lydelloxenis and Metoposisyrops Townsend, 1916 are proposed as subgenera. Keywords: parasitoids, stem boring caterpillars, phylogeny, Exoristinae, Eryciini. Theo Zeegers, Eikenlaan 24, 3768 EV Soest, the Netherlands. [email protected]

Introduction Within the western Palaearctic, the genus Lydella The genus Lydella Robineau-Desvoidy, 1830 is consists of grey tachinids of medium size. The genus known from all major biogeographic regions (O’Ha- is characterized by the bare eye, the erect apical ra 2010). As currently understood, it includes the scutellar setae, presence of one very strong setula subjective synonyms Lydelloxenis Mesnil, 1956 from at base of vein R4+5 and the presence of median the Palaearctic region (Herting 1959) and Dia- discal setae on tergites 3 and 4 (Tschorsnig & Richter traeophaga Towsend, 1916, Metagonistylum Town- 1998). Six species are known (Herting 1984), falling send, 1927 and Metoposisyrops Townsend, 1916 from into two distinct groups. the subtropics and tropics of the old and new world The nomino-typical subgenus consists of four (Woodley 1994). The genus belongs to the tribe species characterized by the presence of fields of spe- Eryciini, of the subfamily Exoristinae (Herting 1960, cialized hairs on the ventral surface of the male fourth Wood 1987, Tachi & Shima 2010). An important tergite (= ‘Sturmia-spots’) (Tschorsnig & Herting feature of this genus is the presence of one very strong 1994), i.e. the type-species L. grisescens Robineau- setula at the base of vein R4+5.AllspeciesofLydella, Desvoidy, 1830 and L. ripae (Brischke, 1885), L. as far as known, are parasitoids of stem boring cater- stabulans (Meigen, 1824) and L. thompsoni Hert- pillars (Woodley 1994) belonging to several families ing, 1959. The subgenus Lydelloxenis,inwhichthese of Lepidoptera, especially Noctuidae and Pyralidae specialized hairs are lacking, consists in the western (Herting 1960), several of them of economic im- Palaearctic of two members: its type-species L. brevi- portance (Cleare 1939, Holloway & Mathes 1940, seria (Pandellé, 1896) and L. lacustris Herting, 1959. Herting 1959). The host choice is a synapomorphy This paper adds a third species: L. slavonica sp. n. for the genus (Woodley 1994). from Montenegro and Poland. Members of the sub- genus Lydelloxenis are very rarely collected.

Tijdschrift voor Entomologie 156: 103–112, Tables 1–3. Figs 1–22. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 15 July 2013. DOI 10.1163/22119434-00002023

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Methods Note: L. matutina Richter, 2003 from Sakhalin, Morphological terminology follows Oosterbroek et Russian Far East, belongs to Lydella (s.s.) but has only al. (2005) and Tschorsnig & Herting (1994). Mea- 2 anterodorsal setae on mid tibia. sures were taken using an Olympus SZ-61 stereo- microscope with an ocular with scale. Photographs Keytothespeciesofthesubgenus were produced with an Olympus SP-500UZ camera Lydelloxenis in the western Palaearctic mounted on a separate phototube on the stereomi- croscope. Composite multi-focal images were pro- 1. Hind tibia with 3 distinct preapical setae, duced by stacking several images using the software the dorsal equal to posterodorsal and an- program CombineZM (Hadley 2007). Phylogenetic terodorsal one. Mid tibia with 2 strong an- relationships have been studied by parsimony ana- terodorsal setae. Facial ridge with setae in lysis, using the computer program TNT 1.1, as de- lowerhalf...... L. breviseria scribed by Goloboff et al. (2008), running on com- – Hind tibia with 2 distinct preapical setae, puter under Windows XP. dorsal one lacking or very small. Mid tibia The acronyms for collections follow Evenhuis with 1–2 anterodorsal setae, the second seta (2012). To these, I add the following: CCBW – per- less than half as long as the first. Facial ridge sonal collection of C. Bystrowski, Warsaw. with setae restricted to lower third or less . . . . 2 2. Katepisternum with 4 setae (Fig. 3). Api- cal scutellar seta as long as scutellum, longer Results than discal scutellar seta (Fig. 7). Third an- × Key to subgenera of Lydella in the western tennal segment slender, more than 4 as Palaearctic long as broad. Basicosta and tegula black. Male: proclinate orbital seta absent (Figs 1, 1. Genal dilation narrow, occupying at most 4). Wing membrane distinctly darkened half of gena, with only 2 rows of hairs along veins (Fig. 11). Claws and pulvilli (Figs 1, 14, 16). Fore and mid tarsus elon- of fore leg longer than fifth tarsal segment × gated, about 1.2 as long as tibia (Fig. 5). (Fig.6)...... L. slavonica sp. n. First posterior intra-alar seta small, dis- – Katepisternum with 3 setae. Apical scutel- tinctly smaller than other intra-alar setae. lar seta shorter than length of scutellum, Notopleuron with 2 setae, otherwise bare or only as long as discal scutellar seta (Fig. 18). at most with 3–4 hairs. Mid tibia with 1–2 Third antennal segment not slender, at anterodorsal setae. Male: Tergite 4 ventrally most 3× as long as broad. Basicosta brown, without fields of specialized hairs (Fig. 10). tegula black. Male: one pair of proclinate Postocular setulae short, nearly straight. orbital setae (Fig. 14). Wing membrane Fronto-orbital plate between frontal setae slightly yellowish, more so at base (Fig. 19). and eye margin with very sparse and short Claws and pulvilli of fore leg distinctly × hairs. Third antennal segment at most 3 shorter than fifth tarsal segment (Fig. 17) aslongassecond...... subgenusLydelloxenis ...... L. lacustris – Genal dilation broad, occupying more than half of gena, with 4 or more rows of hairs (Fig. 20). Fore and mid tarsus of normal Description and redescription of species length, not longer than tibia. First poste- of the subgenus Lydelloxenis rior intra-alar seta strong, as strong as other I have added a note on Lydella (Lydelloxenis) intra-alar setae. Notopleuron with 2 setae columbina from the eastern Palaearctic, so that all and at least 5 hairs but usually many more. known species of Lydelloxenis are dealt with. Mid tibia with 3 or more anterodorsal se- tae. Male: Tergite 4 ventrally with a pair of fields of specialized hairs (‘Sturmia-spots’) Lydella (Lydelloxenis) slavonica sp. n. (Fig. 21). Postocular setulae long, distinctly Figs 1–11 curved forward. Fronto-orbital plate be- Lydella (Lydelloxenis) spec. nov. Zeegers 2010: 6. tween frontal setae and eye margin with Lydella (Lydelloxenis) lacustris: Bystrowski 2005: 3 dense, long hairs, increasing in length to- [misidentification]. wards the frontal setae. Third antennal seg-  ment at least 3× aslongassecond...... Type material. Holotype: , Montenegro, Skadar- sko Jezero [= Skadar Lake] WSW Podgorica, Vran- ...... subgenusLydella jina, 42°17 N; 19°08 E, 13.vi.2009, leg. G. Pennards

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Figs 1–11. Lydella slavonica sp. n., male, holotype. – 1, head, lateral view; 2, thorax, dorsal view; 3, ventral half of thorax, lateral view; 4, head, dorsal view; 5, right fore leg: tibia and tarsus, posterior view; 6, right fore tarsus with claws and pulvilli, posterior view; 7, scutellum, lateral and slightly dorsal view; 8, abdomen, dorsal view; 9, abdomen, view obliquely from behind; 10, abdomen, lateroventral view; 11, right wing.

& Th. Zeegers. The holotype will be deposited in (CCBW);  Poland,Biebrzanski´ NP, Dolny Basen RMNH. Biebrzy, Bagno Ławki, Ambona NFOS,´ 2.vii.2003, Paratypes:  Poland,Biebrzanski´ NP, Dolny Tur zycowisko [= sedge marsh] z Comarum palustre i Basen Biebrzy, Gr. Honczarowska, 25.vi.2003, col- Equisetum palustre,leg.C.Bystrowski(SMNS). lected on Equisetum palustre,leg.C.Bystrowski

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Description 5× as long as broad and 2.5× as long as second, Male. Body length 9.5 mm. apical end obliquely truncated, so that the dorsal Colouration (Figs 2, 9). Generally black with apex becomes somewhat angular. Arista thickened on some silvery-white pruinescence. Parafacial, gena and basal two-fifths only, second segment nearly twice occiput behind eye silvery white, fronto-orbital plate as long as broad. Vibrissa at oral margin, setulae slightly yellow, frontal vitta black. Antenna black, above vibrissa small, ascending on lower third of fa- palpus black with yellow tip. Two to three irregular cial ridge. Prementum short (difficult to see), labella rows of black setulae on occiput behind postocular large. Postocular seta short and straight. row, otherwise occiput covered with white setulae. Thorax (Figs 2–3, 5–7, 11). Thoracic dorsum Thoracic dorsum in ground colour black, with light with 3 + 3 acrostichal setae, 3 + 4dorsocentral greyish pruinescence, leaving four distinct dark vit- setae, 1 + 3 intra-alar setae, the first posterior one tae before the suture, the inner pair narrow and with smaller, 1 + 3 supra-alar setae, third posterior one sharp borders, the outer pair broad and more dif- nearly as strong as the first posterior one. Noto- fuse. Behind the suture, there is an additional central pleuron with 2 strong setae, otherwise bare (with- black vitta present, merging with the inner pair into out hairs). Postpronotum with 4 setae, the basal three a diffuse broad central vitta. Scutellum black, with on a straight line, the anterior and inner basal much some grey pruinescence at the tip. Legs black, pul- smaller than the other two. Katepisternum with four villi yellowish white. Wing distinctly tinted brown setae, the third one the smallest but still well de- along main veins and cross-vein DM-Cu, veins dark veloped; one strong pteropleural seta. Scutellum co- brown, basicosta and tegula black, calyptra white, vered with erect hairs, with a pair of discal setae halter brown. Syntergite 1 + 2 black, tergites 3 and and four pairs of marginal setae: the basal and sub- 4 black with transverse bands of silvery pruinescence apical ones very large, the apical pair large, about anteriorly, broadly interrupted in the middle. Bands as long as scutellum, erect and crossed, the lateral of tergite 3 and 4 covering about one-third of ter- one small, half as long (measured: 0.47×)andmuch gite, the one on tergite 3 distinctly broadening to- thinner than subapical. Posterior thoracic spiracle wards lateral margin. Tergite 5 black with transverse closed by large circular posterior lappet (as usual in band of silvery pruinescence covering anterior fifth Tachinidae). of tergite. Silvery bands of pruinescence continuing Tarsi of all legs elongated, length of fore and mid on ventral side of tergites 4 and 5, but lacking on ter- tarsus 1.2× as long as corresponding tibia, tarsus of gite 3; ventral side of tergites 2–4 completely turning hind leg slightly longer than hind tibia. Claws and silvery in some angles of view. pulvilli elongated, longer than the fifth tarsal seg- Head (Figs 1, 4). Vertex in dorsal view slightly ment (measured: 1.21×). Fore tibia with 2 small narrower than width of one eye (measured 0.93×), posterior setae, mid tibia with 1 strong and com- fronto-orbital plate just before ocelli slightly nar- pletely isolated anterodorsal seta, 1 strong ventral rower than frontal vitta (Fig. 4). Face about as long seta and 2 smaller posterodorsal setae, hind tibia with as frons, in lateral view the frons strongly protruding, 1 large and 7 smaller anterodorsal setae, 1 large and 1 parafacial more than twice as broad at level of anten- smaller posterodorsal and 3–4 smaller anteroventral nal base than as level of lower eye margin (Fig. 1). setae, strong anterodorsal and posterodorsal preapi- Gena narrow, slightly narrower than width of third cal setae and 1 small anteroventral preapical. Hind antennal segment and much narrower than width coxa bare on posterior surface. of parafacial at level of antennal base. Genal dila- Wing with cell r4+5 open, sixth section of costa tion narrow, less than half of height of gena, covered slightly shorter than fourth. Section of vein M be- with only 2 rows of hairs and one row of setae at tween cross-vein DM-Cu and bend half as long lower margin. Fronto-orbital plate with 2 reclinate as section between crossveins R-M and DM-Cu, orbital setae, proclinate orbital setae lacking. Inner slightly shorter than crossvein DM-Cu; apical sec- vertical seta strong, about half as long as greater eye tion of vein M much longer, distinctly concave. Bend diameter, length of outer vertical seta two-fifth of of vein M angular, without appendix. Base of vein inner one. Ocellar seta latero-proclinate, strong, as R4+5 with one very strong, curved setula, as typi- long as anterior reclinate orbital seta, situated just cal for the genus. Costa with setulae up to halfway behind the anterior ocellus. Frontal setae descend- fourth section, costal spine differentiated but small, ing to base of arista, the lower ones curved upwards. about twice as long as other setulae. First section of Fronto-orbital plate with 3 smaller setulae near an- costa with setulae on ventral surface, second section tennal base, otherwise fronto-orbital plate between with a few setulae. frontal setae and eye margin very sparsely covered Abdomen (Figs 8–10). Tergites covered with ad- with minute hairs. Parafacial bare. Antenna predom- pressed hairs. Syntergite 1 + 2 with a pair of median inantly hidden in face, third antennal segment about marginal setae and a pair of lateromarginals, tergite

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3 with a pair of median discals, a pair of median Ecology marginals and a pair of lateromarginals, fourth ter- The type locality is situated at the banks of Skadarsko gite with a pair of median discal setae, laterodiscal Jezero (Figs 12–13). The holotype was collected present but not conspicuous, and a row of marginal around noon at 32°C on the inflorescence of Matri- setae, fifth tergite with a row of discal setae and a row caria along a newly built road entering the marsh. of marginal setae. Ventral surface of sternites uni- Records from the hottest part of the day are formly covered with hairs, some larger setae near in- rather atypical in this group of Tachinidae (Cerretti ner margin, no trace of specialized hairs or setulae. 2005). Genitalia not examined. The female paratypes were collected at the south- Female. The female differs from the male as fol- ern part of the marshes accompanying the Biebrza lows: river in north-east Poland. These marshes are Wings clear. Fronto-orbital plate with 2 pairs amongst the best preserved in central Europe. Both of proclinate orbital setae. Third antennal segment specimens were collected in a stand of Equisetum twice as long as second. Notopleuron with 0–5 hairs palustre. additional to the two notopleural setae. Claws and pulvilli short. Mid tibia with a smaller additional anterodorsal seta above the large one. Lydella breviseria (Pandellé) Roeselia (Frontina) breviseria Pandellé, 1896: 46. Syntypes female, France (MNHN) (teste Herting 1978); (CNC) Diagnosis (O’Hara in litt.) [not seen]. Lydella slavonica is readily distinguished from other Peteina rectangula Pandellé, 1896: 106. Holotype male, France species in the subgenus Lydelloxenis by the presence (MNHN) (teste Herting 1978) [not seen]. of four katepisternal setae. It is most similar to lacustris, in which the third antennal segment is less Description slender and the apical scutellar setae are distinctly Described in detail and illustrated by Mesnil (1956) shorter. The wing is distinctly darkened along the and reviewed by Woodley (1994). O’Hara kindly re- major veins in the male sex. examined on my request the female specimen in the Mesnil collection (currently in CNC) and confirmed Etymology (in litt.) the presence of the following features: The species has so far been found only in the eastern, • notopleuron with 2 large setae and 2–3 addi- i.e. Slavonian, part of Europe. The specific name tional hairs; should be treated as an adjective. • fore tarsus 1.2× as long as fore tibia (mea- sured); Distribution • lateral scutellar seta 0.63× as long as subapical Only known from the types from eastern Europe: scutellar seta (measured). Montenegro and Poland.

Figs 12, 13. Type locality of Lydella slavonica at Skadarsko Jezero, Vranjina, Montenegro. 12, general impression of the habitat of the bank of the lake; 13, type locality at road side.

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Lydella lacustris Herting Figs 14–19 Notopleuron with 2 setae, otherwise bare, in fe- male paratype with 2 minute hairs. Apical scutellar Lydella lacustris Herting, 1959: 424. Holotype male: setae erect, crossed, shorter than the length of the Austria: Neusiedler See (NMW) [examined]. Paratype fe- scutellum and therefore shorter than in any Lydella male, Austria: Neusiedler See (NMW) [examined]. (s.s.) (Fig. 18). Fore tarsus elongated, 1.2× as long as fore tibia in both sexes (measured: male: 1.21×; Description female 1.17×) (Fig. 17). To the original description by Herting (1959), I add Section of vein M between crossvein DM-Cu and the following: 3–4 irregular rows of black setulae on bend long, clearly longer than crossvein DM-Cu occiput behind postocular row. Postocular setulae in and about as long as the apical section of vein M male short and virtually straight. Scutellum slightly (Fig. 19). Costal spine present, though weak and reddish brown at its apex. Wing membrane faintly easily overlooked. tinted yellow, veins in basal half of wing yellow. Tegula black, basicosta dark brown. The bands of Diagnosis grey pruinescence on tergites are narrower in the × The presence of proclinate orbital setae (one pair) in female, especially on tergite 4, occupying 0.25 the male sex is characteristic for this species. Woodley tergal length. (1994), who could not study this species, overlooked Fronto-orbital plate in male with 1 proclinate seta the presence of the proclinate orbital seta in the male (Figs 14), in female with 2 (Fig. 16), otherwise with in the original description. very sparse and short hairs. Ocellar setae situated at level of anterior ocellus.

Figs 14–19. Lydella lacustris, 14–15, 17–19, male, holotype; 16, female, syntype. 14, head, lateral view; 15, head, dorsal view; 16, head, lateral view; 17, tibia and tarsus right fore leg, anterior view; 18, thorax, slightly obliquely lateral view; 19, right wing.

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Lydella columbina Richter taxon, a member of the genus Drino Robineau- Lydella (Lydelloxenis) columbina Richter, 1976: 542. Desvoidy, 1863 was chosen, as suggested by Woodley Holotype and paratypes female, Mongolia:Uvsprovince, (1994): Drino inconspicua (Meigen, 1830). In total, = Uvs Nuur [ UvsLake],50kmfromUlaangom(ZIN) 13 taxa were included. [not seen]. All characters previously mentioned in the liter- ature have been used in the analysis, with the ex- Diagnosis ception of the relative length of the lateral scutellar Richter (1976) compared her species with L. brevise- seta (see below). All the characters newly proposed ria,notwithL. lacustris. Based on my translation of in this article have also been included. In total, 29 the original description (confirmed by V. Richter in discrete characters have been included, most of them litt.), L. columbina is very similar to L. lacustris,to binary (Table 1). The data matrix used is presented in which it keys out in the key above. The difference Table 2. The characters were equally weighted. The found on basis of the description is that the bands of characters for the non-Palaearctic species are based grey pruinescence occupy more than half of the ter- on descriptions in the literature (Townsend 1916, gites, whereas they are narrower in the female of L. 1927; Aldrich 1932; Mesnil 1968). Especially for the lacustris. Moreover, the colouration of the pruines- new characters, these are incomplete (question marks cence is bluish in L. columbina. in Table 2). Table 3 shows the values for the relative length of Distribution the lateral scutellar seta, as compared with the sub- The type locality is situated at the border of the apical one (as suggested by Herting (1959)). There great Uvs lake, east of Ulaangom. No other records is more or less a continuum of values and splitting known. these in separate groups can only be done in an arti- ficial way. Therefore, the relative length of the lateral Phylogenetic analysis scutellar seta has not been used. To study the phylogeny of Lydella (s.l.) (Woodley The analysis yields 8 most parsimonious clado- 1994), a parsimony analysis was carried out. All grams, summarized in the strict consensus cladogram western Palaearctic members were included, together shown in Fig. 22. This strict consensus cladogram with the type species of all nominal genera currently has limited resolution. considered in synonymy with Lydella. As outgroup

Table 1. Characters used for the phylogenetic analysis.

1. Host: other (0), stem-boring Lepidoptera caterpillar (1). 2. Width of fronto-orbital plate relative to height of gena: <1 (0), between 1 and 1.5 (1), >1.5 (2). 3. Setae on facial ridge: restricted to lower half (0), present on upper half as well (1). 4. Height of genal dilation: more than half of height of gena (0), less than half of height of gena (1). 5. Third antennal segment of male: at most 3× as long as second (0), more than 3× as long as second (1). 6. Postocular setulae of male: short and nearly straight (0), long and bent at apex (1). 7. Fronto-orbital plates of male: densely hairy (0), virtually bare (1). 8. Arista thickened: 2/3th or less (0), more than 2/3th (1). 9. Second aristal segment: at most 4× as long as wide (0), more than 4× as long as wide (1). 10. Reclinate orbital setae: 2 pairs (0), 1 pair (1). 11. Proclinate orbital seta in male: absent (0), present (1). 12. Number of humeral setae: two (0), three (1), four (2). 13. Third posterior supra-alar: as strong as first (0), much weaker than first (1). 14. First posterior intra-alar seta: as strong as other intra-alar setae (0), much weaker than other intra-alar setae (1). 15. Posterior lappet of posterior spiracle: large (0), small (1). 16. Hairs on notopleuron: many (>5) present (0), absent or at most 5 present (1). 17. Number of STPL: two (0), three (1), four (2). 18. Apical scutellar setae: reduced, hairlike (0), strong (1). 19. Number of anterodorsal setae on mid tibia: one or two, if two, then the second reduced (0), two strong (1), three or more (2). 20. Claws and pulvilli in male relative to length of fifth tarsal segment: longer (0), shorter (1). 21. Length of fore tarsus: less than 1.15× as long as tibia (0), more than 1.15× as long as tibia (1). 22. Cell r4+5 in wing: open (0), with a petiole (1). 23. Second costal section ventrally: bare (0), hairy (1). 24. Excavation on syntergite 1 + 2: reaching hind margin (0), not reaching hind margin (1). 25. Median discal setae on tergites 3 and 4: absent (0), present (1). 26. Fields of specialized hairs at ventral surface of tergite 4: present (0), absent (1). 27. Hairs on tergites 3 and 4 in male: adpressed (0), erect or semi-erect (1). 28. Inner margin of calypter: round (0), angular (1). 29. Number of preapical setae on hind tibia: two (0), three (1).

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Two monophyletic groups can be distinguished below 60%. For the (Lydelloxenis + Metoposisyrops)- within Lydella: clade, Bremer support is 1; stability under bootstrap- 1). The clade that includes the type species of ping and jackknifing is at or below 70%. Metoposisyrops, Metagonistylum, Diatraeophaga Another main conclusion is that the monophyly as well as Lydella jalisco. It is supported by the of the groups Lydella (s.s.) and Lydelloxenis could following synapomorphy: neither be corroborated nor rejected. – one pair of reclinate orbital setae present. – I will refer to this clade as the ‘Metoposisyrops- clade’. Discussion 2). The clade consisting of the species of Lydello- The status and position of Lydelloxenis and Metopo- xenis and the Metoposisyrops-clade, supported by sisyrops and related taxa has been controversial. Mes- the following two synapomorphies: nil (1953, 1956) considered Lydelloxenis to be a valid – first posterior intra-alar seta small, much genus in the vicinity of Metagonistylum,butHert- weaker than other intra-alar setae (Mesnil ing (1959) included it in Lydella, as did later authors 1956). (Mesnil 1975, Woodley 1994, O’Hara 2010). Mes- – male without fields of specialized hairs on nil (1956) and Woodley (1984) considered Metopo- ventral surface of tergite 4. sisyrops to be closely related to Lydella and Lydello- I will refer to this clade as the ‘(Lydelloxenis + xenis, whereas Crosskey (1976) suggested it was ‘very Metoposisyrops)-clade’. distinctive among the Eryciini’ and it might even be Both clades found have low support. For the considered as a separate tribe Metoposisyropsini. Metoposisyrops-clade, the Bremer support is 2; sta- Based on the phylogenetic analysis presented bility under bootstrapping and jackknifing is at or above, Metoposisyrops, Metagonistylum, Diatraeophaga as well as Lydella jalisco form a monophyletic group Table 2. Character matrix for the phylogenetic analysis. in Lydella (s.l.), though the support is relatively weak. Thesameistrueforthe(Lydelloxenis + Metoposisy- Drino inconspicua 00001000000200002100000000010 rops)-clade. This result supports the first opinion of Lydella stabulans 10001000000200002120001010100 Lydella grisescens 10001001000200002121001010000 Mesnil (1956). Lydella ripae 10001001000200002121001010000 The status of some characters is poorly known to Lydella thompsoni 10001000000200002121001010000 me for taxa in the Metoposisyrops-clade. This reduces Lydelloxenis lacustris 11010110001211011101101011000 the quality of the results of the phylogenetic ana- Lydelloxenis slavonica 11010110000201012100101011000 Lydelloxenis breviseria 11010110000111011011100011001 lysis. For instance, if the fore tarsus would be short Metoposisyrops sesamiae 12??1??11100??0?010??011???0? in the Metoposisyrops-clade, Lydelloxenis would prove Metoposisyrops oryzae 120????10100??0?1001100?1??0? to be monophyletic with the elongated fore tarsus Diatraeophaga striatalis 120?1??11110111?0001010?0??0? as synapomorphy. Since this article focuses on the Metagonistylum minense 12011??11?01110?20?1?0?1???0? Lydella jalisco 101010011111?01?1001?1?101?0? Palaearctic region, study of species from other re-

Figs 20, 21. Lydella thompsoni, male, from The Netherlands. 20, head, lateral view; 21, abdomen, lateroventral view.

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Table 3. Length ratio of lateral scutellar seta versus Lydelloxenis-Metoposisyrops clade is monophyletic and subapical scutellar seta for males of Lydelloxenis (left) supported by two synapomorphies. Lydella slavonica and Lydella (s.s.) (right), in ascending order. Ratios tend belongs to this clade. The monophyly of the Metopo- to be slightly lower in females. sisyrops-clade, containing all tropical and subtropical slavonica 0.47 ripae 0.54 species of Lydella, could be established as well. The lacustris 0.51 stabulans 0.57 support for both clades is not very strong. The ana- breviseria 0.63 thompsoni 0.69 lysis keeps open the possibility of the presence of two grisescens 0.85 other clades, i.e. Lydella (s.s.) and Lydelloxenis.For practical reasons and nomenclatural stability, I pro- pose to use Lydelloxenis and Metoposisyrops at the sub- generic level.

Acknowledgements First and foremost, I thank Gerard Pennards (Amers- foort) for collecting the new species at Skadarsko Jezero and also the other members of the Dutch– German expedition to Serbia and Montenegro in 2009, of whom Axel Ssymank (Bonn) provided the photographs of the habitat at Skadarsko Jezero. Cezary Bystrowki (Warsaw) and Peter Tschorsnig (SMNS) drew my attention to the females of L. slavonica from Poland and made them available for study. Peter Sehnal (NMW) kindly arranged a loan of the types of Lydella lacustris. James O’Hara (Ot- tawa) kindly examined a syntype of L. breviseria at my request. Norman Woodley (Washington) helped with relevant literature on Lydella. Vera Richter (St Petersburg) kindly informed me on Lydelloxe- nis columbina. Hans Duffels (Naturalis Leiden) con- Fig. 22. Strict consensus tree for Lydella (s.l.). Black tributed with useful suggestions on the manuscript. square: synapomorphy supporting a clade. Outgroup The manuscript did benefit from the work of two taxon: Drino inconspicua. Right margin: group names reviewers. Herman de Jong (Naturalis Leiden) and as used in the text. Menno Reemer (EIS-NL Leiden) helped me with TNT. This program was made available with the gions is out of the scope and this awaits further stud- sponsorship of the Willi Hennig Society. ies. Using the names Metoposisyrops and Lydelloxenis at subgeneric level might be unjustified, given the possi- References ble paraphyly of Lydelloxenis and Lydella (s.s.). How- Aldrich, J.M., 1932. New Diptera, or two-winged flies ever, this paraphyly has not been established and it from America, Asia, and Java with additional notes. – seems useful to have existing names available for the Proceedings of the United States National Museum 81: three groups Lydella (s.s.), Lydelloxenis and Metopo- 1–28. sisyrops. One could opt for raising the (Lydelloxenis + Bystrowski, C., 2005. Materials to the knowledge of Ta- Metoposisyrops)-clade to generic level. Given the cur- chinid flies (Diptera: Tachinidae) of the Biebrza Valley. rent uncertainties, I propose to be conservative and – Dipteron 21: 3–4. to use Lydella (s.s.), Lydelloxenis and Metoposisyrops at Cerretti, P., 2005. Revision of the West Palaearctic species of the genus Pales Robineau-Desvoidy (Diptera: Ta- subgeneric level. This analysis did not find support chinidae). – Zootaxa 885: 1–36. to separate Metagonistylum or Diatraeophaga from Cleare, L.D., 1939. The Amazon Fly (Metagonistylum mi- Metoposisyrops, not even at a subgeneric level. nense, Towns.) in British Guiana. – Bulletin of Ento- mological Research 30: 85–102. Crosskey, R.W., 1976. A taxonomic conspectus of the Conclusions Tachinidae (Diptera) of the Oriental Region. – Bulletin The newly described Lydella slavonica exhibits clas- of the British Museum (Natural History), Entomology, sical morphological characters of both Lydella and Supplement 26: 1–357. Lydelloxenis. A phylogenetic analysis shows that the

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