Review of the Palaearctic members of the Lispe tentaculata species-group (Diptera, Muscidae): revised key, synonymy and notes on ecology

Abstract The taxonomic reasons for regarding Lispe draperi Séguy, 1933, sp. rev., as a valid species instead of a synonym of Lispe tentaculata (De Geer, 1776) and for treating Lispe quaerens Villeneuve, 1936, syn. n., as a junior synonym of Lispe sericipalpis Stein, 1904 are given. A revised key for the Palaearctic members of the Lispe tentaculata species-group is given. Data on ecology, distribution and feeding preferences are provided.


Introduction
Th e Lispe tentaculata species-group was proposed by Hennig (1960) and is characterized by the meron setulose above hind coxa and the following leg chaetotaxy: t1 without setae; t2 only with p-seta, without ad or av, t3 with 1ad and 1 weak pd, without av.
Currently eight species are placed in the Lispe tentaculata species-group, six of which are present in the Palaearctic region. Th e taxonomic status of Lispe alpinicola Zhong, Wu & Fan, 1981 has not yet been settled (see below). Lispe sericipalpis Stein, 1904 andL. orientalis Wiedemann, 1824 are also distributed in the Oriental region. Lispe tentaculata (De Geer, 1776) is recorded from the very north of the Oriental region and widespread in Nearctic. Two more species of this group are found in the Nearctic region.

Material and methods
Th e majority of the specimens studied are in the Zoological Museum of Moscow University (not indicated in text). Other material is in Natural History Museum, London (BMNH), Zoological Institute, St Petersburg (ZIN), Zoölogisch Museum, Universiteit van Amsterdam (ZMAN).
Here I suggest a new abbreviation for the tarsi as tar followed by a pair of digits separated by a hyphen: the fi rst digit (1 to 3) gives the leg number and the second digit (1 to 5) the number of the tarsal segment. For example, tar1-4 = 4-th segment of fore tarsus; tar3-1 = hind basitarsus.
Identification key for Palaearctic species of the Lispe tentaculata speciesgroup Fore tarsus modifi ed: tar1-1 yellow to dark, tar1-2 to tar1-4 yellow, tar1-5 black; tar1-1 half as long as tar1-2, with a dense row of brush-like av setulae and on posterior side with a fi nger-like yellow process with black apex, this process reaching middle of tar1-2; tar1-2 projecting ventrally. Male cercal plate as Fig. 1  Median third of f3 with 2-5 av setae 1.5-2 times longer than femoral width. tar3-1 shortened (slightly more than one third as long as t3); ventral median part of tar3-1 concave; basal 1/3 of tar3-1 with a brash of ventral hairs contrasting with shorter hairs in apical 2/3. Scutellum with some fi ne hairs below at apex (these hairs may be invisible in old specimens). Cercal plate with 2 pairs of projections apically, distinctly longer than wide ( Fig. 1.2 Median 1/3 of f3 without long av setae (though in basal 1/3 of f3 with 3-4 av subequal to femoral width). tar3-1 straight, not curved or concave, tar3-1 longer (slightly less than half as long as t3); ventral hairs on tar3-1 of uniform length. Scutellum bare below at apex and bare at apex laterally below apical scutellar bristles. Cercal plate with only 1 pair of projections apically, almost as wide as long ( Fig. 1.1). Usually 4 post dc setae, anterior 2 post pairs short and fi ne (but sometimes 3 strong post dc as in tentaculata!    Scutellum bare below at apex. t2 and t3 yellow, more or less dusted. 2+4 dc, but the anterior 2 post pairs short and fi ne. f3 in median 1/3 without av 1.5-2 times longer than femoral width (but 1-3 av setae subequal to f3 width present at basal 1/3 of f3 Remarks. Described from Lhasa (China, 3500-4000 m asl.). Male fore and hind basitarsus modifi ed as in L. tentaculata, female scutum with a pruinose patch as in L. tentaculata (Xue & Dong, 2005), male cercal plate similar to L. tentaculata (Xue & Chao, 1998). According to Xue and Dong (2005), males of L. alpinicola may be separated by the presence of a p-seta on t1 and 3 pd setae on t2. It is diffi cult to give any opinion on this species without making a personal examination of specimens, but I have some doubts about the validity of this species. Among the rich tentaculata material examined, there were no specimens with both p-seta on t1 and 3 pd setae on t2, but separately the presence of these setae was recorded. Several specimens from the north of Russia have a p-seta on t1. Remarks. Th e conspecifi ty of the material listed above with type of L. draperi Séguy was kindly confi rmed by A.C. Pont (pers. comm.) who examined the holotype of L. draperi in the Muséum national d'Histoire naturelle, Paris. Hennig (1960: 430) examined the type of this species and provisionally maintained it as a good species although he considered that the type might be an aberrant specimen of L. tentaculata. Canzoneri and Meneghini (1966) suggested that L. draperi is North African yellowish-legged subspecies of L. tentaculata but the species was sunk as a synonym of L. tentaculata by Pont (1986). Th is decision could be supported by the fact that the male cercal plate is similar to that of L. tentaculata. However, more careful examination shows that the structure of male terminalia diff ers and it is especially obvious in the structure of sternite 5 (Fig. 2). L. draperi should therefore be restored as a valid species, although closely related to L. tentaculata. I examined the sternite 5 of L. tentaculata collected in the Netherlands, Moscow region, Krasnodarsky Kray, Primorsky Kray (Russian Far East) and high in the Pamir mountains (Tajikistan, Gorno-Badakhshan, 3800 m asl.) and found it to be identical in all cases but diff erent from that in Moroccan L. draperi.  10°E, 20.XII.2006, N.Vikhrev, 1♀;Antalia prov., Silion ruins, 36.989°N 30.985°E, goat drinking bowl, 25.V.2008, N.Vikhrev, 1♂, 1♀;Hatay prov., Arzus env., 36.407°N 35.886°E, 14.IV.2010, N.Vikhrev, 8♂♂, 2♀♀;Hatay prov., Samandag env., Çivlek, 17.IV.2010 194°N 34.810°E, forest stream, 23.IV.2010, N.Vikhrev, 1♀;Bolu prov., 40.498°N 31.890°E, forest stream, 1800m asl., 31.VIII.2009Sarakya prov., Karasu env., 41.03°N 30.79°E, forest stream, 15.VI.2010, N.Vikhrev, 1♀, 28.VIII.2009Zonguldak prov., Alaply env., 41.14°N 31.36°E, forest stream, 21.VI.2010, N.Vikhrev, 3♂♂, 2♀♀. Remarks. Male. Ground colour black. Pollinosity grey, but may be yellowish-grey. Palpus black(ish), but becoming brown in old specimens. Fronto-orbital plate and parafacial whitish, rarely yellowish. Scutum with 3 brown vittae along ac and dc rows, submedian (dc) vittae sometimes almost indistinct. dc 2(1) + 4 (rarely 3), as: presutural: weak to hardly distinct, medium; postsutural: weak, weak, strong, strong. Presutural ac in 4-5 irregular rows. Legs dark. f2 with several (2-7) fi ne pv-setae in basal 1/3, the longest one (1.5 times as long as femoral width) at base. f3 at base with long (1.5-2 times as long as femoral width) and fi ne characteristic seta in v-pv position. Other setae on f3 variable: several av-setae present in apical third or slightly more, 1-3 among them longer, from as long as femoral width to twice as long; pv row may consist of 7-8 setae of which 2-3 are as long as femoral width, or may be reduced to 1-2 hardly distinct pv at base; sometimes chaetotaxy of f3 diff ers on right and left legs of the same specimen. Abdomen whitish-grey to yellowish-grey dusted, with paired Lshaped, more or less extensive dark spots on tergites 3 to 5, divided by dusted median vitta. Female diff ers from male as follows: parafacial more often yellowish; dc 1+4(3), anterior prst dc always absent; v-pv seta at base of f3 shorter to reduced; other ventral setae on f2 and f3 shorter or reduced.

Lispe orientalis
Th e type locality of L. sericipalpis is Indonesia, Java. Th e species has also been recorded from other Indonesian islands (Bali, Sumatra), Taiwan, Vietnam, Th ailand, Myanmar, Sri Lanka, India, Pakistan, Nepal. Th e type locality of L. quaerens is Turkey, Akshehir prov. Th is species, as interpreted by Hennig (1960: 453) who studied the holotype, has also been recorded from Spain, Italy, Croatia, several provinces of Turkey, Azerbaijan, Tajikistan and China. Th us, there appears to be no geographic gap between the natural habitats of L. sericipalpis and L. quaerens, but the former, known as an Oriental species, has never been compared with the latter, regarded as a Palearctic species. Th e examined series from Turkey and Tajikistan (some specimens were identifi ed by Hennig, 1960: 453) had been assigned to L. quaerens, whereas the series from the  Pont, 1970). Photo by Joke van Erkelens.
Remarks. L. tentaculata has a variable t3 chaetotaxy: besides the normal strong ad and short pd setae, an additional seta just below the strong ad but in a more a-position is often present and sometimes a second strong ad and a short pd may also be present. About 5% of females have a male-like scutum: with only 3 strong post dc and without a pruinose patch on the scutum.

Distribution and ecology
L. tentaculata is a very common species across its natural habitat. Th e northernmost specimens were collected near Murmansk (forest-tundra at 69°N) and Vorkuta (tundra at 67.5°N), where this species was found on boulders on river banks and was the only Lispe species recorded at these places. In southern Turkey at 36°N L. tentaculata was common again on boulders in streams and, in springtime, at various muddy places such as temporary pools. In temperate conditions of the East European Plain L. ten-

Feeding
It is well known that all Lispe are predators (e.g. Werner and Pont 2006). Actually there are two poles of this behaviour. Some species are skilful hunters of active Diptera, like Dolichopodidae, Ephydridae, Muscidae or Scathophagidae. Such a hunting style is typical for the Lispe caesia species-group (note that the characteristic ventral spines on f1 and f2 serve to grip the prey), another example is the Oriental Lispe geniseta Stein, 1909 which usually hunts on Musca on the cattle dung in pastures. Th e other type is feeding on invertebrate carrion, and a typical example is the Oriental Lispe binotata Becker, 1914 which sucks dead Arthropoda from forest streams, usually ants and spiders. Species of the Lispe tentaculata group represent an intermediate type as they feed either on dead insects (Fig. 6) or on living prey, but in the later case the prey is usually an insect larvae and less often slow moving imago of a group such as Chironomidae.