A new Palaearctic species of the subgenus Lunatipula (Diptera, Tipulidae) from the West Caucasus with a survey of the caucasica species group

Abstract The caucasica species group in the subgenus Lunatipula is redefined and now consists of five species native to the Caucasus. Tipula (L.) eleniyasp. nov. is described as new to science, and variations in the male terminalia in two populations are noted. Two subspecies (quadridentataquadridentata and quadridentatapaupera) are elevated to species rank. Detailed photo’s complement the descriptions of all five species (caucasica, eleniya, paupera, quadridentata, talyshensis), and data on ecology and distribution patterns are included as well as identification keys to males and females. Tipula caucasica is recorded from the West Caucasus and Tipula quadridentata is recorded from Dagestan (Russia) for the first time. Parallel evolution is traced in the male terminalia of the new species and in several non caucasica species group of Palaearctic Lunatipula.


Introduction
There are 502 species in the subgenus Lunatipula Edwards in the world fauna (Oosterbroek 2021). This is the dominant taxon of crane flies in the Caucasus (Savchenko 1964(Savchenko , 1983, comprising at least 53 known species, which is ca. 32% of the total spe-cies of the family in this region (164). Riedel (1920) pioneered the work on crane flies of the Caucasus and neighboring regions, especially on the subgenus Lunatipula. He provided information on seven species of Lunatipula, three of which were described as new to science: Tipula (Lunatipula) caucasica Riedel, Tipula (Lunatipula) armata Riedel (synonym of zaitzevi Savchenko, 1952), and Tipula (Lunatipula) aurita Riedel. Savchenko (1954, 1957, 1964, 1968) made subsequent invaluable contributions to the knowledge of the regional fauna of the subgenus by describing 57 species, of which 32 were from the Caucasus and only two of these were subsequently synonymized (Oosterbroek 2021). The species group caucasica was first proposed by Savchenko (1964) to comprise four species endemic to the Caucasus. A number of subsequent studies have been published since on the diversity and ecology of some species of the subgenus Lunatipula from the Caucasus (Lantsov 1998(Lantsov , 2002(Lantsov , 2007(Lantsov , 2018. In this study we present new data on the taxonomic status, morphology, and distribution of the caucasica species group, provide a key to all currently known species, and describe Tipula eleniya Lantsov & Pilipenko as new to science. It was found in the Western Caucasus (Krasnodarskiy kray) at altitudes of 1200-2000 m. The new species is characterized by a projection on the inner gonostylus of the male, the structure of this shape is not known in any of the 502 known species, including the 360 Palaearctic species.

Materials and methods
All crane flies were collected by sweep-net and then pinned. The genitalia were macerated in warm 10% KOH for ca. one hour to remove soft tissue, and then rinsed in distilled water. Cleared genitalia are preserved in glycerol in micro vials pinned with their respective specimens. Specimens were studied with an Olympus SZ61 stereo microscope. A Nikon d7000 digital camera equipped with combined Tamron 70-300 / 4-5.6 and EL-Nikkor 50/2.8 lenses or Mitutoyo M Plan Apo 10X Microscope objective lens was used to capture partially focused images of each specimen or structure. These were stacked using the Helicon Focus (version 7.6.4) software (http://www.heliconsoft. com/heliconsoft-products/helicon-focus). Photographs of lateral and dorsal view and details of the structure of head, pronotum and scutum were made with a Canon 5D Mark IV digital camera equipped with a Canon MP-E 65mm f/2.8 1-5 × macro lens and Canon Macro Twin-Lite MT-26EX-RT flash. Adobe Photoshop CC 2019 software was used to edit the pictures. Measurement were made with an MBS-10 microscope with a scale installed in the focal plane of the 8× eyepiece (divisions of scale: 0.1 mm at 8 × 1 magnification, 0.05 mm at 8 × 2 magnification, 0.025 at 8 × 4 magnification).
For citing label data on type specimens, a slash / separates each label. Square brackets [ ] are used to indicate additional information not on the original label. Original spelling is retained, including punctuation. In some cases, holotypes were marked by a red label without text and some paratypes were not initially marked. In such cases, specimens with locality labels corresponding to those in the publications of Savchenko (1964) or Savchenko and Kandybina (1987), were provided with a red label with the word "Holotype" or a white label with the word "Paratype," and pinned with the specimen. These type specimens are deposited in the Zoological Institute of Russian Academy of Science, St. Petersburg, Russia. The holotype and paratypes of the Tipula (Lunatipula) caucasica were unavailable for this study.
We generally follow the terminology of Cumming and Wood (2017) except that for head morphology we follow Crampton (1943) and Savchenko (1983), and regarding wing venation, we follow Savchenko (1983). Veins are indicated by capitals; cells are indicated by lowercase letters.
All measurements of the new species were made on pinned material. Species distributions are given according to Oosterbroek (2021). The original descriptions and measurements by Riedel (1920) and Savchenko (1964)  The caucasica species group. Diagnosis (after Savchenko 1964 with additions). Medium sized grey species. Rostrum with nasus, wings with more or less distinct bluish tint in transmitted light; metakatepisternum with setae; abdomen slate or brownish grey with lateral intermittent dark brown stripe on tergites. Males with tergite 9 transverse with two or three concave notches at apex; inner gonostylus posteriorly simple; sternite 8 with paired appendages composed of short wide base with bristles and an elongate glabrous spine bent medially; no dense brush of setae between bases of paired appendages (except for T. quadridentata). Caucasian endemics. Savchenko included the following species in the caucasica group: caucasica, quadridentata quadridentata, quadridentata paupera, talyshensis. Diagnosis. Dark stripes medially on lighter background extending entire length of scutum. Inner gonostylus posteriorly with wedge-shaped projection directed anteriorly. Tergite 9 at apex with three rounded notches, middle one usually deepest and widest.
Head. Light grey, sometimes bluish (Figs 1B, 13A). Rostrum whitish dorsally, flanks rusty or brownish with whitish bristles; nasus distinct, with long, whitish yellow setae; palpus grey. Frons with rows of brown setae along inner edge of eye. Vertex with faint dark median line, posteriorly with dark grey striation, with group of small rusty setae arranged fan-like on either side of glabrous medial area. Genae rusty; wide dark median stripe ventrally on rostrum, continuing along gula midline. Pharyngeal foramen framed with long, rusty hair-like setae. Postgenae light grey.
Antenna (Fig. 1B) 13-segmented, if bent backwards just reaching wing base. Scape light brown or grey, depending on angle of rotation to light source, with slightly pronounced silvery pruinescence and brown horizontal striations. Pedicel yellow, sometimes brownish at base. Flagellomeres brownish, thickened at base, verticils subequal in length to corresponding flagellomeres. Thorax (Fig. 1B). Grey or blue-grey. Pronotum with dark grey median stripe. Katepisternum, anepisternum and meron grey, glabrous. Scutum with four longitu- dinal wide brownish stripes on lighter background; medial stripes extending length of scutum. Scutal lobes, scutellum and mediotergite grey with light bristles in dark brown sockets. (Fig. 1A) greyish with a bluish tinge in transmitted light, stigma brown; narrow whitish area (lunule) extending to base of discal cell.

Wings
Halteres ( Fig. 1A) with light brown or dark-yellow stem, lighter yellow at base, covered with short small pale setae; knob dark brown to black.
Legs with coxae light grey with long light hairs. Coxae may appear darker when specimen is rotated. Trochanters yellow. Femora yellow at base, rest of femora, tibiae, and tarsi dark brown.
Abdomen (Figs 1A,2A). Grey or dark grey; tergites with dark lateral stripe narrowly interrupted with whitish at posterior margins, lateral margins with wider whitish edge.
Female. Body length 17-21.5 mm, wing 15.5-18 mm. Antennae with scape and pedicel brownish yellow. Legs stocky, black to brown, yellow at base, femora grey. Wings with noticeable bluish tint; whitish area (lunule) of wings barely reaching base of discal cell. Female terminalia ( Fig. 12A-C) comparatively short. Tip of cercus upcurved, length of cercus ~ 0.8 mm. Hypogynial valve only slightly longer than width at base; length of hypogynial valve 0.2 mm. Sternite 9 and furca as in Figure 12A, C.
Comparison with closely related species. This species differs from other species of the caucasica group primarily by the presence of a large, anteriorly directed sharp wedge-shaped projection located posteriorly on the inner gonostylus.
Elevation. Adults were collected at altitudes ranging from sea level (Black Sea coast at 35 m) to 2011 m (in Dagestan).
Flight period. From end of April to the beginning of July.
Habitat. Specimens were captured in diverse forest habitats (see above) predominantly in mesophytic moderately moist deciduous or mixed communities with Fagus orientalis, Taxus baccata, Carpinus orientalis, Quercus iberica, Populus tremula, Acer platanoides, etc., sometimes in communities near springs with Salix caprea.
Distribution. Endemic to the Caucasus. It was recently noted for Russia (Dagestan) for the first time (Lantsov 2020). Currently known from the West Caucasus (first record), East Caucasus and Transcaucasus.
Remarks. According to Riedel (1920), a series of type specimens, four males, two females. "Mts'chet prope Tiflis [Georgia, Tbilisi], 6. 5. 1913 (Mus. Caucas. Ph. Za- (Lunatipula) caucasica A terminal segments of male abdomen, lateral view (dry) B-L male terminalia (after KOH 10% treatment) B hypopygium, caudal view C hypopygium, lateral view D sternite 8, ventral view E tergite 9, caudal view F tergite 9, dorsal view G inner and outer gonostylus, lateral view H inner and outer gonostylus, caudal view I gonocoxal fragment, dorsal view J semen pump and aedeagus, lateral view K adminiculum and paired apical appendages of sternite 9, caudal view L adminiculum and paired apical appendages of sternite 9, lateral view. Abbreviations: adm-adminiculum; as 8-paired appendages of sternite 8; as 9-apical appendages of sternite 9; ig-inner gonostylus; gcx-gonocoxite; og-outer gonostylus; s8-sternite 8; s9-sternite 9; t9-tergite 9. Scale bar: 1 mm. itzev)" were deposited in the "Museum Caucasicum in Tiflis". In the author's publication, neither the holotype nor the paratypes are indicated. Presumably, they are not indicated on the material itself. Because this material turned out to be inaccessible to the authors of this article -it is not yet known whether this material has survived at all -there was no opportunity to designate lectotype and paralectotypes. Diagnosis. Male. General coloration grey with silver pruinescence. Tergite 9 with deep rounded notches either side of slightly grooved cone-shaped projection. Sternite 8 with a pair of small appendages, each bearing a long medially curved spine, broad base of appendages with fringe of long whitish hairs. Outer gonostylus small, triangular, slightly thickened distally, covered with long setae. Inner gonostylus with small rodlike outgrowth in middle of outer edge.
Description. Head (Figs 3, 13B) grey with silvery pubescence. Rostrum grey dorsally, ventrally yellow-brown with sparse light rusty setae in black sockets. Nasus distinct, appearing triangular in dorsal aspect, covered with long, sparse procumbent setae. Frons and gula bare. Vertex (Fig. 13B) framed by rows of sparse setae alongside eye, glabrous in center with indistinct narrow longitudinal dark grey line. Genae, including lateral part of rostrum, with very sparse fine pale bristles. Tempora and postgenae with sparse black setae.
Antennae, bent backwards, reaches base of abdomen. Scape grey with silvery pruinescence, pedicel dirty yellow brown. Flagellomeres dark brown with verticils on slightly thickened bases, longest verticils subequal to length of respective flagellomere.
Thorax (Fig. 3A, B) grey with silvery pruinescence, and short sparse whitish hairs. Pronotum dark brown with a weak dark stripe medially, and short sparse whitish bristles. Scutum (Fig. 3B) with four brownish grey longitudinal stripes, lateral stripe shorter, medial stripes narrowly separated by dirty yellow area. Katepisternum, anepisternum, katepimeron, anepimeron and meron grey, bare, with silvery pruinescence; scutellum and mediotergite light grey with silvery pruinescence; scutal lobes with wide dark grey stripe, bordered by sparse whitish setae and with long yellow setae anterolaterally at wing base. Scutellum framed anteriorly with brown, with sparse yellow setae laterally, and with vague narrow dark grey medial line. Mediotergite light grey with sparse yellow setae and with silvery pruinescence. Laterotergite grey with silvery pruinescence and sparse erect short brown setae.
Wings (Fig. 3A) with typical venation for the subgenus, transparent with brownish tin, (viewed in transmitted light). Stigma dirty yellow. Oblique lunule proximal to stigma. Setae on costal and subcostal veins; group of short black macrotrichia distally on R and base of R 1 . Distal section of R s , R 4 + 5 , r-m, base of pentagonal discal cell lighter and weakened. Stem of cell m 1 approximately half length of cell. Apex of vein Cu 1 sharply curved at wing margin.
Halteres. Stem grey with base covered with short white setae, knob dark grey. Legs. Coxae light grey with silvery pruinescence; laterally with long sparse whitish yellow setae; medially with very short yellowish setae. Trochanters yellow with long yellow setae. Femora brown, narrowly yellow at base, covered with adpressed short brown setae; tibiae brownish; tarsal segments dark brown with short, adpressed brown setae. Tibial spur formula 1-2-2. Tarsal claws with a small tubercle at the base (magnification 10 × 4.5); with short but clearly visible arolium between ones.
Abdomen (Figs 3A, 4A). Tergites and sternites grey with silvery pruinescence, and sparse fine short golden setae. Tergites with lateral dark grey intermittent stripe, no distinct dorsal stripe. Tergites laterally and distally edged with whitish-yellow, proximal segments with thinner distal edge, becoming wider on caudal segments.
Distribution. Endemic to the Caucasus: currently known from the West Caucasus. Etymology. Tipula (Lunatipula) eleniya sp. nov. is named after the mother of the first author, Elena Nikolaevna Lantsova. Diagnosis. Male. Gonocoxite with two elongate, pointed teeth, one dorsally and one ventrally. Tergite 9 with two projections posteriorly, separated by deep wide notch. Paired appendages of sternite 8 widely spaced, base shorter than wide, gap between not masked by setae. Apical appendages of sternite 9 elongate, narrow distally, with dense bundle of relatively short golden yellow setae at tip.
Head (Figs 6B, 13C). Rostrum dorsally light grey to whitish with silvery pruinescence, dark brown procumbent setae, base of rostrum dorsally light beige, laterally dirty rusty, yellow. Nasus well defined with longer whitish procumbent setae. Palpus dark brown. Vertex (Fig. 13C) light sandy color with soft silvery pruinescence, dark grey thin median line extending to occiput; nine or ten short brown setae, behind antenna, four or five longer brown setae alongside eye. Row of long brown setae on temporal region and around occipital foramen. Gula and gena (including rostrum ventrally) yellowish-sandy to light brown, with medially enfolded region ventrally marked by moderately broad, light brown median line; gena near eyes light yellow; postgena yellow to sandy.
Antennae. Scape light grey with silvery pruinescence, pedicel yellow with subtle brown ring at base. Flagellomeres brown.
Thorax (Fig. 6A, B). Pronotum grey with dark broad median stripe with blurred edges. Scutum with four brown stripes, central pair more pronounced, widener anteriorly with dirty yellow between. Scutal lobes brown with silvery pruinescence. Scutellum and mediotergite with thin fuzzy grey median line; scutellum yellow to light grey; mediotergite light grey with silvery pruinescence and scattered whitish short bristles. Katepisternum, anepisternum, katepimeron, anepimeron and meron light grey with silvery pruinescence, glabrous.
Wings. Transparent, without noticeable marble pattern, with light brown stigma. Longitudinal veins with macrotrichia.
Halteres. Stem light brown to dirty yellow, knob brown. Legs. Coxae light grey with silvery pruinescence and long, whitish bristles; trochanters light brown; femora yellow at base, light brown with darkened tips, with adpressed dark brown bristles.
Female. Unknown. Status. The species was described and treated as a subspecies of T. quadridentata. It is elevated here to species rank because of the presence of a number of differences of the species from T. quadridentata and all other species of the caucasica species group (see below).
Comparison with closely related species. This species differs from all other species of the caucasica group by the number and arrangement of setae dorsally on the head (Fig. 13C), the deep median notch at the apex of tergite 9 (Figs 7F, 14C), and the more widely spaced appendages of sternite 8 (Fig. 7D). It differs from T. eleniya sp. nov., T. quadridentata, and T. caucasica in the shape and presence of the dense bun- (Lunatipula) paupera A terminal segments of male abdomen, lateral view (dry) B-L male terminalia (after KOH 10% treatment) B hypopygium, caudal view C hypopygium, lateral view D sternite 8, ventral view E tergite 9, caudal view F tergite 9, dorsal view G inner and outer gonostylus, lateral view H inner and outer gonostylus, caudal view I gonocoxal fragment, dorsal view J semen pump and aedeagus, lateral view K adminiculum and paired apical appendages of sternite 9, caudal view L adminiculum and paired apical appendages of sternite 9, lateral view. Scale bar: 1 mm. dle of short setae on the appendages of sternite 9 (Fig. 7K). This species is similar to T. quadridentata in having two teeth on the gonocoxite (Figs 7G, 14I, J).
Elevation. The holotype was probably collected at 2623 m (height of Zagar Pass) and one of the paratypes was collected "on the descent from the Bassa Pass", the height of which is 3057 m. It can be assumed that this species occurs in high-mountainous habitats.
Flight period. Adults were collected from the last third of July to early August.
Halteres. Stem light brown to yellowish, knob brown. Legs. Femora light brown with procumbent dark brown bristles. Tarsal claws without noticeable tooth at base.
Terminalia (Figs 9, 14D, J). Hypopygium not thickened. Tergite 9 and gonocoxite with silver pruinescence. Upper margin of gonocoxite with two dentate projections. Tergite 9 (Figs 9E, F; 14D) at apex with four widely spaced teeth and with three rounded notches, narrower middle notch bounded by small flat projection bearing sharp thorn beneath; small backwards projecting spine beneath tergite 9. Outer and inner gonostyli (Figs 9G, H; 14J) without significant features. Paired appendages of sternite 8 (Fig. 9D) large, wide base with fringe of thick yellow setae hiding gap between (apparent on pinned specimens). Paired apical appendages of sternite 9 as in Figure 9K, L. Gonocoxal fragment (Fig. 9I) with characteristic small V-shaped fork at base. Semen pump and aedeagus as in Figure 9J.
Comparison with closely related species. This species differs from other species of the caucasica species group (in the male) by tergite 9 with four widely spaced teeth and with three small notches at the apex and a small spine beneath (Figs 9E, F; 14D). Paired appendages of sternite 8 is broadest, with thick yellow bristles that cover the gap between; in the other species this gap is not obscured. Tipula quadridentata is similar to T. paupera by having two teeth on the edge of the gonocoxite and similar coloring of the antenna. The outer gonostylus is somewhat larger than in T. caucasica. The gonocoxal fragment is similar to that of T. talyshensis (Figs 9I, 11I). The female has a parallel-sided, straight cercus whereas it is upturned at the tip in T. caucasica and broader at base in T. talyshensis.
Flight period. Adults were collected from throughout the month of May.
Habitat. Specimens are found in moderately humid woody deciduous communities. Distribution. Endemic to the Caucasus; currently known from the West Caucasus (northern and southern slopes; Krasnodar and Stavropol Territory) and from the East Caucasus (Dagestan; first record). B-L male terminalia (after KOH 10% treatment) B hypopygium, caudal view C hypopygium, lateral view D sternite 8, ventral view E tergite 9, caudal view F tergite 9, dorsal view G inner and outer gonostylus, lateral view H inner and outer gonostylus, caudal view I gonocoxal fragment, dorsal view J semen pump and aedeagus, lateral view K adminiculum and paired apical appendages of sternite 9, caudal view L adminiculum and paired apical appendages of sternite 9, lateral view. Scale bar: 1 mm. Savchenko, 1964 Figs 10, 11, 12G-I;13E, 14E, K Tipula (Lunatipula) (Fig. 10A, B), however, some legs missing: fore legs -left missing, right up to femur; mid legs -left present, right missing; hind legs -left and right up to femur. Paratypes. Azerbaijan •2 males, 2 females; same data as for holotype [printed on white paper] / "Paratypus" [Paratype not initially marked; red label]; ZISP. Diagnosis. Male. Tergite 9 at apex with a denticle on either side of wide, shallow, flat median notch. Paired apical appendages of sternite 9 narrower distally, bearing tassel of long golden setae. Female. Hypogynial valve several times longer than width at base.
Head (Fig. 13E). Vertex light grey with indistinct median grey line, glabrous in middle and around eye with rows of setae between. Rostrum dorsally dark grey, whitish at base, ventrally light brown with grey median stripe. Nasus with long, whitish procumbent setae. Tempora and genae light grey to whitish (color depending on angle of inclination) with faint silvery pruinescence.
Antennae. Scape dark grey with rusty bristles, pedicel yellowish with indistinct brown line in middle, flagellum brown. Thorax (Fig. 10A, B). Pronotum grey with whitish setae. Scutum ( Fig. 10B) with four brownish stripes on grey, short whitish setae between central and side stripes. Scutal lobe grey with faint silvery pruinescence, anterolaterally with group of whitish setae. Pleura and coxae grey with silvery pruinescence; katepisternum dorsally with sparse whitish setae, anepisternum and meron glabrous. Scutellum and mediotergite grey with whitish setae, scutellum with dark grey median line visible as specimen is rotated.
Wings (Fig. 10A). Translucent, grey, with light brown pterostigma. Longitudinal veins C, Sc, and R and bases of A 1 and A 2 with macrotrichia.
Halteres. Stem yellowish with light setae, knob brown. Legs. Coxae grey with silvery pruinescence and long, whitish setae. Femora light brown with procumbent dark brown setae. Claws of fifth tarsal segment without spine at base.
Abdomen (Figs 10A, 11A). Dark grey with short whitish setae. Posterior margins of tergites without noticeable light edging, lateral margins with wide whitish to yellowish edging.
Comparison with closely related species. This species differs from other species of the caucasica group by the broad shallow notch at the apex of tergite 9, the tassel of golden setae at the tip of the apical appendage of sternite 9, and by the small size of the   Inner gonostylus with rod-like outgrowth behind anterior beak (Fig. 14G) Wedge-shaped projection posteriorly on inner gonostylus (Fig. 14F)  Anterior (dorsal) margin of gonocoxite with sharp dentate projection in addition to smaller more ventral tooth (Fig. 14 I, J). Adminiculum and paired apical appendage of sternite 9 as in Figure 7K (Fig. 14K). Apical appendage of sternite 9 narrowed distally, bearing tassel of long golden setae (Fig. 11K, L) ...... T. talyshensis Savchenko, 1964 4 Tergite 9 at apex with four projections, central notch between middle projections small and shallow (dorsal view) (Fig. 14D). Paired appendage of sternite 8 with thick yellow setae on base covering space between. Setal arrangement on vertex as in Figure 13 D. Adminiculum and paired apical appendages of sternite 9 as in Figure 9K ..................... T. quadridentata Savchenko, 1964 -Tergite 9 at apex with only two projections, central notch between quite wide and deep (dorsal view) (Fig. 14C). Gap between paired appendage of sternite 8 usually exposed, not masked by setae. Arrangement of setae on vertex as in Figure 13C. Adminiculum and paired apical appendage of sternite 9 as in Figure  1 Hypogynial valve long, more than three times longer than width at base (Fig. 12G, H)

Discussion
All species belonging to the caucasica species group are morphologically quite distinct including T. quadridentata and T. paupera, previously considered as subspecies.
Their status as separate species is beyond doubt: each is distinguishable by the structure of tergite 9 as well as additional distinctive features mentioned in the descriptions and comparisons above, as well as in the keys. For T. caucasica, this is a characteristic wedge-shaped projection posteriorly on the inner gonostylus (Fig. 14F); for T. eleniya, it is a unique rod-shaped outgrowth of the inner gonostylus (Fig. 14G); T. talyshensis differs by the unique structure of the paired appendage of sternite 9 (Fig. 11K). The inner gonostyli of T. quadridentata, T. paupera, and T. talyshensis are similar in lateral view (Fig. 14I, J, K), but these species differ in size, shape, and coverage of the paired appendages of sternite 8 (Figs 7D, 9D, 11D). T. eleniya, T. quadridentata, and T. talyshensis are similar in the structure of the gonocoxal fragment, semen pump, and aedeagus (Figs 4I, J; 9I, J; 11I, J). The color and arrangement of the setae on the vertex of the head in T. paupera distinguishes it from all other species of the group (Fig. 13C). The variability seen in the outgrowth of the inner gonostylus of T. eleniya (Fig. 5) probably indicates the plasticity of this species and confirms the need for additional studies. Females of only three of the five species of the caucasica species group are known, and they differ sufficiently in the structures of the cerci, hypogynial valves, and sternite 9 (Fig. 12), as reflected in the identification key. Of interest would be the immature stages of the species of the caucasica species group, about which nothing is yet known.
Zoogeographically, the caucasica species group belongs to the Caucasian subgroup, a part of the eastern Mediterranean group, which is, in turn, a part of the Mediterranean species complex (Savchenko 1983). These species are narrow-range endemics to the Caucasus. Ecologically, they are confined to mesophytic habitats within the forest belt, occurring in a fairly wide range of altitudes and can be classified as species characteristic of deciduous forest and mixed communities.
Parallel morphology in the subgenus Lunatipula in the structure of internal gonostylus in Palaearctic species The inner gonostylus of T. (L.) eleniya was compared with those of the 502 known species including the 360 Palaearctic species of the subgenus Lunatipula. No direct matches were found, but outgrowths of various shapes on the middle of the inner gonostylus were found in some Palaearctic species: two species from Turkey (Tipula (Lunatipula) auriculata Mannheims, 1963 and Tipula (Lunatipula) horsti Theischinger, 1982), one from China (Tipula (Lunatipula) oreada Alexander, 1933), and two from Kyrgystan (Tipula (Lunatipula) milkoi Pilipenko, 2005 and Tipula (Lunatipula) zarnigor Savchenko, 1954). The latter species has also been recorded in Tajikistan and northeastern Afghanistan (Oosterbroek 2021). The species belong to the same sub-genus, but are classified in different species groups and are geographically dispersed in this regard, it can be assumed that these various outgrowths to be independently derived and therefor examples of parallel evolution.