Research Article |
Corresponding author: Vladimir I. Lantsov ( lantsov@megalog.ru ) Academic editor: Netta Dorchin
© 2021 Vladimir I. Lantsov, Valentin E. Pilipenko.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lantsov VI, Pilipenko VE (2021) A new Palaearctic species of the subgenus Lunatipula (Diptera, Tipulidae) from the West Caucasus with a survey of the caucasica species group. ZooKeys 1048: 145-175. https://doi.org/10.3897/zookeys.1048.67564
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The caucasica species group in the subgenus Lunatipula is redefined and now consists of five species native to the Caucasus. Tipula (L.) eleniya sp. nov. is described as new to science, and variations in the male terminalia in two populations are noted. Two subspecies (quadridentata quadridentata and quadridentata paupera) 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.
Crane flies, male and female terminalia, new species, Russia, Tipula eleniya
There are 502 species in the subgenus Lunatipula Edwards in the world fauna (
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
We generally follow the terminology of
All measurements of the new species were made on pinned material.
Species distributions are given according to
The original descriptions and measurements by
Abbreviations for institutional and private collections used herein: IEMT Tembotov Institute of Ecology of Mountain Territories of Russian Academy of Sciences, Nalchik, Russia; VPMC private collection of Valentin E. Pilipenko, Moscow, Russia;
Diagnosis (after
Tipula (Lunatipula) caucasica
Georgia • 1 male, 3 females; “ущ. Бани-Хеви бл. Боржоми, Аджаро-Имеретинск. хр. [Gorge Bani-Khevi near Borjomi, Adjaro-Imeretinsk. Ridge]; 26 Jun. 1958; Kurnakov leg.;
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.
Adult male (Fig.
Head.
Light grey, sometimes bluish (Figs
Antenna
(Fig.
Thorax
(Fig.
Wings
(Fig.
Halteres
(Fig.
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
T. (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.
Terminalia
(Figs
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.
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.
Adults were collected at altitudes ranging from sea level (Black Sea coast at 35 m) to 2011 m (in Dagestan).
From end of April to the beginning of July.
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.
Endemic to the Caucasus. It was recently noted for Russia (Dagestan) for the first time (
According to
Holotype: Russia • 1 male; Krasnodarskiy Kray, Sochi env. Psekhako Mt., 43°41'28"N, 40°22'E; alt. ~ 2000 m; 14–18 Jun. 2008; K. Tomkovich leg.;
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 rod-like outgrowth in middle of outer edge.
Adult (Fig.
Head
(Figs
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.
Wings
(Fig.
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
T. (Lunatipula) eleniya sp. nov. A terminal segments of male abdomen, lateral view (dry) B–L male terminalia (after KOH 10% treatment) B hypopygium (without sternite 8), caudal view C hypopygium, lateral view D sternite 8, ventral view E tergite 9, caudal view F tergite IX, 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.
Terminalia
(Figs
T. (Lunatipula) eleniya sp. nov., variation in the structure of the male terminalia A, D, G, J variant I (holotype) B, E, H, K variant II (paratype) C, F, I, L variant III (paratype) A–C inner and outer gonostylus, lateral view D–F inner and outer gonostylus, latero-caudal view G–I tergite 9, caudal view J–L tergite 9, dorsal view. Scale bar: 1 mm.
Female unknown.
Three variants exist of the small rod-like outgrowth of the inner gonostylus: variant I (holotype – Sochi env., Psekhako Mt, alt. 2000 m) (Fig.
The new species is readily separable from all other species of the subgenus by the presence of an outgrowth medially, on the outer edge of the inner gonostylus (Figs
Adults were collected at altitudes ranging from 1200–2000 m.
Adults are active from middle of June through the end of July.
Mixed moderately moist mesophytic plants, shady communities that include common hornbeam (Carpinus betulus), oriental beech (Fāgus orientālis), Nordman fir (Abies nordmanniana), ash vulgaris (Fraxinus excelsior), field maple (Acer campestre), colchis holly (Ilex colchica), etc.
Endemic to the Caucasus: currently known from the West Caucasus.
Tipula (Lunatipula) eleniya sp. nov. is named after the mother of the first author, Elena Nikolaevna Lantsova.
Tipula (Lunatipula) quadridentata paupera
Holotype: Georgia •1 male; “Загор [Загар Сванетия], Груз. ССР” [Zagor Pass Svanetiya, Gruz. SSR. Georgian Soviet Socialist Republic]; alt. 2623 m; 19 Jul. 1957; R. Savenko leg.;
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.
Adult male (Fig.
Head
(Figs
Antennae. Scape light grey with silvery pruinescence, pedicel yellow with subtle brown ring at base. Flagellomeres brown.
Thorax
(Fig.
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.
Abdomen
(Figs
T. (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.
Terminalia
(Figs
Female. Unknown.
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).
This species differs from all other species of the caucasica group by the number and arrangement of setae dorsally on the head (Fig.
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.
Adults were collected from the last third of July to early August.
Data absent.
Endemic to the Caucasus-currently known from the southern slopes of the Greater Caucasus (Georgia).
Tipula (Lunatipula) quadridentata quadridentata
Holotype. Russia • 1 male; ”околицы Ставрополя, байрачн.[ый] лес” [near Stavropol, bairak forest] [small forest in steppe ravines]; 25 May 1954; [S.] Medvedev leg.;
Russia • 9 males, 8 females; “курорт “Горячий Ключ” хр. Котх.[ский] Краснодар.[ский] кр. дуб.[овый] лес” [Goryachiy Klyuch resort Koth[skiy] Ridge, Krasnodar.[sky] District oak. Forest]; 18 May 1956; Gilyarov leg.;
Tergite 9 with four widely spaced teeth and with three rounded notches distally; caudal margin of gonocoxite with two dentate projections. Paired appendages of sternite 8 with wide base bearing thick yellow setae distally covering gap between. Cercus long and straight; hypogynial valve only slightly longer than width at base.
Adult male (Fig.
Head
(Figs
Antennae. Framing around antennal sockets yellow. Scape light brown, pedicel yellow, flagellomeres brown; verticils not longer than corresponding flagellomeres.
Thorax
(Fig.
Wings
(Fig.
Halteres. Stem light brown to yellowish, knob brown.
Legs. Femora light brown with procumbent dark brown bristles. Tarsal claws without noticeable tooth at base.
Abdomen
(Figs
T. (Lunatipula) quadridentata 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.
Terminalia
(Figs
Female. Body length 17.5–19.5 mm, wings 16–18.5 mm. Scape and pedicel brownish yellow. Terminalia (Fig.
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
Adults were collected at altitudes ranging from sea level (54 m in Tiso-samshitovaya rosha) to 800–900 m (in Mount Lysaya, Tuapse district).
Adults were collected from throughout the month of May.
Specimens are found in moderately humid woody deciduous communities.
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).
Tipula (Lunatipula) talyshensis
Holotype: Azerbaijan •1 male, “р-н Лерик, Азербайдж. ССР 26. VI. [1]954 Джафаров” [Lerik region, Azerbaijan. SSR. [Azerbaijanskaya Soviet Socialist Republic] [alt. ~1115 m, 38°46'31"N, 48°24'55"E]; 26 Jun. 1954; Jafarov; / “Tipula talyshensis det. Savchenko sp. n.” [white label] / [Original red label without text] / “Holotypus” [red label];
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.
Adult male (Fig.
Head
(Fig.
Antennae. Scape dark grey with rusty bristles, pedicel yellowish with indistinct brown line in middle, flagellum brown.
Thorax
(Fig.
Wings
(Fig.
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
Terminalia
(Figs
T. (Lunatipula) talyshensis 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.
Female. Adult female body length 18–25 mm, wings 18–18.5 mm. Similar to male. Cercal hypogynial valve (Fig.
Female terminalia of caucasica species group A–C T. (Lunatipula) caucasica D–F T. (Lunatipula) quadridentata G–I T. (Lunatipula) talyshensis A, D, G ovipositor, lateral view B, E, H ovipositor ventral view C, F, I sternite 9 and furca, lateral view. Abbreviations: c–cerci; f–furca; hv–hypogynial valve; s–sternite; t–tergite. Scale bar: 1 mm.
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 paired appendages of sternite 8. The unusually long hypogynial valve of the female distinguishes this species from that of T. caucasica and T. quadridentata, the other known females of the caucasica group.
Adults were collected at altitudes ~ 1115 m.
Adults were collected on 26 June.
No data.
Endemic to the Caucasus – currently only known from Talysh (Azerbaijan).
Comparisons of some components of male terminalia of the caucasica species group A, F T. (Lunatipula) caucasica B, G T. (Lunatipula) eleniya sp. nov. C, I T. (Lunatipula) paupera D, J T. (Lunatipula) quadridentata E, K T. (Lunatipula) talyshensis A–E tergite 9, dorsal view F–K inner and outer gonostylus, lateral view. Arrows point to central notch at apex of tergite 9 (C, D), to the wedge-shaped projection posteriorly on inner gonostylus (F), to the rod-like outgrowth behind anterior beak of inner gonostylus (G), to the anterior (dorsal) margin of gonocoxite (I, J, K). Scale bar: 1 mm.
Males
1 | Inner gonostylus with rod-like outgrowth behind anterior beak (Fig. |
T. eleniya sp. nov. |
– | Without such an outgrowth behind anterior beak on the inner gonostylus (Fig. |
2 |
2 | Wedge-shaped projection posteriorly on inner gonostylus (Fig. |
T. caucasica Riedel, 1920 |
– | Without such a wedge-shaped projection posteriorly on inner gonostylus | 3 |
3 | Anterior (dorsal) margin of gonocoxite with sharp dentate projection in addition to smaller more ventral tooth (Fig. |
4 |
– | Anterior (dorsal) margin of gonocoxite with only traces of a projection at most (Fig. |
T. talyshensis Savchenko, 1964 |
4 | Tergite 9 at apex with four projections, central notch between middle projections small and shallow (dorsal view) (Fig. |
T. quadridentata Savchenko, 1964 |
– | Tergite 9 at apex with only two projections, central notch between quite wide and deep (dorsal view) (Fig. |
T. paupera Savchenko, 1964 |
Females (known for three species of the caucasica group only)
1 | Hypogynial valve long, more than three times longer than width at base (Fig. |
T. talyshensis Savchenko, 1964 |
– | Hypogynial valve short, length approximately equal to width at base (Fig. |
2 |
2 | Cercus upcurved distally (Fig. |
|
– | Cercus straight (Fig. |
T. quadridentata Savchenko, 1964 |
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.
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.
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 (
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 (
The authors express their sincere gratitude to the staff of The Laboratory of Insect Taxonomy (Diptera Department) of the Zoological Institute of Russian Academy of Sciences (St. Petersburg, Russia) – Nikolai Parmonov, Galina Suleimanova, Emilia Narchuk, and Olga Ovchinnikova for the opportunity to work with material from the collection of the institute. The authors would like to express their sincere gratitude to Vladimir Neymorovets (All-Russian Research Institute of Plant Protection, St. Peterburg, Russia) for making the general color photographs of the crane flies. The authors sincerely thank Fenja Brodo (Ottawa, Canada) for valuable advice and assistance in the revision of the English text. We are very grateful for Konstantin Tomkovich (Moscow) for the gift of material collected in the Caucasus Reserve. The authors are sincerely thankful to Nelli Tsepkova (Tembotov Institute of Ecology of Mountain Territories of Russian Academy of Sciences, Nalchik) for providing the geobotanical data in the field and to Nadya Matveyeva (Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg) for her professional comments concerning English names of plant communities. This study has been partly supported by the Russian Science Foundation (project no. 18-04-00961). The research was carried out as part of the Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University No. 12103230063-3.