Corresponding author: Boyan Vagalinski (
Academic editor: Pavel Stoev
The diplopod tribe
Vagalinski B, Golovatch SI (2021) The millipede tribe Brachyiulini in the Caucasus (Diplopoda, Julida, Julidae). ZooKeys 1058: 1–127.
The Caucasus is a vast border region in Eurasia lying between southeastern Europe and western Asia, bordered on the south by Iran, on the southwest by Turkey, on the west by the Black Sea, on the east by the Caspian Sea, and on the north by Russia. Two main parts are distinguished: the Caucasus Major, or the Greater Caucasus, represented by the Main Caucasus Ridge in the north, and the Caucasus Minor, or the Lesser Caucasus, in the south. The Caucasus includes the southern parts of European Russia (Rostov-on-Don Region (with its southernmost parts), Krasnodar and Stavropol provinces, Adygea, Karachay-Cherkess, Kabarda-Balkar, North Ossetia-Alania, Ingush, Chechen, and Dagestan republics), as well as three states in Transcaucasia: Georgia (together with the autonomous republics Abkhazia and Ajara = Ajaria), Armenia, and Azerbaijan. The immediately adjacent parts of Turkey and northwestern Iran are often considered by biologists as belonging to the Caucasus as well (e.g.,
According to the most recent estimates, the millipede fauna of the Caucasus is known to comprise > 160 species, > 50 genera, 14 families, and eight orders. Endemism at the species level is overwhelming, amounting to > 85%, while as many as 25 millipede genera are endemic or subendemic to the Caucasus. All families and orders they belong to, however, are widely distributed at least across the Euro-Mediterranean realm (
Two review papers summarising our knowledge of the millipede fauna of the Caucasus, and both discussing biogeographic issues as well, one by
The Euro-Mediterranean and mostly epigean tribe
All examined material is preserved in 70% ethanol, with particular body parts of some of the type specimens mounted on permanent microscopic slides with Euparal medium. Gonopods of all species represented by more than one male were prepared for scanning electron microscopy (
The diagnosis of the tribe
Identifications and general observations were made under a МБС-10 stereo microscope. Colour pictures of type specimens were prepared by focal stacking of multilayer photographs taken with the aid of a Carl Zeiss Discovery V8 stereo microscope with a Nikon Coolpix S3700 camera mounted on one of the eyepieces. Black-and-white micrographs of various body parts were taken with a ProgRes C7 camera connected to a Zeiss Axio Imager 2 compound microscope. Part of these images were represented as line drawings, after copying with the aid of tracing paper placed on a laptop screen.
All included species are presented with their short synonymy lists (the original combination and the subsequent mentions of any other combination existing in the literature, these concerning only the Caucasian fauna) and known distribution records, except for some more common ones whose distributions are summarised.
Apart from the Caucasus proper (i.e., the two main mountain ranges, the Greater and the Lesser Caucasus), the current study also encompasses the immediately adjacent regions of Ciscaucasia (including its lower parts) and Colchis, which belong to the same biogeographic province. The Hyrcan biogeographic province within both the Republic of Azerbaijan and Iran and the eastern parts of the Pontic Mountains in Turkey, despite often considered as part of the Caucasus sensu lato (see Introduction), are here treated as separate regions and thus not included in the distribution maps. However, unpublished material of the only brachyiulinine to occur in Hyrcania,
All abbreviations of gonopodal and vulval structures are explained in the figure captions. More detailed explanations of the main gonopodal and vulval features in the
A genus of
Differs from its only congener known from the Caucasus,
2 ♂♂, 8 ♀♀ (
From eastern Poland in the west to southwestern Siberia, Russia and western Kazakhstan in the east (
The species is new to the fauna of
Differs from its only congener known from the Caucasus,
Subcosmopolitan (
This ubiquitous, largely anthropochoric species was first listed by
A genus of
Also known from eastern Ukraine, Crimea, and the Kursk, Orel, Voronezh, Belgorod, Rostov-on-Don, Samara, Volgograd, and Saratov regions in central and southern European Russia (
The subspecies
The species is new to the faunas of Georgia and Azerbaijan.
A genus of
Small to medium-sized (L (males) = 15–37 mm) Brachyiulini.
Ommatidia present.
Ozopores right on or tightly behind pro-metazonal suture at least on more anterior body rings.
Epiproct well-developed, from moderately to relatively (not conspicuously) long.
Male hypoproct rounded trapezoidal to semi-circular, ventrally with two distal paramedian setae.
Male mandibular stipites considerably expanded, broadly rounded, without a distinct anterior/anteroventral corner.
Male walking legs ventrally with two well-developed adhesive pads, one each on postfemur and tibia.
Penis short and stout, with very short apical lobes and small terminal lamellae.
In situ considerably protruding from gonopodal sinus, directed caudoventrad.
Promere higher than opisthomere, elongate, bearing a slender distomesal process with a narrow channel connected with a deep distomesal groove on caudal sur face of promere, both designed to envelop the solenomere; median ridge and median groove rather poorly developed; flagellum thin, micro-dentate apically.
Opisthomere rather elongate; basoposterior process weakly pronounced, mostly fused to CBO, ending in a finely branched, papillose, freely protruding, apical outgrowth; anterior process long and flattened, partly enveloping the solenomere; an apicoposterior or lateral process, and a mesomeroidal lobe absent; solenomere fine and slender, with a simple tubular structure; with long spiniform filaments along flagellum channel.
Subcylindrical, mostly symmetrical.
Bursa with a distinct, more or less obtuse, postero-apical margin.
Opening placed apically on bursa.
Operculum subequal in height to bursa.
Receptaculum seminis: central tube narrow; posterior tube long and very narrow, somewhat folded; posterior ampulla small to medium-sized.
Both the distomesal process of the promere and the anterior process of the opisthomere in
A species of
The original record from the valley of Marukha River in the Karachay-Cherkess Republic, Russia almost surely concerns
Some of the examined females had large chunks of tar-like brownish substance behind leg pair 2, tightly stuck to the distal parts of the vulvae and covering the opening. These most likely represent ‘copulatory plugs’ (see, e.g.,
A species of
Emphasising the high-mountainous occurrence of this species. Adjective.
Body rings slightly vaulted. Prozonae with very short and shallow, mostly parallel, longitudinal striae. Metazonae not very deeply, but densely striate, n
Apart from body size and gonopodal structural details,
Two of the four paratype females have chunks of tar-like substance attached to the vulvae, like those observed in
A genus of
A subgenus of the genus
1 ♂ (
The above represents the first record of this species since its original description, suggesting a broader distribution within the western parts of the Greater Caucasus. Nevertheless, considering the high collecting activity that has taken place over the years in this particular area of the Caucasus,
A subgenus of the genus
Apart from the unusually short promere, the gonopods and vulva in
1 ♂, 2 ♀♀ (
20–25 labral setae. Promentum of gnathochilarium rather large, separating both lamellae linguales nearly halfway, each with four setae in a longitudinal row. Collum remarkably long, with five or six well discernible striae at posterolateral corner. Paraprocts very densely setose, without rows of shorter setae at caudal margins. Male leg pair 1 mostly parallel, slightly converging hooks, somewhat more slender compared to the usual brachyiulinine pattern. Male walking legs each with a crested adhesive pad on tibia and postfemur, tibial one very large, strongly protruding, covering the proximal half of tarsus; both pads gradually reduced towards telson, but still visible in last leg pairs; legs 3–8 additionally with a pad rudiment on femur; tarsus and tibia, on midbody legs subequal in length and 3.5–4 × as long as apical claw.
Penis very small, roughly quadrangular, slightly broader than long, with barely discernible apical lobes ending with short, blunt, terminal lamellae directed distad. Gonopods (Fig.
The species is new to the fauna of Armenia. The examined individuals have sparse and short metazonal setae, and very densely setose paraprocts, while the syntypes, according to the original description, completely lack metazonal setae and show only a sparse pilosity on the paraprocts; the type specimens are also somewhat larger. Apart from these differences, all other characters studied completely match
A genus of the tribe
Small to medium-sized Brachyiulini (L (males) = 11–32 mm).
Ommatidia present.
Ozopores right on or tightly behind pro-metazonal suture at least on more anterior body rings.
Epiproct well-developed, markedly long.
Male hypoproct broadly rounded, without setae.
Male mandibular stipites considerably expanded, broadly rounded, anterior/anteroventral corner either indistinct or broadly rounded.
Male walking legs ventrally with two well-developed adhesive pads, one each on postfemur and tibia.
Penis short and stout, with indiscernible or very short apical lobes, as well as small and rounded terminal lamellae.
In situ protruding from gonopodal sinus only with their apical parts, directed completely ventrad.
Promere as high as opisthomere, with a broad base, more or less narrowing distad and bearing a slender, tapering, apicomesal process.
Opisthomere from rather stout to slender; a basoposterior process mostly vestigial, i.e., present as a weakly pronounced vertical ridge running parallel to CBO, ending with a simple, lamellar, apical outgrowth protruding nearly perpendicular to CBO; anterior process fine and tapering; a lateral process absent or represented by a small lobe; a mesomeroidal lobe absent; solenomere complex, consisting of a tapering basomesal process and several other small processes and lobes apically.
Subcylindrical, mostly symmetrical.
Bursa with a distinct, slightly obtuse to subrectangular, postero-apical margin.
Opening placed right on top of bursa.
Operculum shorter than, to subequal to, bursa.
Receptaculum seminis: central tube narrow, digitiform; posterior tube very narrow, more or less folded; posterior ampulla small to medium-sized.
A species of
Iran, Lahijan (type locality); Azerbaijan, Lenkoran [Lankaran] (
A species of
Meaning three-horned in Latin, referring to the apical part of the solenomere which consists of three pointed branches. Adjective.
Body rings slightly vaulted. Prozonae with very short, shallow, longitudinal striae in posterior parts. Metazonae moderately deeply striated, n
A genus of
A subgenus of the genus
Differs from its only congener known from the Caucasus,
Central and southwestern parts of the Balkan Peninsula, northwestern Caucasus.
Apart from the records from the Caucasus region,
Differs from its only congener known from the Caucasus,
Western Caucasus (unspecified type locality) (
This species seems to be a narrow local endemic of the northwestern foothills of the Caucasus Major, showing preferences for limestone terrain.
A genus of
Small to medium-sized Brachyiulini (L (males) = 10–33 mm).
Ommatidia present.
Ozopores set tightly behind pro-metazonal suture at least on more anterior body rings.
Epiproct present, varying in length.
Male hypoproct either broad, dentate at margin and trapezoidal or semi-circular/semi-elliptic.
Male mandibular stipites moderately to considerably expanded, forming a distinct anterior/anteroventral corner.
Male pleurotergum 7 significantly bulging only in some species.
Male walking legs ventrally with two well-developed adhesive pads, one each on postfemur and tibia, postfemoral pads rarely lacking in leg pair 2.
Penis short and stout, usually considerably compressed anterocaudally, with short or indistinct apical lobes.
In situ: promeres slightly to considerably protruding outside gonopodal sinus, directed completely ventrad.
Promere subequal to or higher than opisthomere, mostly roughly leaf-shaped, caudal side sometimes forming small lobes, but bearing no distinct processes; flagellum usually much longer than height of promere.
Opisthomere from stout to relatively slender; basoposterior process usually represented by a differently pronounced vertical lobe, apically forming a variously shaped and freely protruding outgrowth; an anterior process usually present, varying in shape and size; a mesomeroidal lobe always present, mostly rounded, varying in size and prominence; solenomere always conspicuously protruding from CBO, usually slender and tubular.
Subcylindrical.
Bursa usually with a distinct, more or less obtuse, postero-apical margin.
Opening narrow oval or cleft-like, positioned apically.
Operculum usually somewhat longer than or subequal to, rarely shorter than, bursa.
Receptaculum seminis: central tube narrow, mostly straight, forming no distinct central ampulla; posterior tube narrow, usually more or less twisted, ending in an ovoid or spherical posterior ampulla.
Seven species groups can be recognised within the circum-east Mediterranean genus
A species of
A species of
Numerous records from
Also found in Russia, Kalmykia (
Revision of the holotype of
The species is new to the fauna of Armenia, also being among the most common and widespread in the Caucasus region.
The subspecies
Another subspecies,
A species of
A species of
Caucasian endemic with a broad distribution south of the Greater Caucasus watershed; not reported north of the latter.
A species of
The species is new to the fauna of Georgia.
A species of
The species is new to the fauna of Russia.
The subspecies
A species of
Derived from the Latin
Body rings not vaulted. Prozonae with numerous minute grooves in posterior third. Metazonae relatively shallowly striated, n
Despite the considerable morphological similarity between
A species of
Promere rather slender, significantly outreaching the opisthomere, with a narrowly rounded apex; distolaterally micro-papillate. Opisthomere (Fig.
The males in these two samples from Mestia differ from both the original description and drawings, and the currently studied material of
A species of
Meaning armed in Latin, referring to the overall appearance of the male including the opisthomere bearing various spines and denticles, the large tapering mandibular stipites, and the prominent dentate hypoproct. Adjective.
Body rings barely vaulted. Prozonae with short and very shallow longitudinal striae spread across whole surface. Metazonae moderately deeply striated, n
A species of
To emphasise the caudally flattened, serrate, apical outgrowth of the basoposterior process of opisthomere, resembling the rostrum in the sawfishes of the family
Body rings somewhat vaulted (more significantly in caudal part of body). Prozonae with very short and fine longitudinal striae in their hind sections. Metazonae rather shallowly striated, n
This species possesses some peculiar morphological characters, being the only representative of
A species of
Key to the species of the
Character |
|
||
---|---|---|---|
length | 17–19 mm | 14–16 mm | 12–13 mm |
epiproct | with a distinct hyaline tip (usually) turned somewhat dorsad | as in |
straight, ending bluntly without distinct hyaline tip (Fig. |
promere | basal and distal parts of nearly the same width; distolateral lobe weakly to moderately pronounced (Fig. |
distally more or less tapering; distolateral lobe strongly pronounced (Fig. |
of proportions intermediate between |
apical outgrowth of basoposterior process | margin smooth or weakly dentate; lamellar part moderately developed (Fig. |
margin pronouncedly dentate; lamellar part strongly developed (Figs |
margin pronouncedly dentate; lamellar part moderately developed (Figs |
mesomeroidal lobe | without distinct apicolateral part (Fig. |
with distinct apicolateral part (Fig. |
without distinct apicolateral part (Figs |
solenomere | apically with two slender and strongly diverging branches (Fig. |
apically clavate, with two contiguous, irregularly shaped, apical lobes (Figs |
apically with two slender, tightly contiguous branches (Fig. |
Promere (Fig.
Georgia,
The species is new to the fauna of Russia.
The subspecies
A species of
From the Latin
Body rings barely vaulted. Prozonae with very short, fine, mostly parallel, longitudinal striae near pro-metazonal suture. Metazonae moderately deeply striated, n
The legs on body rings 2–6 of the paratype male from
A species of
Honours the friend and long-term collaborator of the first author, Eszter Lazányi, a diplopodologist from the Hungarian Natural History Museum, Budapest, Hungary, whose scientific contributions mostly focus on the tribe
Body rings not vaulted. Prozonae with very short, shallow, scattered, longitudinal striae in posterior parts. Metazonae moderately deeply striated, n
A species of
Length 11–12 mm, height 0.8–0.9 mm. Labral setae: 16. Promentum of gnathochilarium separating both lamellae linguales almost halfway; stipites parabasally each with a row of short, stiff setae, similar to the condition seen in
All remaining external somatic characters as in
The gonopods of the presently examined near-topotypic males show no differences from those seen on the syntype slide.
A species of
Meaning torch-bearer in Latin, after the resemblance of the solenomere to a burning torch, the flame being represented by a fine apical sigmoid process. Noun in apposition.
Body rings gently vaulted. Prozonae with very short, fine, mostly purely longitudinal striae in posterior sections. Metazonae rather deeply striated, n
A species of
Honours Roman Zuev, a myriapodologist from Stavropol, Russia, and the collector of the material on which the description of this new species is based.
Body rings not vaulted. Prozonae with very short and fine, mostly parallel longitudinal striae. Metazonae rather shallowly striated, n
The complete lack of metazonal setae is a rare condition to be observed in the
A species of
Until now, the systematic position of this species has been doubtful.
Unfortunately, the presently examined material has previously been dissected and lacks both male and female copulatory organs. However, the newly described species
Given the general trend for narrow endemism within the
(
A species of
In memory of Eristo Kvavadze (1940–2013), a specialist in earthworms and biological control, the collector of the bulk of the material of this new species.
Body rings very gently vaulted. Prozonae with scattered, short and shallow, mostly parallel longitudinal striae. Metazonae relatively deeply striated, n
As already mentioned under the Remarks section referring to
A species of
Key to the species of the
Character | ||||
---|---|---|---|---|
apical outgrowth of basoposterior process | rather slender, distally tapering, pointing disto-frontad | stout, distally tapering, apex abruptly bent frontad (Figs |
stout, distally tapering, pointing disto-frontad (Fig. |
very fine and slender, pointing disto-frontad (Fig. |
mesomeroidal lobe | weakly pronounced, without apicolateral part | rather weakly pronounced, with an apicolateral part pointing frontad (Figs |
moderately pronounced, with an apicolateral part pointing frontad (Fig. |
strongly pronounced, without apicolateral part (Fig. |
anterior process | well-developed lamellar ridge without free apical part | distally deeply bifurcated, with only a vestigial lamella (Figs |
well-developed lamellar ridge with free apical part (Fig. |
well-developed lamellar ridge without free apical part (Fig. |
solenomere | rather stout, unipartite, apically shortly bifurcate | rather stout, unipartite, apically hollow (Figs |
slender, bipartite (Fig. |
slender, unipartite (Fig. |
After the Greek name of the Black Sea, Pontos Euxeinos, to emphasise the type locality of the new species at the eastern coast of that sea. Adjective.
Body rings somewhat vaulted. Prozonae with scattered, very short and shallow, longitudinal grooves. Metazonae rather deeply striated, n
A species of
Derived from the resemblance of the apical outgrowth of the basoposterior process of the opisthomere to the beak of a hummingbird, family
Body rings weakly vaulted. Prozonae with scattered minute grooves. Metazonae rather shallowly striated, n
A genus of
Honours the Svan people, the indigenous inhabitants of Svanetia, northwestern Georgia, whence the type species of the new genus originates. Gender: masculine.
Medium-sized (L (males) = 23–28 mm) Brachyiulini.
Ommatidia present.
Ozopores set tightly behind pro-metazonal suture at least on more anterior body rings.
Epiproct well-developed, relatively (not conspicuously) long.
Hypoproct rounded trapezoidal to semi-circular, ventrally with two distal paramedian setae.
Male mandibular stipites considerably expanded, forming a distinct ventro-anterior corner.
Male pleurotergum 7 significantly bulging.
Male walking legs ventrally with two well-developed adhesive pads, one each on postfemur and tibia.
Penis short and stout, with very short apical lobes and small terminal lamellae.
In situ: promeres considerably protruding outside gonopodal sinus, directed partly caudad, opisthomeres visible only with their apical parts.
Promere slightly higher than opisthomere, elongate, bearing a long, stout, mesal process on caudal side; flagellum slightly shorter than height of promere.
Opisthomere slender; basoposterior process well-developed, broad, flattened frontocaudally; anterior and apicoposterior processes absent; mesomeroidal lobe shaped as a weakly pronounced ridge; solenomere long and slender, distally bipartite, the frontal part ending in a fine, sharply pointed tip.
Subconical.
Bursa without distinct postero-apical margin.
Opening or median cleft absent – the two valves completely fused to one another.
Operculum subequal in height to bursa.
Receptaculum seminis: central tube short and narrow, ending in a distinct central ampulla; posterior tube long and narrow, ending in a subspherical posterior ampulla.
The new genus resembles
Differs from its only known congener,
Honours Aleksandr B. Ryvkin, a coleopterist and the collector of the specimens used for the description of this new species.
Body rings considerably vaulted. Prozonae with densely set, very short, shallow, parallel, longitudinal striae in their hind sections. Metazonae moderately deeply striated, n
In the absence of a bursal opening and considering the position of the central tube of the vulval receptaculum, which is displaced strongly anteriad, at the border with the operculum, the only way for the male opisthomere to reach the receptaculum would be through widening the gap between the bursa and operculum, with the slender, gradually attenuating solenomere ending with a short and sharply pointed process that seems to serve as a perfect tool for that purpose. This is an obvious and rare example of co-evolution of gonopods and vulvae within
Differs from its only known congener,
Honours Frank Walther, a malacologist and an active collector of various invertebrate groups, including the material used for the present description.
Body rings slightly to moderately vaulted. Male hypoproct (Fig.
All other external somatic characters as in
Despite the distance between the type localities of
1 | Promere nearly half as high as opisthomere |
|
– | Promere subequal to, or somewhat higher than, opisthomere |
|
2 | Opisthomere with a prominent anterior process and a broad and flattened lateral process; without apicoposterior process | ( |
– |
Opisthomere with neither an anterior nor a lateral process, but with a well-developed apicoposterior process protruding perpendicular to |
|
3 | Lateral process of opisthomere finely and densely striated, anterior process strongly bent caudad |
|
– | Lateral process of opisthomere smooth, anterior process only slightly bent caudad |
|
4 | Hypoproct triangular |
|
– | Hypoproct trapezoidal, semi-elliptic or semi-circular |
|
5 |
Opisthomere with a large lateral process nearly level to promere, and a rounded lobe-like basoposterior process; lacking an apicoposterior process (Fig. |
|
– | Opisthomere lacking both lateral and basoposterior processes, but possessing a thumb-like, micro-dentate, apicoposterior process |
|
6 | Promere with a distinct, smaller or larger, (apico/disto)mesal process |
|
– | Promere without prominent processes, only with small to moderate lobes in some species |
|
7 |
Opisthomere with a weakly pronounced basoposterior process ending with a visor-like apical outgrowth protruding nearly perpendicularly to |
|
– | Opisthomere with a more or less well-developed basoposterior process in the shape of a broad lobe with a freely protruding apical part; solenomere simple, slender, apically sharply pointed; mesal process of promere originating subapically or medially |
|
8 | Mesal process stout and long, originating at promere mid-height; promeres positioned mostly anteriorly and only slightly laterally in relation to promeres. Svanetia and Megrelia |
|
– | Mesal process fine and rather short, originating from a distal or apical part of promere; promeres positioned completely laterally in relation to opisthomeres. W Greater Caucasus and N Colchis |
|
9 | Male antennae 1.3–1.4 × as long as head; male pleurotergum 7 ventrally with blunt, spade-like lobes originating entirely from metazona (Fig. |
|
– | Male antennae 1.6–1.8 × as long as head; male pleurotergum 7 ventrally with narrow oar-like lobes with slightly concave apical margins, originating from border zone between pro- and metazona (Fig. |
|
10 | Body length usually > 30 mm, vertical diameter > 2 mm; promere with its distomesal process significantly outreaching the apex (Fig. |
|
– | Body usually < 20 mm in length, vertical diameter < 1.3 mm; promere with its distomesal process being subequal to apex (Fig. |
|
11 | Male hypoproct trapezoidal, with apical margin < half the width at base, tridentate; opisthomere with a distinct basoposterior process, clearly separated from |
|
– | Male hypoproct either rounded or trapezoidal, with apical margin just slightly narrower than base, often forming three or more denticles. Basoposterior process mostly fused to |
|
12 | Body uniformly dark with an orange to dark red mid-dorsal line; promere with mostly parallel side margins, ending with a flat apex |
|
– | Body with a light yellow to ochre dorsum divided by a black axial line; promere significantly tapering all the way to a narrowly rounded apex |
|
13 | Promere considerably higher than opisthomere (cf. Figs |
|
– | Promere subequal in height to opisthomere (cf. Figs |
|
14 | Male hypoproct very broadly trapezoidal, margin with three large denticles (Fig. |
|
– | Hypoproct rounded, edentate; apical outgrowth different in shape |
|
15 | Basoposterior process a well-pronounced lobe ending in a tripartite apical outgrowth; an anterior process absent or vestigial (Fig. |
|
– | Basoposterior process a rather weakly pronounced lobe ending in a leaf-like (from mesal and caudal views) apical outgrowth serrate at margin (Figs |
|
16 | Apical outgrowth of basoposterior process simple, mostly smooth, and more or less tapering distad |
|
– | Apical outgrowth more elaborate, multipartite or bearing various small lobes or denticles |
|
17 | Opisthomere with a very prominent mesomeroidal lobe and with solenomere bent markedly caudad |
|
– | Mesomeroidal lobe either well-developed but moderately pronounced, or positioned at the very base of opisthomere; solenomere directed mostly distad |
|
18 | Apical outgrowth of basoposterior process more or less slender, strongly tapering, bent partly or completely frontad (Figs |
|
– | Apical outgrowth more robust, directed mostly distad; anterior process a short spine or a slender rod. Epiproct with the tip slightly to considerably turned dorsad. Medium-sized species (length usually > 20 mm, height > 1.5 mm) |
|
19 | Apical outgrowth narrow, ending bluntly (Fig. |
|
– | Apical outgrowth broad, roughly diamond-shaped; anterior process long, distally drawn out into a fine rod. Epiproct long |
|
20 | Hypoproct rounded, edentate, rarely with three weakly pronounced and rounded teeth or undulations, with two distal paramedian setae (cf. Figs |
|
– | Hypoproct trapezoidal, margin clearly dentate, usually with a row of more than two submarginal setae (cf. Fig. |
|
21 | Opisthomere with a moderately to strongly pronounced mesomeroidal lobe; anterior process smaller or larger, but always clearly detached from solenomere; solenomere rather thick. Republic of Adygea, Stavropol Territory |
|
– | Mesomeroidal lobe rather weakly pronounced; an anterior process either absent or present as an indistinct ridge, mostly fused to solenomere; solenomere slender. |
|
22 | Apical outgrowth of basoposterior process massive, bent strongly basofrontad, covering the basomesal side of solenomere like a hood (Figs |
|
– | Apical outgrowth flattened, shield-like, oriented distad; mesomeroidal lobe forming an apicolateral and an apicomesal part distally. Metazonal setae absent |
|
23 | Anterior process well-developed, linguiform, visible from most angles; both apicomesal and apicolateral parts of mesomeroidal lobe rather weakly pronounced; solenomere apically only moderately enlarged, bearing a soft sigmoid process directed basad (Fig. |
|
– | Anterior process vestigial, mostly hidden between base of solenomere and distal part of mesomeroidal lobe (Fig. |
|
24 | An anterior process absent. |
|
– | Anterior process well-developed, slender, directed distad |
|
25 | Apical outgrowth of basoposterior process pillow-like, its margin with several small humps and undulations. Epiproct very long and broad |
|
– | Apical outgrowth broad, leaf-shaped, deeply divided into two or three lobes. Epiproct of moderate size |
|
26 | Apical outgrowth mostly symmetrical, with a central part and two side alate parts; anterior process equal to, or slightly higher than, solenomere, the latter bent abruptly caudad (Fig. |
|
– | Apical outgrowth strongly asymmetrical, forming a dentate distal part and a thumb-like lateral part; anterior process significantly outreached by solenomere, the latter nearly straight, directed almost completely distad (Fig. |
|
The present list of 32 species of
Distribution of the genera
The diversity of the
Interestingly, the Caucasus Major’s main ridge does not seem to act as a significant distribution barrier as one might expect. Seven species (excluding the likely anthropochoric
When we talk about the faunogenesis of the Caucasus region, we have to consider the different origin and development of the Greater (
In the light of the aforecited palaeogeographical reconstructions, the genera
The presence of several other species in the Caucasus points to relatively recent zoogeographical connections with the Balkan Peninsula.
The genus
Distribution of the
The current study covers much of the Caucasus, giving a near-to-complete view over the diversity of the
Distribution of the
We are most grateful to Karin Voigtländer and Hans Reip (