Monograph |
Corresponding author: Boyan Vagalinski ( boyanv84@gmail.com ) Academic editor: Pavel Stoev
© 2021 Boyan Vagalinski, Sergei I. Golovatch.
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:
Vagalinski B, Golovatch SI (2021) The millipede tribe Brachyiulini in the Caucasus (Diplopoda, Julida, Julidae). ZooKeys 1058: 1-127. https://doi.org/10.3897/zookeys.1058.68628
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The diplopod tribe Brachyiulini is represented in the fauna of the Caucasus by eight genera and 32 species, of which one genus and 14 species are described as new: Colchiobrachyiulus montanus Vagalinski, sp. nov., Iraniulus tricornis Vagalinski, sp. nov., Omobrachyiulus armatus Vagalinski, sp. nov., O. fasciatus Vagalinski, sp. nov., O. faxifer Vagalinski, sp. nov., O. kvavadzei Vagalinski, sp. nov., O. lazanyiae Vagalinski, sp. nov., O. ponticus Vagalinski, sp. nov., O. pristis Vagalinski, sp. nov., O. trochiloides Vagalinski, sp. nov., O. unugulis Vagalinski, sp. nov., O. zuevi Vagalinski, sp. nov., Svaniulus ryvkini Vagalinski, gen. nov., sp. nov., and S. waltheri Vagalinski, gen. nov., sp. nov. Colchiobrachyiulus Lohmander, 1936, a former subgenus of Megaphyllum, is here elevated to a full genus, and the genus Grusiniulus Lohmander, 1936 is downgraded to a subgenus of the genus Cyphobrachyiulus Verhoeff, 1900, both stat. nov., with their previously described species, Colchiobrachyiulus dioscoriadis (Lignau, 1915) and Cyphobrachyiulus redikorzevi (Lohmander, 1936), respectively, listed as comb. nov. Omobrachyiulus brachyurus (Attems, 1899) is formally established as a junior subjective synonym of O. caucasicus (Karsch, 1881), syn. nov., and Omobrachyiulus implicitus ritsensis (Golovatch, 1981) is formally synonymised with the typical Omobrachyiulus implicitus (Lohmander, 1936), syn. nov. Omobrachyiulus sevangensis (Lohmander, 1932), originally described in the genus Megaphyllum, is here transferred to the former genus, comb. nov. The diagnoses and descriptions of some genera and subgenera are refined and complemented. A key is given to all genera and species of Brachyiulini that occur in the Caucasus, and their distributions are mapped. Several species are recorded as new to the faunas of Armenia, Azerbaijan, Georgia, or Russia. The distribution patterns of the Caucasian Brachyiulini and their biogeographic implications are discussed.
Armenia, Azerbaijan, Georgia, key, new genus, new records, new species, Russia
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 Brachyiulini is the target of the present contribution. With currently > 100 species described (
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 (SEM).
The diagnosis of the tribe Brachyiulini and the diagnoses of the genera and subgenera treated here are updated versions of those found in
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. SEM micrographs were taken with a JEOL JSM-5510 or a JEOL JSM-6510LV scanning electron microscope after mounting on SEM stubs or cover glasses and sputter-coating with gold. Blank maps were generated in ArcGIS and the species distributions were subsequently mapped using Photoshop CC 2019. All image processing was made using Photoshop CC 2019.
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, Iraniulus fagorum (Attems, 1951), is also included in the paper, to compare it with a newly described Caucasian congener, I. tricornis sp. nov.
AE Private collection of Aleksandr Evsyukov, Rostov-on-Don, Russia;
IBER Institute of Biodiversity and Ecosystem Research, Sofia, Bulgaria;
ZMUM Zoological Museum, State University of Moscow, Moscow, Russia.
CAUC broad endemic to the Caucasus region, distributed on both sides of the Greater Caucasus watershed, and both east and west of Mount Kazbek;
CECA central Caucasian endemic (the valley of Kura River and the southern and northern slopes of the Greater and Lesser Caucasus, respectively, east of Likhi (= Surami) Mountain Range);
COLC endemic to Colchis (between the southern and northern foothills of the Greater and Lesser Caucasus, respectively, west of Likhi Mountain Range);
GRCA endemic to the Greater Caucasus (both east and west of Mount Kazbek);
LECA endemic to the Lesser Caucasus;
NWGC endemic to the northwestern parts of the Greater Caucasus (north of the Caucasus Major watershed and west of Mount Kazbek);
SWGC endemic of the southwestern parts of the Greater Caucasus (south of the Caucasus Major watershed and west of Mount Kazbek);
WCIS endemic to western Ciscaucasia (approximately west of the line between the Stavropol Highland and the Pyatigorsk Mountains);
WECA broad endemic to the western parts of the Caucasus region (west of Mount Kazbek, on both sides of the Greater Caucasus watershed including parts of Ciscaucasia, Cochis, or Lesser Caucasus).
AR Autonomous Republic
CBO central body of opisthomere
SIG Sergei I. Golovatch
L body length measured at the ozopore level
H vertical diameter at mid-body
S developmental stadium
nSchub number of striae over a distance equal to the length of the metazona, counted just below the ozopore level in a mid-body ring (after
T telson
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 Brachyiulini can be found in
A genus of Brachyiulini differing from contribal genera by the following combination of characters: promeres positioned completely anteriorly in relation to opisthomeres; promere very short, on average half as long as opisthomere; opisthomere possessing a clearly discernible basoposterior process, the latter sometimes tightly contiguous with CBO, a usually well-pronounced lateral process (very small in some species), which is strongly mesolaterally flattened, and an anterior process; some species with a mesoanterior process.
Brachyiulus jawlowskii Lohmander, 1928: 536–538, fig. 8.
Brachyiulus jawlowskii: Zuev 2014: 351.
Differs from its only congener known from the Caucasus, B. lusitanus Verhoeff, 1898, mainly by the opisthomere possessing a smooth rather than striated lateral process, and a slightly rather than strongly bent anterior process.
2 ♂♂, 8 ♀♀ (ZMUM), Georgia: AR Abkhazia, Pitsunda, Bzyb River Valley, meadow with a few Buxus trees, in litter, 8.IV.1983, SIG leg.
Russia: southern Rostov-on-Don Region and Stavropol (
From eastern Poland in the west to southwestern Siberia, Russia and western Kazakhstan in the east (
The species is new to the fauna of AR Abkhazia. It has already been known from the Caucasus region, reported by
Brachyiulus pusillus, lusitanus (sic!) Verhoeff, 1898: 153–154, fig. 28.
Brachyiulus lusitanus:
Brachyiulus lusitanus calcivagus:
Differs from its only congener known from the Caucasus, B. jawlowskii, mainly by the opisthomere possessing a striated rather than smooth lateral process, and a strongly rather than slightly bent anterior process.
(ZMUM). Georgia: 7 ♂♂, 1 ♀, AR Abkhazia, region of Sukhumi, Nizhnyaya Yashtukha, tobacco plantation, 26.V–19.VI.1981, A. Markosyan leg.; 1 ♂, Tbilisi, canyon of Vere River, 18.X.1980, M. Kokhia leg.; 2 ♂♂, 2 ♀♀, eastern Georgia, Akhmeta District, E of Kasristskali, salty swamp, reed thickets, 7.V.1983, V. Yanushev leg.
Subcosmopolitan (
This ubiquitous, largely anthropochoric species was first listed by
A genus of Brachyiulini differing from contribal genera by the following combination of characters: promeres positioned completely anteriorly in relation to opisthomeres; opisthomere with a mostly vestigial or lobe-like basoposterior process, sometimes bearing one or two apical outgrowths; and a well-developed lateral process; anterior process absent.
Iulus rossicus Timotheew, 1897: 284–291, figs 21–31.
Chromatoiulus (Donbrachyiulus) rossicus:
Chromatoiulus (Donbrachyiulus) rossicus rossicus:
Megaphyllum rossicum:
Byzantorhopalum (Byzantorhopalum) rossicum:
Azerbaijan: 1 ♂ (
Gonopods: Promere (Fig.
Byzantorhopalum rossicum (Timotheew, 1897), ♂ from S of Zərgəran (A–D) and ♀ from near Qurbanəfəndi (E), both in Azerbaijan (
Russia: Environs of Kislovodsk (
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 B. r. strandschanum (Verhoeff, 1937) occurs in southeastern Bulgaria and northeastern Greece (
The species is new to the faunas of Georgia and Azerbaijan.
A genus of Brachyiulini differing from contribal genera by the unique arrangement of the gonopod parts, including the promeres being positioned completely laterally rather than anteriorly in relation to the opisthomeres, and the solenomere being protected by specific grooves/channels in the anterior process of the opisthomere and in the distomesal process of the promere; as well as by the following combination of other characters: opisthomere with a lobe-like basoposterior process being mostly fused to CBO, ending with a freely protruding, branched, papillose, apical outgrowth, a well-developed anterior process, partially enveloping the solenomere, and a very slender, tapering solenomere, without any distinct apical structures.
Both the distomesal process of the promere and the anterior process of the opisthomere in Colchiobrachyiulus seem to be specially designed for providing protection to the solenomere, enveloping the latter from the lateral and caudo-lateral sides, respectively. It is also possible that one or both of these structures take part in copulation, facilitating the penetration of the solenomere into the vulval opening.
Brachyiulus dioscoriadis Lignau, 1915: 382–387, text figs 12–20, tab. IV: figs 4, 5.
Chromatoiulus (Colchiobrachyiulus) dioscoriadis:
Chromatoiulus dioscoriadis:
Megaphyllum dioscoriade
(sic!):
Megaphyllum (Colchiobrachyiulus) dioscoriadis:
Megaphyllum dioscoriadis:
(ZMUM). Georgia: AR Abkhazia: 2 ♂♂, 1 juv., Gulripsh near Sukhumi, 24.X.1953, E. Borutzky leg.; 2 ♂♂, 2 ♀♀, Ochamchira District, Jgerda, Kodorskiy Mountain Range, in litter and under bark, 21.IX.1985, I.A. Ushakov leg.; 2 ♂♂, 3 ♀♀, Pitsunda-Myussera Nature Reserve, Myussera part, 20–130 m, mixed deciduous forest (Castanea, Alnus, etc.), in litter, under bark and stones, 8–10.IV.1983, SIG leg.; 1 ♂, 1 ♀, Pskhu-Gumistinskiy Nature Reserve, Gumistinskiy part, cordon Nizhniy Tsumur, 25.IX.1985, I.A. Ushakov leg.; 3 ♂♂, 1 ♀, 1 juv., Sukhum District, Tsebelda, 300 m a.s.l., Carpinus, Acer and Buxus scrub, in litter, 19.VIII.1986, SIG leg.; 6 ♂♂, 11 ♀♀, 3 juv., Sukhum District, Lake Amtkelis (Azanta) ca. 16 km N of Tsebelda, 550 m, Alnus forest, litter and under bark, 19.VIII.1986, SIG leg.
A species of Colchiobrachiulus differing from its single known congener, C. montanus sp. nov., by the larger size (both sexes > 30 mm in length and 2 mm in height, vs. males < 20 mm in length and 1.3 mm in height, and females < 25 mm in length and 2 mm in height in C. montanus sp. nov.), the longer epiproct, and by certain details of gonopodal structure, more specifically, by the broader apex of the promere and its distomesal process significantly outreaching the apex, vs. the same being subequal to the apex in C. montanus sp. nov.; by the apical outgrowth of the opisthomere’s basoposterior process being relatively broader, with shorter branches at margin; and by the solenomere being bi- rather than unipartite in its distal section.
Colouration
(Fig.
Epiproct
(in Fig.
Gonopods
(Fig.
Colchiobrachyiulus dioscoriadis (Lignau, 1915), gonopods of ♂ from near Lake Amtkelis, Georgia (ZMUM) A right promere, caudal view B right opisthomere, mesal view C apical part of same aspect. Scale bars: 0.2 mm (A, B), 0.1 mm (C). Abbreviations: ap anterior process, ao apical outgrowth, bpp basoposterior process, dmg distomesal groove, dmp distomesal process, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, mg median groove, mr median ridge, s solenomere, sf spiniform filaments, si anteromesal sinus.
Georgia: AR Abkhazia: New Athos Monastery [S of Armianskoe Ushchelie]; at Besla [Basla] River [by Sukhumi]; Kamani Monastery [ca. 3 km N of Sukhumi]; Tsebelda; Lata; by the influx of Chkhalta River in Kodori River; village “Aschary” [Azhara]; on the right bank of Chkhalta River; near the influx of Gvandra River in Sekeni [must be Kodori] River (original localities); Cave Mikhailovskaya near Sukhumi (
COLC-SWGC.
The original record from the valley of Marukha River in the Karachay-Cherkess Republic, Russia almost surely concerns Colchiobrachyiulus montanus sp. nov. All remaining records of C. dioscoriadis comb. nov. come from the Colchis Lowland and the southwestern foothills of the Greater Caucasus, and it is fairly unlikely that the species appears sympatric with its single known congener on the other side of the main Caucasus Major watershed.
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.,
(all from Russia, Karachay-Cherkess Republic, Teberda Biosphere Nature Reserve). Holotype: ♂ (ZMUM) (unbroken), Mount Mussa-Achitara in Dombai, 2700–2800 m, alpine meadow, under stones, 29.VII.1986, SIG leg. Paratypes: 1 ♂ (ZMUM) (head to ring 6 and rest of body, gonopods prepared for SEM), 2 ♀♀ (ZMUM) (one unbroken, the other with head to ring 3 and rest of body, left vulva dissected), same collecting data as for holotype; 1 ♂ (ZMUM) (head to ring 2 (head damaged, gnathochilarium separated), rings 3 to 6, and rest of body (half-broken in the middle, gonopods dissected)), 2 ♀♀ (ZMUM) (one unbroken, the other broken into two pieces), Alibek Canyon near Dombai, 2000–2100 m, sparse Betula stand, litter, under stones, 25.VIII.1986, SIG leg.; 1 ♂ (
A species of Colchiobrachiulus differing from its single known congener, C. dioscoriadis comb. nov., by its smaller size (males < 20 mm in length and 1.3 mm in height, mature females < 25 mm in length and 2 mm in height, vs. > 30 mm in length and 2 mm in height for both sexes in C. dioscoriadis comb. nov.), by a shorter epiproct, and by gonopodal details, more specifically by the more narrow apex of the promere and its distomesal process being subequal to the apex, vs. that same process significantly outreaching the apex in C. dioscoriadis comb. nov.; by the apical outgrowth of the opisthomere’s basoposterior process being relatively more narrow, with longer branches at margin; and by the solenomere being uni- rather than bipartite in its distal section.
Emphasising the high-mountainous occurrence of this species. Adjective.
Measurements: holotype ♂ in S XI, 43+1+T, L = 16 mm, H = 1.2 mm; paratype ♂♂ in S X, 42–45+1+T, H = 1.15–1.2 mm; paratype ♀♀ in S X–XI, 42–45+1+T, L = 18–22 mm, H = 1.5–1.85 mm.
Colouration
(after > 30 years of preservation in alcohol) (Fig.
External structures: Eye patches in adults consisting of 35–40 ommatidia arranged in easily countable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 18–22, respectively. Antennae 1.4–1.5 × as long as head in males and 1.1–1.2 × in females; antennomere 2 > 3 > 4 ~ 5 > 6. Promentum of gnathochilarium separating lamellae linguales in their proximal 2/5 or so, each of the latter with three or four setae in a longitudinal row. Collum mostly smooth, with just two or three shallow grooves at posterolateral corners.
Body rings slightly vaulted. Prozonae with very short and shallow, mostly parallel, longitudinal striae. Metazonae not very deeply, but densely striate, nSchub = 10 or 11; setae apparently mostly abraded, ca. 1/2–2/3 of metazonal length. Ozopores relatively large, placed right in pro-metazonal suture in more anterior body rings, and ca. 1× their diameter in more posterior ones; sutures not sinuous in front of ozopores. Tarsus of mid-body legs ca. 1.2 × as long as tibia and 3 × as long as apical claw.
Telson
(Fig.
Male sexual characters: Mandibular stipites considerably expanded, protruding mostly ventrad, forming no distinct corner. Leg pair 1 compact, rounded, parallel hooks. Walking legs with crested adhesive pads, both tibial and postfemoral ones gradually reduced posteriad, but still visible until caudalmost pairs. Pleurotergum 7 ventrally forming elongated, narrowly rounded lobes (Fig.
Colchiobrachyiulus montanus sp. nov., paratypes (ZMUM) A–C ♂ from near Arkhyz, Russia A left flange of pleurotergum 7, ventro-lateral view B penis, caudal view C right gonopods, oral view D–H ♂ from Mount Mussa-Achitara, Russia D right promere, caudal view E distal part of left opisthomere, mesal-oral view F right opisthomere, caudal view G apical part of left opisthomere, with the flagellum remained in its channel, oral view H branches of apical outgrowth of basoposterior process I ♀ from Mount Mussa-Achitara, Russia left vulva, caudal view. Scale bars: 0.2 mm (A–C, I), 0.1 mm (D, F), 0.05 mm (E, G), 0.01 mm (H). Abbreviations: ap anterior process, ao apical outgrowth, bpp basoposterior process, ct central tube, dmg distomesal groove, dmp distomesal process, f flagellum, fc flagellum channel, hp hyaline protrusions, mg median groove, mr median ridge, p promere, pa posterior ampulla, pt posterior tube, s solenomere, sf spiniform filaments.
Gonopods
(5C, D–H): Promere (Fig.
Female sexual characters: Leg pairs 1 (significantly) and 2 (slightly) shorter and thicker than following legs. Vulva (Fig.
NWGC.
Apart from body size and gonopodal structural details, C. montanus sp. nov. differs from C. dioscoriadis comb. nov. also by the shape of the vulval receptaculum (cf. fig. 182 in
Two of the four paratype females have chunks of tar-like substance attached to the vulvae, like those observed in C. dioscoriadis comb. nov.
A genus of Brachyiulini differing from contribal genera by the following combination of characters: promeres positioned completely anteriorly in relation to opisthomeres; opisthomere lacking a basoposterior process or this represented by only a small remnant, also lacking an anterior process.
A subgenus of the genus Cyphobrachyiulus differing from other subgenera by the following combination of characters: opisthomere possessing an apicoposterior process and lacking a lateral process, or the latter represented by a weakly developed lobe; promere ending in a broad margin, rather than in a distinct apex, protruding in a conspicuous mesal tip; vulval operculum subequal to bursa; male coxae 2 not enlarged.
Julus litoreus Lignau, 1903: 137–138, figs 55–58.
Chromatoiulus (Chromatoiulus) litoreus:
Megaphyllum litoreum:
Cyphobrachyiulus (Diaxylus) litoreus:
1 ♂ (ZMUM), Georgia: Kutaisi District, 8 km E of Orpiri, environs of Cave Tsutskhvati, deciduous forest, rock, litter, 24.X.1981, SIG leg.
Russia: Adler (type locality).
WECA?
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, C. litoreus seems to be a rare species. The species is new to the fauna of Georgia.
A subgenus of the genus Cyphobrachyiulus differing from other subgenera by the following combination of characters: promere significantly shorter than opisthomere; opisthomere possessing an apicoposterior process and lacking a lateral process or lobe; male coxae 2 not enlarged.
Apart from the unusually short promere, the gonopods and vulva in Grusiniulus match the diagnosis of the genus Cyphobrachyiulus. Consequently, the former taxon is downgraded to a subgenus of the latter genus. Examination of the gonopods of the sole species, Cyphobrachyiulus (Grusiniulus) redikorzevi (Lohmander, 1936), shows that the opisthomeral ‘horizontally protruding protecting branch directed mesocaudad’ [translated from
Grusiniulus redikorzevi Lohmander, 1936: 148–152, figs 126–130.
Grusiniulus redikorzevi: Kobakhidze, 1964: 191; 1965: 394;
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.
Cyphobrachyiulus (Grusiniulus) redikorzevi (Lohmander, 1936), comb. nov., gonopods of ♂ from Sevan, Armenia (ZMUM) A right promere, caudal view B right opisthomere, lateral view. Scale bars: 0.1 mm. Abbreviations: apl lamellae of apicoposterior process, app apicoposterior process, f flagellum, lr lateral ridge, mr median ridge, s solenomere, sl lamella of solenomere, sp central process of solenomere.
Georgia: Akhalkalaki and Borjomi (original localities).
CECA-LECA.
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 Brachyiulini differing from contribal genera by the unique, complex structure of the solenomere which consists of a basomesal process terminating the flagellum and seminal channels, a basocaudal process, and a branched apical part; as well as by the following combination of other characters: promeres positioned completely anteriorly in relation to opisthomeres; opisthomere with a lamellar basoposterior process ending with a visor-like apical outgrowth protruding mostly caudad, and an anterior process; a lateral process absent or poorly developed, in the shape of a rather weakly pronounced lobe.
Chromatoiulus (Iraniulus) fagorum Attems, 1951: 421–422, figs 39–41.
Chromatoiulus fagorum:
Megaphyllum loeffleri:
Megaphyllum fagorum:
Iraniulus fagorum:
Azerbaijan: 1 ♀ (
A species of Iraniulus differing from its only known congener, I. tricornis sp. nov., by a larger body (males > 25 mm long and higher than 1.7 mm, females > 30 mm long and ca. 2.5 mm high, vs. males < 20 mm long and ca. 1 mm high, females < 25 mm long and ca. 1.5 mm high in I. tricornis sp. nov.), by having a proportionately longer, more slender opisthomere, and by certain details of solenomere structure: directed completely distad, with a larger basocaudal process, and with an apical part ending in two short rounded branches of equal size on the mesal side, and one slender sigmoid branch on the lateral side, vs. the same turned somewhat caudad in I. tricornis sp. nov., with a proportionately smaller basocaudal process, and with an apical part bearing three sharply pointed branches of equal size.
Gonopods: Promere (Fig.
Iraniulus fagorum (Attems, 1951), gonopods of ♂ from near Astara, Azerbaijan (ZMUM) A left promere, caudal view B right opisthomere (with flagellum in its channel), mesal view C left opisthomere, oral-lateral view D apical part of right opisthomere, mesal view E apical part of left opisthomere, oral-lateral view. Scale bars: 0.1 mm (A–C), 0.05 mm (D, E). Abbreviations: amp apicomesal process, ao apical outgrowth of basoposterior process, ap anterior process, aps apical part of solenomere, bpp basoposterior process, cps basocaudal process of solenomere, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, lp lateral lobe, mg median groove, mps basomesal process of solenomere, mr median ridge, s solenomere, sf spiniform filaments, si anteromesal sinus.
Iran, Lahijan (type locality); Azerbaijan, Lenkoran [Lankaran] (
Iraniulus fagorum can easily be recognised in the field by both its greyish green colouration and its specific, strong odour (Dragan Antić, pers. comm.). The latter is the result of the large proportion of p-cresol (> 90%) in its defensive secretion. This compound is known to be produced by only a few species of Julidae, while it is typical of the order Callipodida (
(all from Georgia, Svanetia): Holotype: ♂ (unbroken) (ZMUM), Khumpreri River (left affluent of Enguri), near Dizi, 1.5–2 km before the influx, ca. 1000 m a.s.l., leaf litter, 14.IX.1986, A. Ryvkin leg. Paratypes: 6 ♂♂ (ZMUM) (one in 4 pieces, gonopods prepared for SEM, leg pair 2 and penis dissected; another in head and 2 pieces, gonopods in situ, the rest unbroken), 1 ♂ (
A species of Iraniulus differing from its only known congener, I. fagorum, by a smaller body: males < 20 mm long and ca. 1 mm high, females < 25 mm long and ca. 1.5 mm high, vs. males > 25 mm long and higher than 1.7 mm, females > 30 mm long and ca. 2.5 mm high in I. fagorum), by having a proportionately shorter and stouter opisthomere, and by details of the solenomere: turned somewhat caudad, with a proportionately smaller basocaudal process, and with an apical part bearing three sharply pointed branches of equal size, vs. the same directed completely distad in I. fagorum, with a larger basocaudal process, and with an apical part ending in two short, rounded branches of equal size on the mesal side and one slender sigmoid branch on the lateral side.
Meaning three-horned in Latin, referring to the apical part of the solenomere which consists of three pointed branches. Adjective.
Measurements: holotype ♂ in S IX, 43+2+T, L = 16 mm, H = 1.05 mm; paratype ♂♂ in S VII–X, 38–43+2–3+T, L = 11–13 mm, H = 0.85–1.1 mm; paratype ♀♀ in S VII–X, 38–45+1–2+T, H = 1.2–1.6 mm, L = 14–21 mm.
Colouration: (after > 30 years in alcohol) (Fig.
External structures: Eye patches in adults consisting of 20–38 ommatidia arranged in hardly traceable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 17–20, respectively. Antennae 1.5–1.6 × as long as head in males and ca. 1.4 × in females; antennomere 2 > 5 ≥ 3 ~ 4 > 6. Gnathochilarium with promentum separating both lamellae linguales in their basal 1/3–2/5, each with three setae in a longitudinal row. Collum mostly smooth, with only 2–3 short striae near posterolateral corners.
Body rings slightly vaulted. Prozonae with very short, shallow, longitudinal striae in posterior parts. Metazonae moderately deeply striated, nSchub = 7 or 8 in males and 9 or 10 in females; metazonal setae from 2/5 (in more anterior rings) to 1/2 (in more posterior rings) of metazonal length. Ozopores placed right on pro-metazonal suture in first several rings, gradually taking a more posterior position to ~ 1.5 their diameter in caudalmost rings; sutures sinuous in front of ozopores in some rings. Tarsus of mid-body legs slightly shorter than tibia and ca. 3 × as long as apical claw.
Telson
(Fig.
Male sexual characters: Mandibular stipites (in Fig.
Iraniulus tricornis sp. nov., ♂ (A, B) and ♀ (C) paratypes from near Dizi, Georgia (ZMUM) A right flange of pleurotergum 7, ventro-lateral view B penis, caudal view C left vulva, caudo-lateral view. Scale bars: 0.2 mm (A, B), 0.1 mm (C). Abbreviations: ct central tube, op operculum, pa posterior ampulla, pt posterior tube.
Gonopods
(Fig.
Iraniulus tricornis sp. nov., gonopods of ♂ paratype from near Dizi (ZMUM) A left gonopods, mesal view B left promere, caudal view C right opisthomere, mesal, slightly oral view D left opisthomere, lateral view E right opisthomere, mesal view. Scale bars: 0.2 mm (A), 0.1 mm (B–E). Abbreviations: amp apicomesal process, ao apical outgrowth of basoposterior process, ap anterior process, aps apical part of solenomere, cps basocaudal process of solenomere, dml distomesal process, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, lp lateral lobe, lr lateral ridge, mg median groove, mps basomesal process of solenomere, mr median ridge, s solenomere, sc seminal channel, sf spiniform filaments, si anteromesal sinus.
Female sexual characters: Leg pairs 1 and 2 somewhat thicker and shorter than following legs. Vulva (Fig.
SWGC.
A genus of Brachyiulini differing from contribal genera by the following combination of characters: promeres positioned slightly to considerably laterally (up to ca. 45° in species of Megaphyllum s. str.) rather than completely anteriorly in relation to opisthomeres; promere visibly broader than opisthomere, with a well-developed median groove partly enveloping the opisthomere; opisthomere possessing a well-developed basoposterior process, clearly detached from CBO in at least ~ its distal 1/3, a differently developed, sometimes vestigial anterior process, and lacking lateral process.
A subgenus of the genus Megaphyllum differing from other subgenera and certain isolated congeners by the following combination of characters: promeres positioned considerably laterally (ca. 45° in most species) rather than completely anteriorly in relation to opisthomeres; promere considerably broader than opisthomere, with a deep median groove tightly enveloping the opisthomere anterolaterally; opisthomere with a small or vestigial anterior process, solenomere composed of two processes, an anterior and a posterior one, the former positioned at the end of the flagellum channel; vulva with an apically or subapically positioned opening, operculum shorter than, or rarely subequal to, bursa; hypoproct trapezoidal or broadly rounded.
Brachyiulus (Chromatoiulus) unilineatus hercules Verhoeff, 1901: 97–98, figs 18–20.
Brachyiulus unilineatus hercules:
Chromatoiulus unilineatus hercules:
Chromatoiulus unilineatus hercules:
Megaphyllum hercules:
Differs from its only congener known from the Caucasus, M. (s. str.) spathulatum (Lohmander, 1936), mainly by the colour pattern: body uniformly dark with an orange to dark red mid-dorsal line, vs. body with a light yellow to ochre dorsum divided by a black axial line in M. spathulatum; and by the promere having mostly parallel side margins, ending with a flat apex, vs. the same significantly tapering all the way to a narrowly rounded apex in M. spathulatum.
Russia: Environs of Novorossiysk (
Central and southwestern parts of the Balkan Peninsula, northwestern Caucasus.
Apart from the records from the Caucasus region,
Chromatoiulus (Chromatoiulus) spathulatus Lohmander, 1936: 104–109, figs 80–84.
Chromatoiulus (Chromatoiulus) spathulatus:
Chromatoiulus spathulus
(sic!):
Megaphyllum spathulatum:
(
Differs from its only congener known from the Caucasus, M. (s. str.) hercules, mainly by the colour pattern: body with a light yellow to ochre dorsum divided by a black axial line, vs. body uniformly dark with an orange to dark red mid-dorsal line in M. hercules; and by the promere significantly tapering all the way to a narrowly rounded apex, vs. the same having mostly parallel side margins, ending with a flat apex in M. hercules.
Western Caucasus (unspecified type locality) (
WCIS?
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 Brachyiulini differing from other contribal genera by the following combination of characters: promeres positioned completely anteriorly in relation to opisthomeres; opisthomere with a mesomeroidal lobe on anterior surface, and with a basoposterior process either completely absent (rarely) or present as a vertical lobe or ridge, with only apical part freely protruding; sometimes with a faint lateral lobe, but never with a well-developed lateral process; solenomere well differentiated, mostly slender.
Seven species groups can be recognised within the circum-east Mediterranean genus Omobrachyiulus: The caucasicus group is the most diverse one and includes both Caucasian and Balkan-south Carpathian species; the hortensis, implicitus, roseni, and sevangensis groups each comprises three or four Caucasian endemics; O. mesorientalis Vagalinski & Golovatch, 2019 from Israel and Lebanon, as well as O. beroni (Strasser, 1973) from southern Bulgaria are morphologically the most isolated/disjunct congeners, each representing a group of its own (
Characterisation. Both gonopod pairs subequal in height. Promere with a well-developed distal groove. Opisthomere rather slender, with a well-pronounced mesomeroidal lobe, a basoposterior process with a moderately to strongly pronounced proximal part, ending with a variously shaped apical outgrowth, an anterior process in the shape of a small spine or rod of various length, a deep anteromesal sinus, a flagellum channel overgrown with short to moderately long spiniform filaments, and a more or less slender, apically often bipartite solenomere. Vulva with operculum subequal (from slightly lower to slightly higher) to bursa.
Included species.
O. adsharicus (Lohmander, 1936)
O. caucasicus (Karsch, 1881), comb. nov. (= Brachyiulus brachyurus Attems, 1899, the type species of the genus)
O. curvocaudatus (Lignau, 1903)
O. divaricatus (Lohmander, 1936)
O. geniculatus (Lohmander, 1928)
O. macrourus (Lohmander, 1928)
O. platyurus (Latzel, 1884) (South Carpathians, Romania; Banat, Romania/Serbia)
O. strasseri Vagalinski & Lazányi, 2018 (Andros, Greece)
O. unugulis Vagalinski, sp. nov.
Chromatoiulus (Omobrachyiulus) adsharicus Lohmander, 1936: 123–126, figs 99–101.
Chromatoiulus (Omobrachyiulus) adsharicus:
Chromatoiulus adsharicus:
Megaphyllum adsharicum:
Omobrachyiulus adsharicus:
Georgia: AR Ajara: 1 ♂ (
A species of Omobrachyiulus differring from congeners mainly by the solenomere being apically bifurcate into two pointed branches of nearly equal size and shape, in combination with the complete absence of an anterior process on the opisthomere.
Georgia, AR Ajara, near mouths of Chorokhi and Acharistskali rivers (type locality).
LECA.
Julus caucasicus Karsch, 1881: 20.
Brachiulus (sic!) brachyurus Attems, 1899: 326–328, figs 72–75, syn. nov.
Chromatoiulus brachyurus:
Chromatoiulus (Omobrachyiulus) brachyurus:
Chromatoiulus (Omobrachyiulus) brachyurus dagestanus
Lohmander, 1936: 121–123, figs 96–98;
Chromatoiulus brachyurus brachyurus:
Chromatoiulus brachyurus dagestanus:
Chromatoiulus (Omobrachyiulus) brachyurus brachyurus:
Megaphyllum brachyurum brachyurum:
Megaphyllum brachyiurum
(sic!):
Megaphyllum caucasicum caucasicum:
Megaphyllum brachyurum:
Omobrachyiulus brachyurus:
Georgia: Holotype: juvenile ♀ (by monotypy) (
A species of Omobrachyiulus differing from congeners mainly by the opisthomere having a thumb-like apical outgrowth of the basoposterior process, bent somewhat anteriad, and by the solenomere ending with a fine, short, sharply pointed, apical process.
Gonopods: Promere (Fig.
Omobrachyiulus caucasicus (Karsch, 1881) comb. nov., ♂ (A–D) and ♀ (E) from S of Zərgəran, Azerbaijan (
Numerous records from Georgia; Russia: Republic of Dagestan (
Also found in Russia, Kalmykia (
Revision of the holotype of Julus caucasicus Karsch, 1881 revealed that the juvenile specimen belongs to the tribe Brachyiulini. This can be inferred from the relatively short epiproct with the apex turned dorsad, in combination with the ozopores set tightly behind the pro-metazonal suture. Of the Caucasian Brachyiulini occurring in the region of Borjomi, only O. brachyurus (Attems, 1899) and O. macrourus Lohmander (Lohmander, 1928) show the aforementioned condition of the epiproct. However, in O. macrourus the latter is both much longer and broader, as the name of this species implies. Furthermore, the dorsolaterally dark trunk and contrasting yellow legs (according to the original description) is exactly the colour pattern seen in O. brachyurus. Thus, there can be little doubt that O. brachyurus is a junior subjective synonym of J. caucasicus.
The species is new to the fauna of Armenia, also being among the most common and widespread in the Caucasus region.
The subspecies O. C. dagestanus was described by
Another subspecies, O. C. thassensis (Mauriès, 1985), was described from the Greek island of Thassos (
Julus curvocaudatus Lignau, 1903: 135–136, figs 55–58.
Chromatoiulus (Omobrachyiulus) curvocaudatus: Lohmander, 1936: 126–129, figs 108–110;
Chromatoiulus curvocaudatus:
Megaphyllum curvocaudatum:
Omobrachyiulus curvocaudatus:
Georgia: 1 ♂ (
A species of Omobrachyiulus differing from congeners mainly by the opisthomere bearing a large, flattened, roughly diamond-shaped, apical outgrowth of the basoposterior process, and a shortly bifurcate solenomere, with the anterior branch being clavate, and the posterior one fine and pointed, bent anteriad.
Russia: Krasnodar Province, Pseashkho Pass (type locality); Sochi (
CAUC.
Chromatoiulus (Omobrachyiulus) divaricatus Lohmander, 1936: 135–140, figs 117–119.
Chromatoiulus (Omobrachyiulus) divaricatus:
Chromatoiulus divaricatus:
Megaphyllum divaricatum:
Omobrachyiulus divaricatus:
Armenia: 3 ♂♂, 2 ♀♀ (ZMUM), “Prov. Zori, Mount Polat”, 30.VIII.1925, A. Schelkovnikov leg. [probably H. Lohmander det.]. Georgia: 1 ♂ (
A species of Omobrachyiulus very similar to O. unugulis sp. nov. by the general shape of the promere and by the opisthomere (Fig.
Omobrachyiulus divaricatus (Lohmander, 1936), gonopods of ♂ from near Nakra, Georgia (ZMUM) A left opisthomere, mesal view B same, oral, slightly lateral view C same, caudal, slightly mesal view. Scale bars: 0.2 mm (A), 0.1 mm (B, C). Abbreviations: ao apical outgrowth of basoposterior process, ap anterior process, fc flagellum channel, ml mesomeroidal lobe, s solenomere, sf spiniform filaments.
Opisthomere
(Fig.
Georgia: shore of Lake Tabatskuri; Turkey/Georgia, Ajara, “Post Arsian” [most likely a locality in the Arsiani Range shared between Turkey and Georgia] (original localities); Ipari; Rveli; Lentekhi District: Ushguli; Koruldashi; at the foot of Mt Shkhara (
Caucasian endemic with a broad distribution south of the Greater Caucasus watershed; not reported north of the latter.
Chromatoiulus geniculatus Lohmander, 1928: 236–238, figs 12–14.
Chromatoiulus (Omobrachyiulus) geniculatus: Lohmander, 1936: 129–131, figs 108–110;
Omobrachyiulus geniculatus:
Georgia: 1 ♂ (
A species of Omobrachyiulus differing from congeners by the following combination of opisthomere characters: basoposterior process in the shape of a strongly pronounced lobe forming a nearly right-angled corner at base, ending with a simple and smooth apical outgrowth with a blunt tip; anterior process shaped as a stout spine; mesomeroidal lobe strongly pronounced; solenomere simple, bent strongly caudad.
Russia, Karachay-Cherkess Republic, by Bolshoy Zelenchuk River (type locality), “Post Akssaut” [probably on Aksaut River] (
GRCA.
The species is new to the fauna of Georgia.
Chromatoiulus macrourus Lohmander, 1928: 542–544, figs 15–17.
Chromatoiulus (Omobrachyiulus) macrourus abchasicus Lohmander, 1936: 131–135, figs 111–114.
Chromatoiulus (Omobrachyiulus) macrourus macrourus:
Chromatoiulus (Omobrachyiulus) macrourus abchasicus:
Chromatoiulus macrourus abchasicus:
Chromatoiulus macrourus macrourus:
Megaphyllum macrourum macrourum:
Megaphyllum macrourum abchasicum:
Omobrachyiulus macrourus:
Georgia: 2 ♂♂ (
A species of Omobrachyiulus closely resembling O. divaricatus and O. unugulis sp. nov. by the general shape of both pro- and opisthomere. Differs from these two species mostly by the bulging pillow-shaped, rather than antero-caudally strongly flattened, apical outgrowth of the basoposterior process, as well as by the very large (both long and broad) epiproct.
Georgia: Tbilisi; Borjomi (original localities); Zeskho near Lentekhi (
CAUC.
The species is new to the fauna of Russia.
The subspecies O. m. abchasicus was described by
(
A species of Omobrachyiulus very similar to O. divaricatus by the general shape of the promere and by the opisthomere having a slender, rod-like, anterior process directed distad, a broad, strongly flattened, multilobed, apical outgrowth of the basoposterior process, and a similarly finely and asymmetrically bifurcate solenomere. Differs from O. divaricatus by the following gonopodal characters: promere with a more broadly rounded apex; apical outgrowth of opisthomeral basoposterior process asymmetrically bilobed, forming a dentate distal part and a thumb-like lateral part, vs. the same being more or less symmetrical, tripartite in O. divaricatus; anterior process of opisthomere significantly lower rather than equal to, or higher than, solenomere; solenomere mostly straight, directed almost completely distad, vs. the same being abruptly bent caudad in O. divarictus; and by the male hypoproct being tri-, rather than penta- or septidentate.
Derived from the Latin unugula meaning paw, after the apical outgrowth of the basoposterior process of the opisthomere, which resembles an animal foot with claws and a large thumb. Adjective.
Measurements: holotype ♂ in S in XI, 50+1+T, L = 29 mm, H = 1.95 mm. Paratype subadult ♂ in S X, 47+2+T.
Colouration
(Fig.
External structures: Eye patches consisting of ca. 45 ommatidia arranged in easily recognisable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 18, respectively. Antennae (Fig.
Body rings not vaulted. Prozonae with numerous minute grooves in posterior third. Metazonae relatively shallowly striated, nSchub = 10 or 11; setae (apparently) mostly abraded, rather short based on the few remaining. Ozopores set tightly behind pro-metazonal suture in more anterior rings, gradually moved further back, to nearly equal to their diameter behind the suture in caudalmost rings, sutures gently sinuous in front of ozopores in some rings. Tarsus of mid-body legs slightly shorter than tibia, and slightly over 3 × as long as apical claw.
Telson
(Fig.
Male sexual characters: Mandibular stipites (in Fig.
Gonopods
(Fig.
Omobrachyiulus unugulis sp. nov., gonopods of ♂ holotype A left gonopods (without the basal parts), mesal view B right promere, caudal view C right opisthomere, oral-lateral view D distal half of the same, oral view E same, caudal view. Scale bars: 0.4 mm (C), 0.3 mm (B), 0.2 mm (A, D, E). Abbreviations: ao apical outgrowth of basoposterior process, ap anterior process, bpp basoposterior process, fc flagellum channel, ml mesomeroidal lobe, p promere, s solenomere, sf spiniform filaments.
Female sexual characters: unknown.
Despite the considerable morphological similarity between Omobrachyiulus divaricatus and O. unugulis sp. nov., they undoubtedly represent two separate species, this being inferred from the few, but significant differences concerning gonopodal characters. Furthermore, the gonopod structure of O. divaricatus is remarkably consistent (in comparison with the condition in, e.g., O. caucasicus), with almost no visible variations between specimens from all over the species’ distribution area.
SWGC.
Characterisation. Promere significantly higher than opisthomere, distolaterally with a micro-squamose lobe or field; with a distinct distal groove. Opisthomere stout, compact, with a weakly to moderately pronounced mesomeroidal lobe, a basoposterior process with a moderately to well-developed proximal part, ending with a variously shaped apical outgrowth, a ridge-like anterior process of various size, being mostly fused to solenomere, a deep anteromesal sinus, a flagellum channel overgrown with very long, erect, spiniform filaments, and a simple, finger- or rod-like solenomere bent more or less caudad. Vulva with operculum shorter than bursa.
Included species.
O. hortensis (Golovatch, 1981)
O. armatus Vagalinski, sp. nov.
O. pristis Vagalinski, sp. nov.
Chromatoiulus hortensis Golovatch, 1981: 110–112, figs 14–26.
Megaphyllum hortense:
Omobrachyiulus hortensis:
Georgia: 2 ♂♂, 2 ♀♀ (ZMUM), NE of Poti, Chaladidi, Alnus, Quercus, Fraxinus forest on swamp, in litter, 13.IV.1983, SIG leg.; 1 ♂, 1 ♀ (ZMUM), AR Abkhazia, Sukhum District, Nizhnyaya Yashtukha, forest nursery, 29.III.1985, А. Markossian; 1 ♂, 2 juv. (ZMUM), same place, tobacco plantation, 16.VI.1980 and 25.VII.1980, А. Markossian.
A species of Omobrachyiulus most similar to O. armatus sp. nov. by the promere significantly outreaching the opisthomere, the latter possessing a massive lobe-like basoposterior process forming two distinct corners, a basal and a distal one, and having a micro-spiculate mesal side, and a unipartite solenomere with a slender rod-like ending. Differs from O. armatus sp. nov. mainly by the clearly tripartite apical outgrowth of the basoposterior process, consisting of a mesal lamellar, a median fan-shaped, and a lateral spiniform part, vs. that same outgrowth being broad, unipartite, collar-shaped and dentate at the margins in the latter species.
Promere rather slender, significantly outreaching the opisthomere, with a narrowly rounded apex; distolaterally micro-papillate. Opisthomere (Fig.
A Omobrachyiulus hortensis (Golovatch, 1981), ♂ from near Poti, Georgia (ZMUM) B, C O. cf. hortensis from Mestia, Georgia (ZMUM) B right promere, caudal view A, C right opisthomere, mesal view. Scale bars: 0.1 mm. Abbreviations: aod median part of basoposterior process’ apical outgrowth, aol lateral part of basoposterior process’ apical outgrowth, aos mesal part of of basoposterior process’ apical outgrowth, ap anterior process, bpp basoposterior process, fc flagellum channel, g (supposed) gonocoxal gland, ml mesomeroidal lobe, s solenomere, sf spiniform filaments, si anteromesal sinus.
Georgia: AR Abkhazia, Sukhum Botanical Garden (type locality).
COLC-SWGC.
Georgia: 3 ♂♂, 2 ♀♀ (ZMUM), Mestia, 1500 m a.s.l., Betula, Rhododendron on moraine, litter and under stones, 5 and 15.IX.1986, SIG leg.; 1 ♂ (
The males in these two samples from Mestia differ from both the original description and drawings, and the currently studied material of O. hortensis by certain structural details of the opisthomere (Fig.
(all from Georgia). Holotype: ♂ (unbroken) (ZMUM), Manglisi, Quercus forest, 12.XI.1984, E. Kvavadze leg. Paratypes: 2 ♂♂ (ZMUM) (one in head to ring 2, ring 3 to ring 6, pleurotergum 7, and rest of body; right antenna, leg pair 2, penis, legs 6, 7, and mid-body leg dissected, opisthomeres and left promere prepared for SEM; the other unbroken with removed hypoproct), 3 ♀♀ (ZMUM), same collecting data as for holotype; 1 ♂ (ZMUM) (partly broken in anterior section), 2 ♀♀ (ZMUM) (one unbroken, the other in 3 pieces), AR Ajara, E of Khulo, decidous forest near spring, rock, litter, 10.X.1981, SIG leg.; 2 ♂♂ (ZMUM) (one in three pieces, with dissected gonopods, the other with a missing posterior body half), 1 ♂ (
A species of Omobrachyiulus most similar to O. hortensis by the promere significantly outreaching the opisthomere, the latter possessing a massive lobe-like basoposterior process forming two distinct corners, a basal and a distal one, and having a micro-spiculate mesal side, and a unipartite solenomere with a slender rod-like ending. Differs from O. hortensis mainly by the broad, unipartite, collar-shaped, apical outgrowth of the basoposterior process dentate at margin, that same outgrowth being clearly tripartite in the latter species, consisting of a mesal lamellar, a median fan-like, and a lateral spiniform part.
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.
Measurements: holotype ♂ in S IX, 45+1+T, L = 17 mm, H = 1.4 mm; paratype ♂♂ in S IX, 44–46+1+T, L = 16–20 mm, H = 1.35–1.55 mm; paratype ♀♀ in S IX, 44–46+1+T, L = 17.5–22 mm, H = 1.7–1.9 mm.
Colouration
(most specimens considerably faded in ethanol) (Fig.
External structures: Eye patches in adults consisting of 30–35 ommatidia, usually arranged in easily countable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 19–23, respectively. Antennae (in Fig.
Body rings barely vaulted. Prozonae with short and very shallow longitudinal striae spread across whole surface. Metazonae moderately deeply striated, nSchub = 7 or 8 in males and 9 or 10 in females; metazonal setae rather sparse, 2/3 (in middle to caudal parts of body)–3/4 (in frontal part of body) of metazonal length. Ozopores set tightly behind pro-metazonal suture in more anterior rings, gradually moved further backwards, to nearly equal to their diameter behind the suture in caudalmost rings; sutures straight to gently sinuous in front of ozopores. Tarsus of a mid-body leg slightly shorter than to equal to tibia, and nearly 3 × as long as apical claw.
Telson
(Fig.
Omobrachyiulus armatus sp. nov., ♂ (A–F) and ♀ (G) paratypes from near Khulo, AR Ajara, Georgia (ZMUM) A hypoproct, ventral view B leg 6 (coxa not shown) C right flange of pleurotergum 7, ventro-lateral view D penis, caudal view E right opisthomere, caudo-mesal view F same, latero-oral view G left vulva, caudal, slightly mesal view. Scale bars: 0.2 mm (C, G), 0.1 mm (A, B, E, F) 0.6 mm (D). Abbreviations: ao apical outgrowth of basoposterior process, ap anterior process, bpp basoposterior process, ct central tube, ml mesomeroidal lobe, pa posterior ampulla, pt posterior tube, s solenomere, sf spiniform filaments.
Male sexual characters: Mandibular stipites (in Fig.
Gonopods
(Figs
Omobrachyiulus armatus sp. nov., gonopods of ♂ paratype from near Manglisi, Georgia (ZMUM) A left gonopods, mesal view B left promere, caudal view C right opisthomere, mesal view D left opisthomere, lateral view E distal part of right opisthomere, mesal view. Scale bars: 0.1 mm. Abbreviations: ao apical outgrowth of basoposterior process, ap anterior process, bpp basoposterior process, dg distal groove, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, lp lateral lobe, mg median groove, ml mesomeroidal lobe, mr median ridge, s solenomere, sc seminal channel, sf spiniform filaments, si anteromesal sinus.
Female sexual characters: Leg pairs 1 and 2 shorter, leg pair 2 also thicker, than following legs. Vulva (Fig.
LECA-COLC.
(all from Georgia). Holotype: ♂ (unbroken) (ZMUM), AR Ajara, 15 km W of Adigeni, Abies, Picea, Fagus, Acer, etc., 1500–1700 m, litter, logs, under stones, 14–15.V.1983, SIG leg. Paratypes: 1 ♂ (ZMUM) (in head and 4 pieces, gonopods, penis, legs 2, 3, 7 and a mid-body leg dissected), 1 ♂ (
A species of Omobrachyiulus most clearly differing from congeners by the caudally flattened, serrate, elongated, apical outgrowth of the basoposterior process of opisthomere, and by the conspicuously large anterior process which is almost as high as the solenomere.
To emphasise the caudally flattened, serrate, apical outgrowth of the basoposterior process of opisthomere, resembling the rostrum in the sawfishes of the family Pristidae when observed from caudal or caudomesal view. Noun in apposition.
Measurements: holotype ♂ in S IX, 41+1+T, L = 15 mm, H = 1.3 mm; paratype ♂♂ in S IX–X, 41–44+1+T, L = 15–17 mm, H = 1.25–1.4 mm; paratype ♀♀ in S IX–X, 38–42+1–2+T, L = 15–19 mm, H = 1.4–1.65 mm.
Colouration
(considerably faded from ethanol) (Fig.
External structures: Eye patches in adults consisting of 30–35 ommatidia, mostly arranged in easily countable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 22–26, respectively. Antennae (in Fig.
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, nSchub = 8 or 9 in males and 11 or 12 in females; metazonal setae 2/5–1/2 of metazonal length. Ozopores set tightly behind pro-metazonal suture in more anterior rings, gradually moved further back, to nearly equal to their diameter behind the suture in caudalmost rings, sutures not sinuous in front of ozopores. Tarsus of mid-body legs 1.1–1.2 as long as tibia and ca. 4 × as long as apical claw.
Telson
(in Fig.
Male sexual characters: Mandibular stipites (in Fig.
Omobrachyiulus pristis sp. nov., ♂ (A–C) and ♀ (D) paratypes from near Adigeni, AR Ajara, Georgia (ZMUM) A right flange of pleurotergum 7, ventro-lateral view B penis, caudal view C distal part of left opisthomere, caudo-mesal view D left vulva, caudo-lateral view. Scale bars: 0.1 mm (A, D), 0.06 mm (C), 0.05 mm (B). Abbreviations: ao apical outgrowth of basoposterior process, ap anterior process, ca central ampulla, ct central tube, pa posterior ampulla, pt posterior tube, s solenomere.
Gonopods
(Figs
Omobrachyiulus pristis sp. nov., gonopods of ♂ paratypes from Adigeni (A) and Danisparauli (B–E), AR Ajara, Georgia (ZMUM) A right gonopods, mesal view B right promere, caudal view C right opisthomere (with flagellum in its channel), caudo-mesal view D right opisthomere, lateral view E distal part of the same aspect. Scale bars: 0.1 mm (A–D), 0.05 mm (E). Abbreviations: al apicolateral part of mesomeroidal lobe, ao apical outgrowth of basoposterior process, ap anterior process, bpp basoposterior process, dg distal groove, dl distolateral lobe, f flagellum (broken near base), fc flagellum channel, g (supposed) gonocoxal gland, lp lateral lobe, mg median groove, ml mesomeroidal lobe, mr median ridge, p promere, s solenomere, sc seminal channel, sf spiniform filaments, si anteromesal sinus.
Female sexual characters: Leg pairs 1 and 2 markedly thicker, first also shorter, than following legs. Vulva (Fig.
This species possesses some peculiar morphological characters, being the only representative of Omobrachyiulus that shows a vulval seminal receptaculum with a differentiated central ampulla, as well as the only one that lacks postfemoral adhesive pads on male leg pair 2.
LECA.
Characterisation. Both gonopod pairs subequal in height. Promere possessing a distinct distal groove. Opisthomere rather slender, with a weakly to moderately pronounced mesomeroidal lobe, a basoposterior process with a weakly developed proximal part in the shape of a lamellar ridge, ending with a broad, collar-like, apical outgrowth showing a soft mesal part, a vestigial (if present) anterior process in the shape of an indistinct ridge, a narrow and not too deep anteromesal sinus, a flagellum channel overgrown with sparse, short to moderately long, spiniform filaments, and a slender, tubular, unipartite or distally branched solenomere directed mostly distad. Vulva with operculum shorter than, to subequal to, bursa.
Included species.
O. implicitus (Lohmander, 1936)
O. fasciatus Vagalinski, sp. nov.
O. lazanyiae Vagalinski, sp. nov.
Chromatoiulus (Omobrachyiulus) implicitus Lohmander, 1936: 140–143, figs 120–122.
Chromatoiulus (Omobrachyiulus) implicitus:
Chromatoiulus implicitus:
Chromatoiulus implicitus ritsensis Golovatch, 1981: 108–110, figs 8–11, syn. nov.
Megaphyllum implicitum implicitum:
Megaphyllum implicitum ritsense:
Megaphyllum implicatum
(sic!):
Omobrachyiulus implicitus:
Georgia: AR Abkhazia: 6 ♂♂, 7 ♀♀, 1 juv. (ZMUM), Pitsunda-Myussera Nature Reserve, Myussera part, 120–130 m a.s.l., mixed deciduous forest (Castanea, Alnus etc.), in litter, under bark and stones, 8–10.IV.1983, SIG leg.; 1 ♂, 3 ♀♀, 1 juv. (ZMUM), Avadhara, 1600–1700 m a.s.l., Abies, Fagus, fern, Rubus, Galium, 18.IX.1985, I.A. Ushakov leg.; 2 ♂♂, 3 ♀♀, 1 juv. (ZMUM), N of Lake Ritsa, Abies, Fagus, in litter, 13.IX.1985, I.A. Ushakov leg. Russia: Republic of Adygea: 1 ♂, 2 ♀♀ (AE), near Maykop, Polkovnitskaya Ravine, 44°20.72'N, 40°11.37'E, pitfall traps, 01–17.X.2011, Yu. Chumachenko leg.; 1 ♂, 1 juv. ♂ (AE), same place, collecting method and collector, 17.X–01.XI.2011; 1 ♂, 1 juv. (ZMUM), confluence of Kisha and Belaya rivers, mixed forest, ca. 200 m a.s.l., 11.VI.2013, R. V. Zuev leg.; 5 ♂♂, 6 ♀♀, 5 juv. (ZMUM), Krasnodar Province, 5.5 km NE of Krasnaya Polyana, lower course of Achipse River, 43°15'25"N, 40°15'25"E, ca. 600 m a.s.l., 19–23.VIII.2014, K. Makarov and A. Matalin leg.
A species of Omobrachyiulus most similar to O. fasciatus sp. nov. and O. lazanyiae sp. nov. by the overall shape of the promere, the weakly developed basoposterior process and mesomeroidal lobe of the opisthomere, and the broad, collar- or scarf-shaped, apical outgrowth of the basoposterior process, the latter having a characteristic, wrinkled, lamellar part partially covering the mesal side of the solenomere. Differs from these two species mainly by the very slender and tripartite solenomere apically forked into two mostly symmetrical, strongly diverging branches, with a minute ear-like lobe mesally at the base of the bifurcation; and by the distally broader promere with a very broad and deep distal groove on the caudal surface, this being a small pit in O. fasciatus sp. nov. and O. lazanyiae sp. nov.; as well as in other gonopod details summarised in a tabular key (Table
Key to the species of the Omobrachyiulus implicitus group based on male gonopodal and external somatic characters.
Character | O. implicitus | O. fasciatus sp. nov. | O. lazanyiae sp. nov. |
---|---|---|---|
length | 17–19 mm | 14–16 mm | 12–13 mm |
epiproct | with a distinct hyaline tip (usually) turned somewhat dorsad | as in O. implicitus (Fig. |
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 O. implicitus and O. fasciatus sp. nov.; distolateral lobe weakly pronounced (Fig. |
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.
Omobrachyiulus implicitus (Lohmander, 1936), gonopods of ♂ from Myussera, AR Abkhazia, Georgia (ZMUM) A right promere, caudal view B apical part of flagellum C right opisthomere, mesal view D distal part of the same aspect E right opisthomere, oral-mesal view F same, oral view. Scale bars: 0.1 mm (A, C, E, F), 0.05 mm (D), 0.01 mm (B). Abbreviations: ao apical outgrowth of basoposterior process, bpp basoposterior process, dg distal groove, dl distolateral lobe, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, mg median groove, ml mesomeroidal lobe, mr median ridge, s solenomere, si anteromesal sinus.
Georgia, AR Abkhazia, Gagry (type locality), near Lake Ritsa (the type locality of O. i. ritsensis)
WECA.
The species is new to the fauna of Russia.
The subspecies O. i. ritsensis was described by
(all from Russia). Holotype: ♂ (in two pieces, gonopods dissected, right antenna missing) (ZMUM), Republic of Adygea, Caucasian Nature Reserve, Lagonaki Plateau, Azishskiy Pass, forest, pitfall traps, 14–26.VIII.2013, Yu. Chumachenko leg. Paratypes: 1 ♂, 1 ♀, 1 subad. ♂, 2 juv. ♀♀ (
A species of Omobrachyiulus most similar to O. implicitus and O. lazanyiae sp. nov. by the overall shape of the promere, the weakly developed basoposterior process and mesomeroidal lobe of the opisthomere, and the broad, collar- or scarf-shaped, apical outgrowth of the basoposterior process which has a characteristic, wrinkled, lamellar part and partially covering the mesal side of the solenomere. Differs from these two species mainly by the apically clavate solenomere, and by the apical outgrowth of the opisthomeral basoposterior process consisting of a relatively narrow cockscomb-like distal part and a strongly developed scarf-shaped mesal part; as well as by other morphological details summarised in Table
From the Latin fascia, meaning scarf or band, after the strongly developed lamellar part of the apical outgrowth of the basoposterior process of the opisthomere, which is wrapped around the basomesal part of the solenomere. Adjective.
Measurements: holotype ♂ in S IX, 45+3+T, L = 15.5 mm, H = 1.05 mm; paratype ♂♂ in S VIII–IX, 44–46+1–3+T, L = 14–16 mm, H = 1–1.1 mm; paratype ♀♀ in S VIII–X, 41–46+2–3+T, L = 11.9–19.9 mm, H = 0.85–1.3 mm.
Colouration
(Fig.
External structures: Eye patches in adults consisting of 30–40 ommatidia, mostly arranged in easily countable vertical rows. Vertigial, supralabral, and labral setae: two, four (one ♂ with five), and 16–18, respectively. Antennae 1.3–1.4 × as long as head in males and 1.2–1.3 × in females; antennomere 2 > 4 = 5 ≥ 3. Gnathochilarium with a promentum separating both lamellae linguales over ca. 2/5 of their length, each latter with three or four setae in a longitudinal row; male stipites parabasally each with a longitudinal row of several short and stiff setae. Collum mostly smooth, with two or three broad shallow grooves near posterolateral corners, and with several densely set, oblique, shallow striae laterally at anterior margin.
Body rings barely vaulted. Prozonae with very short, fine, mostly parallel, longitudinal striae near pro-metazonal suture. Metazonae moderately deeply striated, nSchub = 8 or 9; metazonal setae 1/3 to nearly half of metazonal length. Ozopores set tightly behind pro-metazonal suture in more anterior rings, gradually moved further backwards, to nearly equal to their diameter behind the suture in caudalmost rings, sutures not being sinuous in front of ozopores. Tarsus of mid-body legs equal to, to slightly longer than, tibia, and nearly 3 × as long as apical claw.
Telson
(Fig.
Omobrachyiulus fasciatus sp. nov., ♂ (A–D) and ♀ (E) paratypes from near Mount Fisht, Krasnodar Prov., Russia (ZMUC) A hypoproct, ventral view B left flange of pleurotergum 7, ventro-lateral view C penis, caudal view D distal part of right opisthomere, lateral, somewhat oral view E right vulva, mesal view. Scale bars: 0.2 mm (B, E), 0.1 mm (D), 0.06 mm (A, C). Abbreviations: ao apical outgrowth of basoposterior process, bpp basoposterior process, ct central tube, pa posterior ampulla, pt posterior tube, s solenomere.
Male sexual characters: Mandibular stipites weakly to moderately expanded, forming a blunt rectangular or slightly obtuse anterior corner. Leg pair 1 rather slender, parallel to slightly converging hooks. Adhesive leg pads large and crested, gradually reduced in caudal direction, postfemoral ones disappearing in caudal third of body, tibial ones discernible until caudalmost leg pairs; femora without modifications. Pleurotergum 7 ventrally forming rather elongated, distally somewhat clavate and broadly rounded lobes (Fig.
Gonopods
(Figs
Omobrachyiulus fasciatus sp. nov., gonopods of ♂ paratype from near Mount Fisht, Krasnodar Prov., Russia (
Female sexual characters: Leg pairs 1 and 2 thicker and considerably shorter than following legs. Vulva (Fig.
NWGC.
The legs on body rings 2–6 of the paratype male from
(
A species of Omobrachyiulus most similar to O. implicitus and O. fasciatus sp. nov. by the overall shape of the promere, the weakly developed basoposterior process and mesomeroidal lobe of the opisthomere, and the broad, collar- or scarf-shaped, apical outgrowth of the basoposterior process, having a characteristic, wrinkled, lamellar part, partially covering the mesal side of the solenomere. Differs from those two species mainly by the solenomere which is apically slender (not clavate as in O. fasciatus sp. nov.), divided in two parallel, tightly contiguous (not strongly diverging as in O. implicitus) branches, subapically on mesal side bearing a minute lobe with several denticles, protruding perpendicular to the main axis of the solenomere.
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 Brachyiulini as well.
Measurements: holotype ♂ in S IX, 41+2+T, L = 12 mm, H = 0.95 mm; paratype ♂ in S IX, 42+1+T, L = 12.5 mm, H = 0.9 mm.
Colouration
(strongly faded from the ethanol conservation) (Fig.
External structures: Eye patches in adults consisting of 20–30 ommatidia arranged in unclear vertical rows. Vertigial, supralabral, and labral setae: two, four (one ♂ with five), and 16, respectively. Antennae (in Fig.
Body rings not vaulted. Prozonae with very short, shallow, scattered, longitudinal striae in posterior parts. Metazonae moderately deeply striated, nSchub = 7 or 8; metazonal setae from 1/2 (in frontal and mid-body rings) to 2/3 (in caudalmost rings) of metazonal length. Ozopores set tightly behind pro-metazonal suture in more anterior rings, moved slightly backwards, to ca. half their diameter behind the suture in more posterior rings, sutures being gently sinuous in front of ozopores. Tarsus of mid-body legs ca. 1.3 × as long as tibia and ca. 3 × as long as apical claw.
Telson
(Fig.
Omobrachyiulus lazanyiae sp. nov., ♂ paratype (
Male sexual characters: Mandibular stipites (in Fig.
Gonopods
(Figs
Omobrachyiulus lazanyiae sp. nov., gonopods of ♂ paratype (
Female sexual characters: unknown.
SWGC.
Characterisation. Both gonopod pairs subequal in height. Promere showing a distinct, deep, distal groove. Opisthomere more or less robust and compact, with a strongly pronounced mesomeroidal lobe, a basoposterior process with a moderately pronounced proximal part in the shape of a lamellar ridge or lobe, ending with an apical outgrowth with a shield-like median part and a more or less pronounced lamellar mesal part turned anteriad, a well-differentiated, freely protruding anterior process of various size and shape, a moderately to very deep anteromesal sinus, a flagellum channel overgrown with rather long and erect spiniform filaments, and a stout, tubular, unipartite solenomere, this being broadened apically and turned more or less anteriad. Vulva with operculum subequal in height to bursa.
Included species.
O. roseni (Verhoeff, 1921)
O. faxifer Vagalinski, sp. nov.
O. zuevi Vagalinski, sp. nov.
Brachyiulus roseni Verhoeff, 1921: 44–45, figs 8, 9.
Chromatoiulus roseni:
Omobrachyiulus roseni:
Syntype
slide (gonopods, flange of male body ring 7) (
A species of Omobrachyiulus morphologically very similar to O. zuevi sp. nov. Differing from the latter species by having generally more slender and less strongly embossed gonopods, and by certain details of opisthomere structure, as follows: anterior process well-developed, linguiform, clearly visible from most aspects, vs. the same being vestigial, visible only in oral and oral-mesal views in O. zuevi sp. nov.; mesomeroidal lobe with a rather weakly developed apicolateral part, vs. the same being massive, nearly reaching the level of the solenomere in O. zuevi sp. nov.; solenomere apically only moderately enlarged, with a soft sigmoid process directed basad, vs. the same ending with a very broad, flower-like apex in O. zuevi sp. nov.
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 O. ponticus sp. nov. Tarsus of mid-body legs somewhat shorter than tibia and ca. 3 × as long as apical claw. Leg pair 1 parallel hooks; ventral lobes of ring 7 protruding entirely from metazona.
Omobrachyiulus roseni (Verhoeff, 1921), gonopods of ♂ from Mount Shibaba, Adygea, Russia (
All remaining external somatic characters as in O. zuevi sp. nov.
Russia: Republic of Adygea, 40 km NE of Mount Oshten, at an influx of Belaya River [some 20 km S of Maykop] (type locality).
WCIS.
The gonopods of the presently examined near-topotypic males show no differences from those seen on the syntype slide.
(all from Russia): Holotype: ♂ (unbroken) (ZMUM), Republic of Adygea, Caucasian Biosphere Nature Reserve, Lagonaki Plateau, Azishskiy Pass, subalpine meadow, pitfall traps, 26.VIII–23.IX.2013, Yu. Chumachenko leg. Paratypes: 1 ♂ (
A species of Omobrachyiulus differing from congeners mainly by the apically strongly enlarged and hollow solenomere, postero-apically bearing a fine, pointed, sigmoid process, and by the well-developed lateral lobe of the opisthomere ending with a pointed process which is turned anteriad.
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.
Measurements: holotype ♂ in S IX, 42+1+T, H = 1.1 mm, L = 18.5 mm; paratype ♂♂ in S VIII–IX, 40–43+1+T, L = 15–19 mm, H = 1.05–1.3 mm; paratype ♀♀ in S VIII–IX, 42–45+0–1+T, L = 15–21 mm; H = 1.5–1.8 mm.
Colouration
(Fig.
External structures: Eye patches in adults consisting of 22–35 well-pronounced ommatidia mostly arranged in easily countable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 22–28, respectively. Antennae (in Fig.
Body rings gently vaulted. Prozonae with very short, fine, mostly purely longitudinal striae in posterior sections. Metazonae rather deeply striated, nSchub = 6 or 7 in males and 7 or 8 in females; metazonal setae 1/2–2/3 of metazonal length. Ozopores touching or set barely behind pro-metazonal sutures along entire body; sutures not sinuous in front of ozopores. Tarsus of mid-body legs equal to, to slightly longer than, tibia, and ca. 3 × as long as apical claw.
Telson
(Fig.
Omobrachyiulus faxifer sp. nov., ♂ paratypes from Lagonaki Plateau, Adygea, Russia (ZMUM) (A–D) and Mamayskiy Forest, Stavropol, Russia (ZMUM) (E), and ♀ paratype (F) from Tamanskaya Dacha Forest, Stavropol, Russia (ZMUM) A hypoproct, ventral view B leg 2 C left flange of pleurotergum 7, ventro-lateral view D penis, caudal view E left opisthomere, oral-mesal view F right vulva, caudal view. Scale bars: 0.2 mm (A, C, D), 0.1 mm (B, E, F). Abbreviations: ao apical outgrowth of basoposterior process, bpp basoposterior process, ct central tube fc flagellum channel, g (supposed) gonocoxal gland, lh horn-like process of lateral lobe, ml mesomeroidal lobe, pa posterior ampulla pt posterior tube s solenomere, sc seminal channel, sf spiniform filaments, si anteromesal sinus, sp sigmoid process of solenomere.
Male sexual characters: Mandibular stipites (in Fig.
Gonopods
(Figs
Omobrachyiulus faxifer sp. nov., gonopods of ♂ paratype from Mamayskiy Forest, Stavropol, Russia (ZMUM) A left promere (somewhat damaged in proximal part), caudal view B apical part of flagellum C left opisthomere, mesal view D right opisthomere, lateral view E distal part of left opisthomere, mesal view F distal part of right opisthomere, lateral view. Scale bars: 0.1 mm (A, C, D), 0.05 mm (E, F), 0.01 mm (B). Abbreviations: ao apical outgrowth of basoposterior process, bpp basoposterior process, dg distal groove, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, lh horn-like process of lateral lobe, lp lateral lobe, mg median groove, ml mesomeroidal lobe, mr median ridge, s solenomere, si anteromesal sinus, sp sigmoid process of solenomere.
Female sexual characters: Vulva (Fig.
NWGC-WCIS.
(ZMUM). Holotype: ♂ (in head to ring 6 and rest of body, opisthomeres dissected), Stavropol, Mamayskiy Forest, 31.III.2013, R. Zuev leg. Paratypes: 1 ♂ (in head to ring 2, ring 3 to ring 6, pleurotergum 7 (broken into two pieces), and rest of body; penis and pleurotergum 7 dissected, gonopods prepared for SEM), 1 ♀ (in head to body ring 2 + rest of body, left vulva dissected), same collecting data as for holotype.
A species of Omobrachyiulus being morphologically very similar to O. roseni (Verhoeff, 1921). Differing from the latter species by having generally stouter, more robust gonopods, and by details of the opisthomere, as follows: anterior process minute, vestigial, mostly hidden between base of solenomere and distal part of mesomeroidal lobe, vs. the same being significantly larger, clearly visible from most angles in O. roseni; mesomeroidal lobe protruding in a massive, rounded, apicolateral part, and a much smaller mesoapical part, vs. the apicolateral part being much less pronounced, nearly same size as the mesoapical one in O. roseni; solenomere with a strongly enlarged, flower-like apex, vs. the same being apically only moderately enlarged, with a soft sigmoid process directed basad in O. roseni.
Honours Roman Zuev, a myriapodologist from Stavropol, Russia, and the collector of the material on which the description of this new species is based.
Measurements: holotype ♂ in S VIII or IX, 40+2+T, L = 10.5 mm, H = 0.8 mm; paratype ♂ in S VIII or IX, 42+2+T, L = 12 mm, H = 0.85 mm; paratype ♀ in S IX or X, 44+1+T, L = 14.5 mm, H = 1.15 mm.
Colouration
(Fig.
External structures: Eye patches in adults consisting of ca. 30 weakly pronounced ommatidia, arranged in hardly recognisable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 13 or 14, respectively. Antennae (in Fig.
Body rings not vaulted. Prozonae with very short and fine, mostly parallel longitudinal striae. Metazonae rather shallowly striated, nSchub = 6 or 7; setae absent. Ozopores set tightly behind pro-metazonal suture in more anterior rings, gradually moved further back, to nearly equal to their dm behind the suture in caudalmost rings, sutures slightly (in the males) to strongly (in the female) sinuous in front of ozopores in anterior body section. Tarsus of mid-body legs as long as tibia, and slightly > 3 × as long as apical claw.
Telson
(Fig.
Male sexual characters: Mandibular stipites (in Fig.
Omobrachyiulus zuevi sp. nov., ♂ (A–D) and ♀ (E) paratypes A left flange of pleurotergum 7, ventro-lateral view B penis, caudal view C same, lateral view D right opisthomere, oral-mesal view E left vulva, caudal, somewhat lateral view. Scale bars: 0.2 mm. Abbreviations: ao apical outgrowth of basoposterior process, bpp basoposterior process, ct central tube, ml mesomeroidal lobe, pa posterior ampulla, pt posterior tube, s solenomere, sf spiniform filaments.
Gonopods
(Figs
Omobrachyiulus zuevi sp. nov., gonopods of ♂ paratype A right promere, caudal view B right opisthomere, mesal, somewhat oral view C left opisthomere, lateral, somewhat oral view D distal part of right opisthomere, oral-mesal view E apex of solenomere, same aspect. Scale bars: 0.1 mm (A–C), 0.05 mm (D), 0.01 mm (E). Abbreviations: al apicolateral part of mesomeroidal lobe, am apicomesal part of mesomeroidal lobe, ao apical outgrowth of basoposterior process, ap anterior process bpp basoposterior process, d subapical denticle dg distal groove f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, mg median groove ml mesomeroidal lobe, mr median ridge, s solenomere, sf spiniform filaments, si anteromesal sinus.
Female sexual characters: Leg pairs 1 and 2 slightly thicker than following legs. Vulva (Fig.
The complete lack of metazonal setae is a rare condition to be observed in the Brachyiulini: apart from O. zuevi sp. nov. and its close sibling, O. roseni, this can only be seen in their Aegean congener, O. strasseri Vagalinski & Lazányi, 2018.
WCIS.
Characterisation. Both gonopod pairs subequal in height. Promere lacking a distinct distal groove. Opisthomere rather elongated, with a moderately to strongly pronounced mesomeroidal lobe developed mostly or entirely within the basalmost part of the opisthomere, a basoposterior process with variously pronounced proximal part, ending with a narrow, more or less tapering apical outgrowth turned partly to completely anteriad, a well-developed, ridge-like, at least partly lamellar, anterior process, a variously developed anteromesal sinus, a flagellum channel overgrown with rather short and slanting spiniform filaments, and a more or less slender, uni- or bipartite solenomere directed (almost) completely distad. Vulva with the operculum considerably higher than the bursa.
Included species.
O. sevangensis (Lohmander, 1932), comb. nov.
O. kvavadzei Vagalinski, sp. nov.
O. ponticus Vagalinski, sp. nov.
O. trochiloides Vagalinski, sp. nov.
Chromatoiulus sevangensis Lohmander, 1932: 178–180, fig. 9.
Chromatoiulus (Armeniobrachyiulus) sevangensis:
Megaphyllum cf. sevangense:
“Megaphyllum” sevangense:
Armenia: Elenovka [Sevan], 2 ♂♂ (
A species of Omobrachyiulus most simillar to O. kvavadzei sp. nov., O. ponticus sp. nov., and O. trochiloides sp. nov. in gonopod structure, as well as in the shape of the male telson. Differs from these species mostly by details of the opisthomere: from O. kvavadzei sp. nov. by the significantly lower position of the mesomeroidal lobe devoid of a distinct apicolateral part; from O. ponticus sp. nov. by both the anterior process and the apical outgrowth of the basoposterior process being subequal to, rather than considerably exceeded by, the solenomere; and from O. trochiloides sp. nov. by having a much more obscure, poorly differentiated basoposterior process ending in an unciform, rather than a fine rod-like, apical outgrowth.
Armenia: Sevan (type locality); ?Turkey: vil. Artvin, above Göktaş (
LECA.
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 Omobrachyiulus kvavadzei sp. nov., O. ponticus sp. nov., and O. trochiloides sp. nov. undoubtedly belong to the same species group, together with sevangense. Thus, the latter is treated here as a member of Omobrachyiulus. We refrain from the usage of the subgenus Armeniobrachyiulus for reasons explained in the Remarks section under O. kvavadzei sp. nov.
Given the general trend for narrow endemism within the sevangensis group and the big distance between Sevan, Armenia (the type locality of O. sevangensis) and the province of Artvin, Turkey, the latter record of “Megaphyllum cf. sevangense” (Enghoff, 2006) may also refer to another closely related, yet undescribed species.
(all from Georgia). Holotype: ♂ (unbroken) (ZMUM), AR Ajara, Kintrishi Nature Reserve, mouth of Khekpara River, 2.X.1984, E. Kvavadze leg. Paratypes: 8 ♂♂ (ZMUM) (five unbroken, one in head to ring 3, ring 4 to pleurotergum 7, and rest of body, with dissected penis and gonopods, one in head to pleurotergum 7 and rest of body, gonopods dissected, one broken in 2 pieces, one in 3 pieces), 1 ♂ (
(ZMUM): 1 ♂, 7 ♀♀ (ZMUM) (all fragmented in 2 or more pieces), Georgia, AR Ajara, Kintrishi Nature Reserve, Zeraboseli, 450–600 m a. s. l., 1–3.VI.1981, SIG and J. Martens leg.
A species of Omobrachyiulus most similar to O. sevangensis comb. nov., O. ponticus sp. nov., and O. trochiloides sp. nov. in gonopod structure, as well as in the shape of the male telson. Differs from all these species mainly by the opisthomere having a bifid anterior process and a better developed, higher mesomeroidal lobe ending with a distinct apicolateral part.
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.
Measurements: holotype ♂ in S IX, 40+1+T, L = 15.5 mm, H = 1.05 mm; paratypes in S VIII–IX, 39–42+1–2+T, L = 13–16.5 mm and 19–24 mm, H = 1–1.1 mm and 1.4–1.6 mm, in males and females, respectively.
Colouration
(after > 30 years in alcohol) (Fig.
External structures: Eye patches in adults consisting of 30–35 ommatidia arranged in easily countable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 15, respectively. Antennae (in Fig.
Body rings very gently vaulted. Prozonae with scattered, short and shallow, mostly parallel longitudinal striae. Metazonae relatively deeply striated, nSchub = 6 or 7 (males) and 7 or 8 (females); metazonal setae ca. 3/4 metazonal length in most rings, this ratio becoming 1:1 in caudalmost rings. Ozopores placed right in pro-metazonal suture in first several rings, gradually taking a more posterior position to ~ 1/2 of their diameter behind suture in caudalmost rings; sutures gently sinuous in front of ozopores in only some rings. Tarsus of mid-body legs 1.3–1.5 × as long as tibia and 3.5–5 × as long as apical claw.
Telson
(Fig.
Omobrachyiulus kvavadzei sp. nov., ♂ (A–F) and ♀ (H) paratypes from near mouth of Khekpara River, AR Ajara, Georgia, and ♂ paratype from Zeraboseli, AR Ajara, Georgia (ZMUM) (G) A hypoproct, ventral view B leg 7 C right flange of pleurotergum 7, ventro-lateral view D penis, caudal view E right gonopods, mesal view F right opisthomere, caudo-lateral view G distal part of right opisthomere, mesal view H right vulva, caudo-lateral view. Scale bars: 0.1 mm (A), 0.2 mm (B–H). Abbreviations: al apicolateral part of mesomeroidal lobe, ao apical outgrowth of basoposterior process, ct central tube, dt distal tip of anterior process f flagellum, op operculum, pa posterior ampulla, pt posterior tube s solenomere, sf spiniform filaments, si anteromesal sinus, tp proximal tip of anterior process.
Male sexual characters: Mandibular stipites (in Fig.
Gonopods
(Figs
Omobrachyiulus kvavadzei sp. nov., gonopods of ♂ paratype from near Chakhati, AR Ajara, Georgia (ZMUM) A left promere, caudal view (flagellum broken off near base) B right opisthomere (with flagellum in its channel), mesal view C left opisthomere, latero-caudal view D distal part of the same aspect E distal part of right opisthomere (with flagellum tip protruding out of its channel), mesal view. Scale bars: 0.1 mm (A–C), 0.05 mm (D, E). Abbreviations: al apicolateral part of mesomeroidal lobe, ao apical outgrowth of basoposterior process, ap anterior process, bpp basoposterior process, dml distomesal lobe, dt distal tip of anterior process, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, mg median groove, ml mesomeroidal lobe, mr median ridge, s solenomere, sf spiniform filaments, si anteromesal sinus, tp proximal tip of anterior process.
Female sexual characters: Leg pairs 1 and 2 somewhat thicker than, and ca. as long as, following legs. Vulva (Fig.
As already mentioned under the Remarks section referring to Omobrachyiulus sevangensis comb. nov.,
LECA.
(ZMUM). Holotype: ♂ (in head, collum to ring 3, ring 4 to ring 6, and rest of body; leg pair 1, penis, right leg 3, gonopods, left flange of pleurotergum 7, and a mid-body leg dissected) (ZMUM), Georgia, AR Abkhazia, Pitsunda-Myussera Nature Reserve, Myussera part, 20–130 m, mixed deciduous forest (Castanea, Alnus, etc.), litter, under bark and stones, 8–10.IV.1983, SIG leg. Paratypes: 2 ♀♀ (one unbroken, the other in head to ring 3 and rest of body, right vulva dissected), same collecting data as of the holotype.
A species of Omobrachyiulus most similar to O. sevangensis (Lohmander, 1936) comb. nov., O. kvavadzei sp. nov., and O. trochiloides sp. nov.; sharing with O. sevangensis and O. trochiloides sp. nov. the presence of a lamellar ridge-like anterior process of the opisthomere. Differs from the former species by the presence of a distinct, rounded, apicolateral part of the mesomeroidal lobe of the opisthomere, as well as by the solenomere considerably outreaching both anterior process and apical outgrowth of the basoposterior process, and from the latter species by the much more obscure, poorly differentiated basoposterior process ending in an unciform, rather than a fine rod-like, apical outgrowth. Resembling O. kvavadzei sp. nov. mostly by the promere having a distomesal lobe and by the mesomeroidal lobe of the opisthomere bearing an apicolateral part, but differing in other gonopod characters included in the tabular key to the sevangensis group (Table
Character | O. sevangensis comb. nov. | O. kvavadzei sp. nov. | O. ponticus sp. nov. | O. trochiloides sp. nov. |
---|---|---|---|---|
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.
Measurements: holotype ♂ in S VII, 35+1+T, L = 10 mm, H = 0.9 mm; paratype ♀♀ in S VII–VIII, 34–35+2+T, L = 11.5–12 mm, H = 1.2–1.35 mm.
Colouration
(Fig.
External structures: Eye patches in adults consisting of 20–23 relatively large, clearly convex ommatidia arranged in mostly easily recognisable vertical rows. Vertigial, supralabral, and labral setae: two, four, and 17, respectively. Antennae ca. 1.4 × as long as head in males and 1.1–1.15 × in females; antennomere 2 > 5 > 4 ≥ 3 > 6. Gnathochilarium with a conspicuously large promentum separating lamellae linguales > halfway, the latter with three or four setae in a longitudinal row; stipites in males parabasally each with a longitudinal row of several setae, non-setose in females. Collum smooth, with a single deep groove running along entire lateral margin.
Body rings somewhat vaulted. Prozonae with scattered, very short and shallow, longitudinal grooves. Metazonae rather deeply striated, nSchub = 6 or 7; metazonal setae relatively short, ca. 2/5 of (in more anterior rings), to equal to (in more posterior rings), metazonal length, arranged in very dense whorls. Ozopores placed right in pro-metazonal suture in more anterior rings, and tightly behind suture in more posterior ones; sutures not being sinuous in front of ozopores. Tarsus of mid-body legs 1.3 × as long as tibia and 3.8 × as long as apical claw.
Telson
(Fig.
Male sexual characters: Mandibular stipites rather weakly expanded, forming a rounded ventro-anterior corner. Leg pair 1 (Fig.
Omobrachyiulus ponticus sp. nov., ♂ holotype (A–C) and ♀ paratype (D, E) A leg pair 1, caudal view B left flange of pleurotergum 7, ventro-lateral view C penis, caudal view D right vulva, lateral view (receptaculum seminis seen by transparency, but drawn with solid lines for emphasis; setae omitted) E same, caudal view. Scale bars: 0.2 mm (D), 0.1 mm (A–C, E). Abbreviations: ct central tube, op operculum, pa posterior ampulla, pt posterior tube.
Gonopods
(Fig.
Omobrachyiulus ponticus sp. nov., gonopods of ♂ holotype A left gonopods, mesal view B right promere, caudal view C right opisthomere, mesal view D same, caudo-lateral view E distal part of the same aspect. Scale bars: 0.1 mm. Abbreviations: abs anterior branch of solenomere, al apicolateral part of mesomeroidal lobe, ao apical outgrowth of basoposterior process, ap anterior process, bpp basoposterior process, dml distomesal lobe, f flagellum, fc flagellum channel, ml mesomeroidal lobe, mr median ridge, p promere, pbs posterior branch of solenomere, s solenomere.
Female sexual characters: Leg pairs 1 and 2 somewhat shorter, first also thicker, than following legs. Vulva (Fig.
SWGC.
Holotype: ♂ (in head to ring 6, pleurotergum 7, and rest of body; distal part of left antenna broken off, gonopods and right flange of pleurotergum 7 on permanent slide) (ZMUM), Georgia, Borjomi Nature Reserve, Baniskhevi Valley, 800–900 m, Picea, Fagus and Carpinus forest, in litter, logs and under stones, 12 and 16.V.1983, SIG leg. Paratypes: 1 ♂ (in head to ring 2, ring 3 to ring 6, and rest of body; gonopods prepared for SEM, penis, leg pair 2, leg 11 and mid-body leg on permanent slide) (ZMUM), 4 adult ♀♀ (one in head to ring 3 + rest of body, left vulva dissected) (ZMUM), 2 subadult ♀♀ (ZMUM), same collecting data as of the holotype; 1 ♂ (in head, collum to ring 6, pleurotergum 7 (in pieces), and ring 8 to rest of body; right antenna, leg pair 1, left flange of pleurotergum 7, and a mid-body leg dissected) (
A species of Omobrachyiulus most simillar to O. sevangensis comb. nov., O. kvavadzei sp. nov., and O. ponticus sp. nov. in gonopod structure, as well as in the shape of the male telson. Easily distinguishable from all these species by the massive, well-differentiated basoposterior process of the opisthomere, ending with a very fine and slender apical outgrowth, as well as by the solenomere being unipartite rather than bifurcated apically.
Derived from the resemblance of the apical outgrowth of the basoposterior process of the opisthomere to the beak of a hummingbird, family Trochilidae. Adjective.
Measurements: male holotype in S VIII, 39+2+T, L = 17.5 mm, H = 1.1 mm; male paratypes in S VIII, 37+2+T, L = 12.8 mm, H = 1 mm, and 42+2+T, L = 18 mm, H = 1.2 mm; adult female paratypes in S IX–X, 40–46+0–1+T; L = 21–24 mm, H = 1.4–1.95 mm; subadult female paratypes in S VIII, 36–42+2+T.
Colouration
(after > 35 years in alcohol) (Fig.
External structures: Eye patches in adults consisting of 28–35 ommatidia arranged in easily recognisable vertical rows. Vertigial, supralabral and labral setae: two, four, and 17–20, respectively. Antennae ca. 1.6 × as long as head in males and 1.2 × in females; antennomere 2 > 3 ~ 5 > 4 > 6. Gnathochilarium with a promentum separating lamellae linguales in their proximal 2/5, the latter with three or four setae in a longitudinal row; stipites with a group of short, stiff setae basomedially near the borders with the lamellae. Collum mostly smooth, with four or five very short and shallow striae at posterolateral corners.
Body rings weakly vaulted. Prozonae with scattered minute grooves. Metazonae rather shallowly striated, nSchub = 9 or 10; metazonal setae relatively short (ca. 1/2 of metazonal length) and erect. Ozopores placed right in pro-metazonal suture in first several rings, gradually taking a more posterior position to ~ 1/2 of their diameter in caudalmost rings; sutures not sinuous in front of ozopores. Tarsus of mid-body legs 1.4–1.5 × as long as tibia and 3.6–3.9 × as long as apical claw.
Telson
(Fig.
Male sexual characters: Mandibular stipites moderately expanded, protruding ventro-anteriad, forming a broadly rounded anterior corner. Leg pair 1 (Fig.
Omobrachyiulus trochiloides sp. nov., ♂ (A, B, D, E) and ♀ (F) paratypes from Baniskhevi Valley, Georgia (ZMUM), and ♂ paratype from near Nabeglavi, Georgia (
Gonopods
(Fig.
Omobrachyiulus trochiloides sp. nov., gonopods of ♂ paratypes from Baniskhevi Valley, Georgia (ZMUM) (A–D) and from near Nabeglavi, Georgia (
Female sexual characters: Leg pairs 1 and 2 somewhat thicker than following legs, leg pair 3 also slightly thicker and somewhat longer than neighbouring pairs. Vulva (Fig.
LECA.
Svaniulus ryvkini sp. nov., by present designation.
Svaniulus ryvkini sp. nov.
Svaniulus waltheri sp. nov.
A genus of Brachyiulini differing from contribal genera by the following combination of characters: promeres positioned anteriorly and only slightly laterally in relation to opisthomeres; caudal surface of promere bearing a strongly developed mesal process; opisthomere possessing a broad and flattened basoposterior process and a faint mesomeroidal lobe, and lacking lateral and anterior processes; solenomere simple, more or less slender, ending with a sharply pointed tip; vulva subconical, bursa with completely fused valves (forming neither an opening nor a cleft), a supposed autapomorphy.
Honours the Svan people, the indigenous inhabitants of Svanetia, northwestern Georgia, whence the type species of the new genus originates. Gender: masculine.
The new genus resembles Omobrachyiulus in the opisthomere having a mesomeroidal lobe (although weakly pronounced), a character not seen in other members of Brachyiulini. On the other hand, the long and freely protruding solenomere and the micro-papillary branches (albeit very small) apically on the opisthomeral basoposterior process suggest proximity to Colchiobrachyiulus, while the mesal process caudally on the promere might be homologous to the similarly positioned structure characteristic of the subgenus Rhamphidoiulus Attems, 1905 of the genus Cyphobrachyiulus Verhoeff, 1900. Anyhow, this unique combination of gonopodal characters, coupled with the supposedly autapomorphic condition of the vulva make the establishment of the new genus warranted.
(ZMUM): Holotype: ♂ (in head and 5 pieces, left antenna, penis, leg pairs 1 and 2, penis, gonopods, right flange of pleurotergum 7, mid-body leg, and hypoproct dissected), Georgia, Svanetia, mouth of Nenskra River, Lukhi, N of Khaishi, ca. 800 m a.s.l., leaf litter, 2.IX.1986, A. Ryvkin leg. Paratype: 1 ♀ (in 2 pieces, left vulva dissected), 5 juv. (unbroken), same collecting data as for holotype.
Differs from its only known congener, S. waltheri gen. nov., sp. nov., mainly by being on average slightly smaller, by the relatively shorter male antennae, by the position and shape of the ventral protrusions of male body pleurotergum 7, and by the following details of gonopod structure: promere proportionally broader, abruptly narrowing only apically, with the mesal process significantly exceeding the apex, vs. the same being more slender, gradually narrowing distally, and with the mesal process just slightly exceeding the apex in S. waltheri gen. nov., sp. nov.; opisthomere possessing a distomesal process vs. this being absent from S. waltheri gen. nov., sp. nov., having a broad and flat apical margin of the basoposterior process, vs. the same forming a distinct apical corner in S. waltheri sp. nov., and with the solenomere gradually narrowing all the way to the top, vs. the same being somewhat enlarged apically in S. waltheri gen. nov., sp. nov.
Honours Aleksandr B. Ryvkin, a coleopterist and the collector of the specimens used for the description of this new species.
Measurements: holotype in S IX, 42+1+T, L = 23 mm, H = 1.6 mm; paratype ♀ in S X, 46+1+T, L = 27 mm, H = 2.2 mm.
Colouration
(> 30 years in alcohol) (Fig.
External structures: Eye patches consisting of 27 or 28 and 37 or 38 markedly convex ommatidia, in the male and female, respectively; developmental rows easily countable. Vertigial, supralabral, and labral setae: two, four, and 20, respectively. Antennae ca. 1.35 × as long as head in the male, and 1.25 × in the female; antennomere 2 > 3 ≥ 5 ≥ 4 > 6. Gnathochilarium with a moderately sized promentum separating lamellae linguales by nearly half their length, each latter with four setae in a longitudinal row; male stipites basolaterally each with a faint oblong hump bearing several setae. Collum mostly smooth, with only several sparsely set, short and shallow grooves near posterolateral corners; frontolateral margin gently concave.
Body rings considerably vaulted. Prozonae with densely set, very short, shallow, parallel, longitudinal striae in their hind sections. Metazonae moderately deeply striated, nSchub = 9 or 10 in the male, and 10 or 11 in the female; setae erect to somewhat slanting, from ca. 2/5 (in mid-body and caudal rings) to equal to metazonal length (in anteriormost rings). Ozopores set tightly behind pro-metazonal suture in more anterior rings, and ca. half their diameter behind in more posterior ones; sutures gently to considerably sinuous in front of ozopores in most rings. Tarsus of mid-body legs equal to tibia, and slightly > 3 × as long as apical claw.
Telson
(Fig.
Svaniulus ryvkini gen. nov., sp. nov., ♂ holotype (A–C) and ♀ paratype (D) A hypoproct, ventral view B right flange of pleurotergum 7, ventro-lateral view C penis, caudal view D left vulva, mesal view (setae omitted). Scale bars: 0.1 mm. Abbreviations: ca. central ampulla, ct central tube, pa posterior ampulla, pt posterior tube.
Male sexual characters: Mandibular stipites (Fig.
Gonopods
(Fig.
Svaniulus ryvkini gen. nov., sp. nov., gonopods of ♂ holotype A left gonopods, mesal view B distal part of right promere, lateral view C right promere, caudal view D right opisthomere, meso-caudal view E same, oral view F distal part of the same, meso-caudal view. Scale bars: 0.1 mm (A–E), 0.05 mm (F). Abbreviations: bpp basoposterior process, dmp distomesal process, dr distal ridge, f flagellum, fc flagellum channel, g (supposed) gonocoxal gland, ml mesomeroidal lobe, mp mesal process, mr median ridge, msl mesal lobe of basoposterior process, p promere, pt posterior tube, s solenomere, sf spiniform filaments, sp apical process of solenomere.
Female sexual characters: Leg pairs 1 and 2 somewhat shorter, 1st also slightly thicker, than following legs. Vulva (Fig.
SWGC.
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 Julidae, the family being generally characterised by highly specialised posterior gonopods with species-specific apical parts, in contrast to the more simply and uniformly built vulvae which are often barely distinguishable even between different genera of the same tribe.
(
Differs from its only known congener, S. ryvkini gen. nov., sp. nov., mostly by being on average slightly larger, by the relatively more slender male antennae, by the position and shape of the ventral protrusions of male pleurotergum 7, and by the following details of gonopod structure: promere more slender, distally significantly narrowing, with the mesal process barely exceeding the apex, vs. the same being somewhat stouter, only apically abruptly narrowing, and with the mesal process significantly exceeding the apex in S. ryvkini gen. nov., sp. nov.; opisthomere lacking a distomesal process, vs. this being present in S. ryvkini gen. nov., sp. nov., having the basoposterior process with a distinct apical corner, vs. the same ending in a broad and flat margin in S. ryvkini gen. nov. sp. nov., and with the solenomere being somewhat enlarged apically, vs. the same narrowing all the way to the top in S. ryvkini gen. nov., sp. nov.
Honours Frank Walther, a malacologist and an active collector of various invertebrate groups, including the material used for the present description.
Measurements: holotype in S X, 43+1+T, L = 26 mm, H = 1.8 mm; paratype ♂ in S IX, 43+1+T, L = 27.5 mm, H = 1.75 mm; paratype ♀ in S IX, 44+1+T, L = 37.5 mm, H = 2.65 mm.
Colouration
(apparently somewhat faded) (Fig.
External structures: Eye patches in adults vertically rather elongate, due to partial reduction of the higher developmental (vertical) rows, consisting of 21–35 markedly convex ommatidia. Vertigial, supralabral, and labral setae: two, four, and 16–18, respectively. Antennae (Fig.
Body rings slightly to moderately vaulted. Male hypoproct (Fig.
Svaniulus waltheri gen. nov., sp. nov., ♂ (A–D) and ♀ (E) paratypes A hypoproct, ventral view B right flange of pleurotergum 7, ventro-lateral view C penis, caudal view D right opisthomere, oral view E left vulva, latero-caudal view. Scale bars: 0.1 mm. Abbreviations: ct central tube, g (supposed) gonocoxal gland, ml mesomeroidal lobe, msl mesal lobe of basoposterior process, pt posterior tube, s solenomere, sp apical process of solenomere.
Male sexual characters: Walking legs with adhesive pads still present in caudalmost pairs. Pleurotergum 7 ventrally forming rather narrow and oar-like lobes (Fig.
All other external somatic characters as in S. ryvkini gen. nov., sp. nov.
Gonopods
(Figs
Svaniulus waltheri gen. nov., sp. nov., gonopods of ♂ paratype A right promere, caudal view B apical part of flagellum C right opisthomere, oral view D distal part of the same, latero-oral view E distal part of left opisthomere, caudo-mesal view F apical part of the same, meso-caudal view. Scale bars: 0.1 mm (A, C–E), 0.05 mm (F), 0.01 mm (B). Abbreviations: bpp basoposterior process, dr distal ridge, f flagellum, g (supposed) gonocoxal gland, ml mesomeroidal lobe, mp mesal process, mr median ridge, msl mesal lobe of basoposterior process, s solenomere, sf spiniform filaments, sp apical process of solenomere.
Female sexual characters: First two leg pairs of same condition as in S. ryvkini gen. nov., sp. nov. Vulva (Fig.
Despite the distance between the type localities of S. waltheri gen. nov., sp. nov. and S. ryvkini gen. nov., sp. nov. being only ca. 35 air-km, the distribution areas of the two species are probably divided by the Egrisi Mountain Range. With summit parts above 3000 m a.s.l., it could have split the population of the common ancestor of these two closely related species during colder periods in the geological past.
SWGC.
1 | Promere nearly half as high as opisthomere | 2 |
– | Promere subequal to, or somewhat higher than, opisthomere | 4 |
2 | Opisthomere with a prominent anterior process and a broad and flattened lateral process; without apicoposterior process | (genus Brachyiulus) 3 |
– |
Opisthomere with neither an anterior nor a lateral process, but with a well-developed apicoposterior process protruding perpendicular to CBO (Fig. |
Cyphobrachyiulus (Grusiniulus) redikorzevi (Lohmander, 1936) |
3 | Lateral process of opisthomere finely and densely striated, anterior process strongly bent caudad | Brachyiulus lusitanus Verhoeff, 1898 |
– | Lateral process of opisthomere smooth, anterior process only slightly bent caudad | Brachyiulus jawlowskii Lohmander, 1928 |
4 | Hypoproct triangular | 5 |
– | Hypoproct trapezoidal, semi-elliptic or semi-circular | 6 |
5 |
Opisthomere with a large lateral process nearly level to promere, and a rounded lobe-like basoposterior process; lacking an apicoposterior process (Fig. |
Byzantorhopalum (s. str.) rossicum (Timotheew, 1897) |
– | Opisthomere lacking both lateral and basoposterior processes, but possessing a thumb-like, micro-dentate, apicoposterior process | Cyphobrachyiulus (Diaxylus) litoreus (Lignau, 1903) |
6 | Promere with a distinct, smaller or larger, (apico/disto)mesal process | 7 |
– | Promere without prominent processes, only with small to moderate lobes in some species | 11 |
7 |
Opisthomere with a weakly pronounced basoposterior process ending with a visor-like apical outgrowth protruding nearly perpendicularly to CBO; solenomere complex, multibranched (Fig. |
Iraniulus tricornis sp. nov. |
– | 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 |
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 | (genus Svaniulus) 9 |
– | 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 | (genus Colchiobrachyiulus) 10 |
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. |
Svaniulus ryvkini gen. nov., sp. nov. |
– | 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. |
Svaniulus waltheri gen. nov., sp. nov. |
10 | Body length usually > 30 mm, vertical diameter > 2 mm; promere with its distomesal process significantly outreaching the apex (Fig. |
Colchiobrachyiulus dioscoriadis (Lignau, 1915) |
– | Body usually < 20 mm in length, vertical diameter < 1.3 mm; promere with its distomesal process being subequal to apex (Fig. |
Colchiobrachyiulus montanus sp. nov. |
11 | Male hypoproct trapezoidal, with apical margin < half the width at base, tridentate; opisthomere with a distinct basoposterior process, clearly separated from CBO in its distal half or so, a mesomeroidal lobe absent | (genus Megaphyllum , subgenus Megaphyllum) 12 |
– | 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 CBO, with only a freely protruding apical outgrowth; anterior surface of opisthomere with a more or less well-developed mesomeroidal lobe | (genus Omobrachyiulus) 13 |
12 | Body uniformly dark with an orange to dark red mid-dorsal line; promere with mostly parallel side margins, ending with a flat apex | Megaphyllum (s. str.) hercules (Verhoeff, 1901) |
– | 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 | Megaphyllum (s. str.) spathulatum (Lohmander, 1936) |
13 | Promere considerably higher than opisthomere (cf. Figs |
14 |
– | Promere subequal in height to opisthomere (cf. Figs |
16 |
14 | Male hypoproct very broadly trapezoidal, margin with three large denticles (Fig. |
Omobrachyiulus armatus sp. nov. |
– | Hypoproct rounded, edentate; apical outgrowth different in shape | 15 |
15 | Basoposterior process a well-pronounced lobe ending in a tripartite apical outgrowth; an anterior process absent or vestigial (Fig. |
Omobrachyiulus hortensis (Golovatch, 1981) |
– | Basoposterior process a rather weakly pronounced lobe ending in a leaf-like (from mesal and caudal views) apical outgrowth serrate at margin (Figs |
Omobrachyiulus pristis sp. nov. |
16 | Apical outgrowth of basoposterior process simple, mostly smooth, and more or less tapering distad | 17 |
– | Apical outgrowth more elaborate, multipartite or bearing various small lobes or denticles | 20 |
17 | Opisthomere with a very prominent mesomeroidal lobe and with solenomere bent markedly caudad | Omobrachyiulus geniculatus (Lohmander, 1928) |
– | Mesomeroidal lobe either well-developed but moderately pronounced, or positioned at the very base of opisthomere; solenomere directed mostly distad | 18 |
18 | Apical outgrowth of basoposterior process more or less slender, strongly tapering, bent partly or completely frontad (Figs |
the Omobrachyiulus sevangensis group (Table |
– | 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 |
19 | Apical outgrowth narrow, ending bluntly (Fig. |
Omobrachyiulus caucasicus (Karsch, 1881) |
– | Apical outgrowth broad, roughly diamond-shaped; anterior process long, distally drawn out into a fine rod. Epiproct long | Omobrachyiulus curvocaudatus (Lignau, 1903) |
20 | Hypoproct rounded, edentate, rarely with three weakly pronounced and rounded teeth or undulations, with two distal paramedian setae (cf. Figs |
21 |
– | Hypoproct trapezoidal, margin clearly dentate, usually with a row of more than two submarginal setae (cf. Fig. |
24 |
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 | 22 |
– | Mesomeroidal lobe rather weakly pronounced; an anterior process either absent or present as an indistinct ridge, mostly fused to solenomere; solenomere slender. AR Abkhazia, Krasnodar Province, Republic of Adygea |
the Omobrachyiulus implicitus group (Table |
22 | Apical outgrowth of basoposterior process massive, bent strongly basofrontad, covering the basomesal side of solenomere like a hood (Figs |
Omobrachyiulus faxifer sp. nov. |
– | Apical outgrowth flattened, shield-like, oriented distad; mesomeroidal lobe forming an apicolateral and an apicomesal part distally. Metazonal setae absent | 23 |
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. |
Omobrachyiulus roseni (Verhoeff, 1921) |
– | Anterior process vestigial, mostly hidden between base of solenomere and distal part of mesomeroidal lobe (Fig. |
Omobrachyiulus zuevi sp. nov. |
24 | An anterior process absent. AR Ajara | Omobrachyiulus adsharicus (Lohmander, 1936) |
– | Anterior process well-developed, slender, directed distad | 25 |
25 | Apical outgrowth of basoposterior process pillow-like, its margin with several small humps and undulations. Epiproct very long and broad | Omobrachyiulus macrourus (Lohmander, 1928) |
– | Apical outgrowth broad, leaf-shaped, deeply divided into two or three lobes. Epiproct of moderate size | 26 |
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. |
Omobrachyiulus divaricatus (Lohmander, 1936) |
– | 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. |
Omobrachyiulus unugulis sp. nov. |
The present list of 32 species of Brachyiulini (more than a quarter of all described species of the tribe (see
Distribution of the genera Brachyiulus (purple), Cyphobrachyiulus (green), Colchiobrachyiulus (black), Iraniulus (blue) Megaphyllum (brown), and Svaniulus (red) in the Caucasus: Brachyiulus jawlowskii (open square), B. lusitanus (ring), Cyphobrachyiulus (Diaxylus) litoreus (drop), C. (Grusiniulus) redikorzevi comb. nov. (star), Colchiobrachyiulus dioscoriadis comb. nov. (cross), C. montanus sp. nov. (diamond), I. tricornis sp. nov. (triangle pointing down), Megaphyllum hercules (pentagon), M. spathulatum (filled circle), S. ryvkini gen. nov., sp. nov. (filled square), and S. waltheri gen. nov., sp. nov. (triangle pointing up); solid line representing the approximate south border of the distribution range of Byzantorhopalum rossicum rossicum.
The diversity of the Brachyiulini within the study region is markedly concentrated in the Colchidan biogeographical province (as understood by, e.g.,
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 Brachyiulus jawlowskii and Megaphyllum hercules, with the latter’s presence in the Caucasus remaining dubious), viz. Byzantorhopalum rossicum, Cyphobrachyiulus litoreus, Omobrachyiulus caucasicus, O. curvcaudatus, O. geniculatus, O. implicitus, and O. macrourus occur on both sides of the mountain range, while most of the remaining species are narrow local endemics of various parts of the Caucasus, and are thus restricted by other geographical or ecological factors. The same holds true for the Caucasian millipedes in general. However, the two highly similar species of Colchiobrachyiulus, C. dioscoriadis and C. montanus sp. nov., seem to lead their existence from splitting the population of their ancestral species into two by the summit parts west of Elbrus, these probably becoming inaccessible to this particular brachyiulinine lineage at some point during the global cooling in the late Pliocene.
When we talk about the faunogenesis of the Caucasus region, we have to consider the different origin and development of the Greater (GC) and the Lesser Caucasus (LC). GC existed as an archipelago and later as an isolated landmass surrounded by the Eastern Paratethys at least throughout most of the Miocene, becoming connected to LC only by the end of the epoch, and with the East European Platform not earlier than the late Pliocene. In contrast, LC is an older landmass that has remained connected with Anatolia and Hyrcania through much of the Cenozoic (
In the light of the aforecited palaeogeographical reconstructions, the genera Colchiobrachyiulus and Svaniulus gen. nov., each known from two species from GC, likely owe their present endemic status to relatively recent migrations followed by declines and extinctions in their ancestral distribution areas. Such extinction events also might have led to the present condition of the highly distinct subgenus Grusiniulus of the mostly Aegean-Anatolian genus Cyphobrachyiulus, which is represented nowadays by a single species, apparently endemic to LC and Transcaucasia. Another Anatolian element in the Caucasian fauna is the Western Caucasian endemic Cyphobrachyiulus (Diaxylus) litoreus, the only representative of the subgenus Diaxylus in the study region.
The presence of several other species in the Caucasus points to relatively recent zoogeographical connections with the Balkan Peninsula. Byzantorhopalum rossicum is represented by two subspecies, the nominate one being widespread in the northern Caucasus, eastern Ukraine and Crimea, and B. r. strandschanum occurring in the East Balkans (
The genus Omobrachyiulus is one of the large genera within the Brachyiulini. With its 20 species known from the Caucasus alone, it is by far the most speciose brachyiulinine genus in the region. Outside the study region, Omobrachyiulus is represented only by four species and one subspecies scattered from the South Carpathians to the Middle East (
Distribution of the Omobrachyiulus caucasicus (red and green) and O. sevangensis (black) species groups in the Caucasus (excluding O. caucasicus comb. nov. known from numerous records in the region, and with a distribution range exceeding the map coverage): O. adsharicus (ring), O. curvocaudatus (diamond), O. geniculatus (pentagon), O. divaricatus (dotted line indicating the approximate distribution range), O. macrourus (open square), O. unugulis sp. nov. (filled circle), O. sevangensis comb. nov. (filled square), O. cf. sevangensis comb. nov. (filled square with white dot), O. kvavadzei sp. nov. (cross), O. ponticus sp. nov. (star), and O. trochiloides sp. nov. (triangle).
The current study covers much of the Caucasus, giving a near-to-complete view over the diversity of the Brachyiulini in the region. However, single records like those of Svaniulus waltheri gen. nov., sp. nov., S. ryvkini gen. nov., sp. nov., Omobrachyiulus unugulis sp. nov., and O. zuevi sp. nov. suggest that certain narrow local endemics could still await discovery even in areas that have already been the target of collecting efforts. The Armenian and Azerbaijani parts of the Lesser Caucasus remain somewhat less thoroughly explored compared to the Georgian part of the massive and to the Greater Caucasus as a whole. Despite the generally drier climate in the east, one or several new, relict brachyiulinine species may exist there in favourable places like high mountain forests or deep river gorges. And considering that three of the four brachyiulinine (sub)species known only from caves belong to the mostly Caucasian genus Omobrachyiulus (the fourth one being Titanophyllum spiliarum Akkari, Stoev & Enghoff, 2011), it would not be surprising if O. lazanyiae sp. nov. was not the only local member of the genus to occur in the subterranean realm (either as trogloxene or as a specifically adapted form).
Distribution of the Omobrachyiulus roseni (red and blue), implicitus (black) and hortensis (green) groups: O. roseni (filled circle), O. faxifer sp. nov. (star), O. zuevi sp. nov. (filled square), O. implicitus (cross), O. fasciatus sp. nov. (triangle), O. lazanyiae sp. nov. (drop), O. hortensis (ring), O. cf. hortensis (broken ring), O. armatus sp. nov. (pentagon), O. pristis sp. nov. (open square).
We are most grateful to Karin Voigtländer and Hans Reip (