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The millipede genera Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914 in the Caucasus, with notes on their distributions (Diplopoda, Julida, Julidae)
expand article infoAleksandr P. Evsyukov, Sergei I. Golovatch§, Dragan Ž. Antić|
‡ Don State Technical University, Rostov-on-Don, Russia
§ Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia
| University of Belgrade, Belgrade, Serbia
Open Access

Abstract

In the Caucasus, the genera Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914 are shown to include two and four species, respectively: Amblyiulus georgicus Lohmander, 1932, from Georgia and Armenia, A. hirtus sp. nov., from Azerbaijan and Dagestan, Russia, Syrioiulus adsharicus (Lohmander, 1936), from Georgia, S. continentalis (Attems, 1903), from Azerbaijan and Iran, S. taliscius (Attems, 1927), from Azerbaijan, and S. armeniacus sp. nov., from Armenia. All these six species are described, illustrated, and keyed, and their distributions are mapped and discussed, based on the literature data and abundant new samples.

Keywords

Faunistic records, key, map, new species, Pachyiulinae, Pachyiulini, taxonomy

Introduction

The very large family Julidae, of the basically Holarctic order Julida dominates the millipede faunas of Europe and the Mediterranean, marginally extending into the Oriental realm as well (Enghoff et al. 2015; Kime and Enghoff 2017). The subfamily Pachyiulinae, often referred to as the tribe Pachyiulini, encompasses between 15 and 20 genera, or 16–22 genera or subgenera, according to Antić et al. (2018) and Vagalinski (2020), respectively, and is characterised by the anterior gonopods being devoid of flagella, with a distinct sternum, both being fused mediobasally, and the posterior gonopods showing a mesomeral process, if any, only as an anterior branch of the opisthomere (Attems 1940; Tabacaru 1978). This group is monophyletic (Enghoff et al. 2013), temperate trans-Palaearctic, mostly restricted to the Mediterranean and ranges from Macaronesia in the west, through the entire Mediterranean, Central Asia, and central China, to Japan in the east. In the Caucasus proper, including the near-Caspian part of the Republic of Azerbaijan, but excluding the one in Iran (= both parts forming the Hyrcanian biogeographic province), this subfamily/tribe is currently known to be represented by three genera only.

The genus Pachyiulus Berlese, 1883 contains ca. 15 species of mostly very large julids which are largely confined to Southern and Southeastern Europe, the Near East, and the Caucasus (Enghoff et al. 2015). The Caucasus actually supports a single native species, P. krivolutskyi Golovatch, 1977, recently revised (Evsyukov 2016) and endemic to the western Caucasus (= Colchidan biogeographic province) within both Georgia and Russia (Kokhia and Golovatch 2020). One more congener, the eastern Mediterranean and synanthropic P. flavipes (C.L. Koch, 1847), has been introduced to the western Caucasus (Lohmander 1936), also being a very common, “tramp” species across Crimea (e.g., Golovatch 2008; albeit perhaps erroneously referred to as P. varius (Fabricius, 1781)).

The remaining two known genera of this tribe/subfamily which inhabit the Caucasus are Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914. The diagnoses and species compositions of these two genera, the main focus of the present contribution, remained unclear and confused for a long time, sometimes the latter genus being treated even as a synonym of the former (Tabacaru 1978, 1995). Mauriès (1982), in contrast to Tabacaru (1978), elevated Syrioiulus to a full genus and defined it primarily through a deeply bipartite posterior gonopod. He considered Syrioiulus, together with the monotypic Promeritoconus Verhoeff, 1943, from Turkey, and the species-rich genus Amblyiulus, as a single eastern Mediterranean lineage in the subfamily Pachyiulinae that is distinguished by the presence of eyes and, with a few exceptions only, 1+1 frontal setae, the development of apicoventral lobes on the male mandibular stipites, and of subequally high posterior gonopodal mesomeral process and opisthomere. Most, but not all, of the species from the Levant, Sporades (Greece), Caucasus, Iran and even Japan were thereby formally transferred to Syrioiulus. Some others from the same regions remained in or newly reassigned to Amblyiulus.

Enghoff (1992), in his review of Dolichoiulus Verhoeff, 1900, mentioned Syrioiulus only in passing. He, in his own outline of the pachyiuline generic classification, rediagnosed Amblyiulus and put the main emphasis on the structure of the posterior gonopods which show three apical processes, not two as is characteristic of several other genera, including Syrioiulus. Furthermore, he clearly illustrated the gonopods of A. barroisi (Porat, 1893), the type species of Amblyiulus.

The situation has become fully clarified only very recently, when first Golovatch (2018) and then Vagalinski (2020) confirmed the distinctions between Amblyiulus and Syrioiulus as lying solely in posterior gonopod conformation, also refining their diagnoses and scopes. Both these genera appear to be very similar, but differ clearly in the structure of the posterior gonopods: each of these being strongly divided into two branches (“bipartite”), i.e., a frontal (= mesomeral) and a caudal (= opisthomere) branch in Syrioiulus spp., vs. “tripartite”, with a third branch, a flagelliform rod, in Amblyiulus spp. (Golovatch 2018). A further refined account of the main differences between both these genera compared is given below.

Considering the above distinctions, Amblyiulus in the Caucasus appears to comprise only one described species: A. georgicus Lohmander, 1932. In addition, both Bababekova (1969, 1996) and Rakhmanov (1972) listed in Azerbaijan a dubious species, A. faliocius Attems, likely a misspelling of taliscius. The genus Syroiulus in the Caucasus presently includes three old species, all valid: S. adsharicus (Lohmander, 1936), S. continentalis (Attems, 1903), and S. taliscius (Attems, 1927). Vagalinski (2020), in his review and a provisional checklist of Syrioiulus species, emphasised the still-poor distinctions of Syrioiulus as opposed to a few pachyiuline genera other than Amblyiulus.

Materials and methods

All material has been shared between the collections of the Zoological Museum of the Moscow State University, Russia (ZMUM), the Senckenberg Museum of Natural History in Görlitz, Germany (SMNG), and the Institute of Zoology, University of Belgrade, Serbia (IZB). The specimens are stored in 70% ethanol. Some parts of males (antennae, gonopods, legs, etc.) and females (vulvae and leg pairs 2) were dissected and mounted in glycerol on temporary microscopic slides. Photographs were taken using a Zeiss StereoDiscovery V.20 microscope and processed with Zeiss ZEN software. Line drawings were executed using a camera lucida attached to a Radical light-transmission microscope. Scanning electron micrographs were taken with a Zeiss CrossBeam 340 (Rostov-on-Don State Technical University, Rostov-on-Don, Russia) or a JEOL JSM-6510LV (SMNG) scanning electron microscope (SEM). For some SEM micrographs, the gonopods were glued to a small sticky plastic triangle, placed on an SEM-stub, air dried for two days in a glass filled with Silica gel and finally coated with gold. After examination, material was removed from stubs and returned to alcohol. Live animals were photographed in situ using a Canon PowerShot SX120 IS digital camera.

The distribution map was created using Google Earth Pro 7.3.3 and processed in Adobe Photoshop CS6.

A “body ring formula” indicates the number of podous (including the gonopod-bearing segment/ring) and apodous segments/rings in an individual. This formula is p+a+T where p is the number of podous body rings, a the number of apodous body rings, and T represents the telson (Enghoff et al. 1993). Only adults have been analysed in the present study. For morphological descriptions, we largely used the terminology from Minelli (2015), for descriptions of the gonopods, that of Enghoff (1992) with changes in Vagalinski (2020).

The biogeographic regionalisation of the Caucasus follows the botanical one by Menitsky (1991).

No type material of the previously described species has been revised (mostly stored in the Zoological Institute, Russian Academy of Sciences, St. Petersburg) because of the 2020–2021 COVID pandemic, and the descriptive accounts and illustrations available in the literature are sufficiently complete and clear to allow a safe species identification. Colouration is largely described from preserved material. In the catalogue sections, D stands for a description or descriptive notes, R for new or repeated records, while M is a mere mention.

Refined characteristics of Amblyiulus vs. Syrioiulus

As shown below in the descriptions of individual species, the use of SEM allows for the distinctions between both genera concerned to be further refined. It is the opisthomere, not the entire posterior gonopod, that is bifid in Syrioiulus: a solenomere (with a distinct fovea or a deep saddle-like structure on top) and an anterior process (an anterior, lamellar branch adjacent to the solenomere) (Figs 7D, 10D, 12D–F, 14D). Using standard light microscopy, this anterior process often remains unnoticed, since it is very tightly appressed to the solenomere. In contrast, a third, anteromesal or lateral process/rod of the opisthomere is characteristic of Amblyiulus (Figs 3D, F, 5A, D–F, I, J).

As a result, it is only the structure of the opisthomere of the posterior gonopods that allows for the genera Amblyiulus and Syrioiulus to be more or less confidently diagnosed and separated. At the same time, species of Syrioiulus are mostly very similar in gonopodal conformation (Figs 7A–C, E, F, 10A–C, E, F, 12A–C, I, J, 14A–C, E–G), but they differ well in somatic characteristics, such as the presence or absence of eyes, frontal setae, and hairs on the rings (see also Key below). Similarly, within the genus Pachyiulus, several species were synonymised based solely on shared gonopodal characters (Mauriès et al. 1997), but later some have been revalidated on the basis of somatic features such as colouration (Frederiksen et al. 2012).

Taxonomic part

Amblyiulus Silvestri, 1896

Type species

Julus barroisi Porat, 1893, by original designation.

Diagnostic remarks

Here we follow Tabacaru’s (1978) opinion that the genera of the subfamily/tribe Pachyiulinae/-ini are best to be diagnosed using a complex of characters, both gonopodal and somatic. In the latest review of this tribe (Mauriès 1982), all genera are divided into three groups depending on the structure of the apical part of the opisthomere, viz., the presence/absence of a fovea and the presence/absence of a pseudoflagellum. He mistakenly assigned the genus Amblyiulus to group 3 (along with many other genera like Dolichoiulus), which have neither a fovea nor a pseudoflagellum. However, in accordance with our and earlier descriptions (e.g., Golovatch 2018), Amblyiulus has a fovea, however small, on the top of the solenomere. By the presence of a fovea and the absence of a pseudoflagellum, the genera Parapachyiulus Golovatch, 1979 and Dangaraiulus Golovatch, 1979 also join this group (Golovatch 1979, 2018). According to a number of other characters, such as the presence of frontal setae, apicoventral lobes on the male mandibles, and the mesomeral process being as high as the opisthomere, Amblyiulus belongs to Mauriès’ subgroup 3aa, together with the genera Syrioiulus and Promeritoconus. However, it seems noteworthy that sometimes frontal setae can be absent, while male mandibular stipites can remain unmodified.

The promere in Amblyiulus is narrowed in the basal third, in contrast to that in Promeritoconus, which is narrowed apically; in the apical part it may have one or two denticles, but sometimes none. The head can be with or without frontal setae. The eyes are mostly absent. The opisthomere of the posterior gonopod is tripartite: a solenomere, an anterior process, and an anteromesal or lateral rod, vs. bipartite in Syrioiulus.

Species included

Amblyiulus barroisi (Porat, 1893), Amblyiulus cedrophilus (Attems, 1910), Amblyiulus festae (Silvestri, 1895), Amblyiulus georgicus Lohmander, 1932, Amblyiulus hirtus sp. nov., and possibly several others, but their identity requires verification (Golovatch 2018).

Amblyiulus georgicus Lohmander, 1932

Figs 1A, 2, 3, 15A, 16

Amblyiulus georgicus Lohmander, 1932a: 180–182, figs 10–12 (D).

Amblyiulus georgicusLohmander 1936: 170 (M); Kobakhidze 1965: 395 (M); Lokšina and Golovatch 1979: 385 (M).

Syrioiulus georgicusMauriès 1984: 43 (M); Kokhia and Golovatch 2020: 207 (M).

Material examined

Armenia: 3 ♂♂, 5 ♀♀, 2 juv. (ZMUM), SW of Shnokh halfway between Alaverdi and Bagratashen, ca. 1500 m a.s.l., Carpinus forest, litter, 24.V.1987; 2 ♀♀ (ZMUM), Odzun W of Alaverdi, 1500–1550 m a.s.l., Quercus, Fagus, Carpinus, etc. forest, litter and under stones with ants, 23–24.V.1987; all leg. S. Golovatch, K. Eskov.

Diagnosis

Differs from A. hirtus sp. nov., apparently the most similar and geographically the closest congener known to date, by the following combination of somatic and gonopodal characteristics. Head without frontal setae; collum and metazonae of body rings without setae. Male mandibular stipites expanded. Promere wide, with two apical denticles. Solenomere with a membranous lobe notched apically. Rod of opisthomere relatively short. See also Key below.

Redescription

Length of adults 27–30 mm (♂♂) or 28–31 mm (♀♀), width 1.6–1.7 mm (♂♂) or 1.7–1.9 mm (♀♀). Number of body rings in adults, 65–67+2+T (♂♂) or 67–69+2+T (♀♀). Body subcylindrical (typical of Julidae), metazonae brownish grey, prozonae yellowish grey (Fig. 1A). Head, a few postcollum rings and telson lighter than other body rings. Collum more vividly red-brown. Antennae, mouthparts, and legs yellow (Fig. 2A–C). Eyes absent. Metazonae with weak and irregular striations, 14–16 striae between dorsal axial line and ozopore (Fig. 2D). Ozopores relatively large, with a stria in front and lying behind suture without touching it (Fig. 2H).

Figure 1. 

Habitus of Amblyiulus and Syrioiulus species, males, lateral views A A. georgicus Lohmander, 1932 from Shnokh, Armenia (ZMUM) B A. hirtus sp. nov., paratype from Bash-Layski, Azerbaijan (ZMUM) C S. adsharicus (Lohmander, 1936) from Adigeni, Georgia (ZMUM) D S. continentalis (Attems, 1903) from Istisu, Azerbaijan (ZMUM) E S. taliscius (Attems, 1927) from Avrora, Azerbaijan (ZMUM) F S. armeniacus sp. nov., paratype from Shikahoh, Armenia (ZMUM). Scale bars: 1.0 mm.

Antennae relatively long, in situ reaching segment 4. Head without frontal setae, but with 8+8–9+9 labral and 2+2 supralabral setae (Fig. 2A–C). Gnathochilarium with four setae on each lamella lingualis, stipites with a group of several short setae in medial part and three long setae at anterolateral margin (Fig. 2I). Collum and metazonae without setae at posterior margin (Figs 1A, 2A–G). Epiproct undeveloped (Fig. 2E, F). Hypoproct subtriangular, with several long setae (Fig. 2G). Telson and anal valves sparsely setose, setae being long (Fig. 2E–G).

Figure 2. 

Amblyiulus georgicus Lohmander, 1932, ♂ from Shnokh, Armenia (ZMUM) A–C anterior part of body, lateral, dorsal and ventral views, respectively D midbody part, lateral view E–G posterior part of body, lateral, dorsal and ventral views, respectively H ozopores on midbody rings, lateral view I gnathochilarium, ventral view J leg pair 1, caudal view K leg 2, caudal view L ventral edge of pleurotergum 7, lateral view. Scale bars: 0.5 mm (A–G); 0.1 mm (H–L).

Male. Mandibular stipites expanded, slightly swollen in distal part (Fig. 2A). Leg pair 1 small, unciform, with a group of setae on each coxa and at base of telopodite; telopodite relatively long (Fig. 2J). Leg pair 2 with pads on postfemur and tibia (Fig. 2K). Penes short, bifurcate on top. Ventral edge of male segment 7 with elongated and rounded lamellae bordering the gonopodal aperture (Fig. 2L).

Gonopods (Fig. 3). Promere spoon-shaped, relatively wide, constricted in basal third; mesal ridge in apical part forming a small mesal denticle (Fig. 3B, E). Lateral denticle large, well-developed. Mesomeral process simple, flattened, ribbon-shaped, notched on top (Fig. 3A, C, F). Opisthomere tripartite (Fig. 3D). Solenomere long, slightly curved, with caudomesal lamella notched apically; apical part with a fovea and a pointed membranous process. Anterior process of opisthomere appressed to solenomere, with a rounded apex. Anterolateral part of opisthomere with a helicoid rod.

Figure 3. 

Amblyiulus georgicus Lohmander, 1932, ♂ from Shnokh, Armenia (ZMUM) (A–D) or holotype ♂, after Lohmander (1932) (E, F). A gonopod, mesal view B promere, caudal view C posterior gonopod, caudal view D end of solenomere, mesal view E promere, caudal view F posterior gonopod, lateral view. Abbreviations: ap anterior process cl caudomesal lamella fo fovea ld lateral denticle md mesal denticle mr mesal ridge ms mesomeral process op opisthomere pr promere r rod sl solenomere. Scale bars: 0.1 mm (A–C); 0.01 mm (D); not to scale (E, F).

Female. First two leg pairs unmodified. Vulva rounded, operculum and bursa equal in height (Fig. 15A). Operculum at apical margin oblique, undivided. Bursa asymmetric, lateral valve higher than mesal one. Each valve with two rows of long setae. Median field of bursa very short, narrow; emargination of median field suboval.

Remarks

This species was described from Borjom (= Borjomi), Georgia (Lohmander 1932a). The above samples represent the first formal records of this species from Armenia. It seems to populate high-montane deciduous forests in the western part of the Caucasus Minor (= Lesser Caucasus) (Fig. 16).

Amblyiulus hirtus sp. nov.

Figs 1B, 4, 5, 15B, 16

Material examined

Holotype ♂ (ZMUM), Azerbaijan, NW above Bash-Layski ca. 20 km NNW of Sheki, 1250 m a.s.l., Fagus, Carpinus, Acer, etc. forest, litter, 3.V.1987, leg. S. Golovatch, K. Eskov. Paratypes: 5 ♂♂, 3 ♀♀ (ZMUM), same collection data as holotype.

Non-type material

Azerbaijan: 2 ♂♂, 6 ♀♀ (ZMUM), SW of Kuba, 750 m a.s.l., Fagus, Quercus, Carpinus, etc. forest, litter and under bark, 23.IV.1987, leg. S. Golovatch, K. Eskov; Russia, Dagestan: 1 ♂, 2 ♀♀ (ZMUM), Kurush, 2550 m a.s.l., S slope, subalpine and alpine meadows, 20.VIII.1990, leg. G. Magomedov.

Diagnosis

Assigned to the genus Amblyiulus primarily because of the presence of a rod on the posterior gonopod opisthomere. Differs from A. georgicus, perhaps the most similar congener known to date, by the following combination of somatic and gonopodal characters. Head with frontal setae; collum and metazonae of body rings each with a posterior whorl of setae. Promere narrow, with two side ridges. Solenomere apically with small filament-like processes. Rod of opisthomere relatively long.

Name

To emphasise the presence of metazonal setae; adjective.

Description

Holotype : length 27 mm, width 1.3 mm, number of body rings 51+2+T. Paratypes: length 25–30 mm, width 1.2–1.4 mm, number of body rings in adults, 45–67+1–3+T (♂♂); or length 27–28 mm, width 1.1–1.3 mm, number of body rings, 46–55+2–3+T (♀♀). Body subcylindrical (typical of Julidae), metazonae and prozonae yellowish grey (Fig. 1B). Head, a few postcollum rings and telson slightly lighter than other body rings. Collum slightly more vividly reddish. Antennae, mouthparts, and legs yellow (Fig. 4A–C). Eyes absent. Metazonae with weak, dense, and regular striations, 21–23 striae per quarter of metazonal surface, i.e., that between dorsal axial line and ozopore (Fig. 4A–G). Ozopores relatively large, with a stria in front, lying behind suture without touching it (Fig. 4H).

Figure 4. 

Amblyiulus hirtus sp. nov., paratype ♂ from Bash-Layski, Azerbaijan (ZMUM) A–C anterior part of body, lateral, dorsal and ventral views, respectively D midbody part, lateral view E–G posterior part of body, lateral, dorsal and ventral views, respectively H ozopores on midbody rings, lateral view I gnathochilarium, ventral view J, K leg pair 1, lateral and caudal views, respectively L leg 2, caudal view M ventral edge of pleurotergum 7, lateral view. Scale bars: 0.5 mm (A–G) or 0.1 mm (H–M).

Antennae relatively long, in situ reaching ring 4. Head with 1+1 frontal, 8+8–9+9 labral and 2+2 supralabral setae (Fig. 4A–C). Gnathochilarium with three thick setae on each lamella lingualis; stipites without setae in medial part, but with three long setae at anterolateral margin (Fig. 4I). Collum and each following metazona with a whorl of setae at posterior margin (Fig. 4A). Epiproct poorly developed, triangular, with several setae (Fig. 4E, F). Hypoproct subtriangular, covered with long setae (Fig. 4G). Telson and anal valves densely setose, setae being long.

Male. Mandibular stipites unmodified (Fig. 4A). Leg pair 1 small, unciform, with a group of setae on coxa and at base of telopodite; telopodites curved anteriad, not anteromesad as in other species of Julidae (Fig. 4J, K). Leg pair 2 with a large pad on tibia and a small one on postfemur (Fig. 4L). Penes short and bifurcate. Ventral edge of male pleurotergum 7 with small subtriangular lamellae bordering the gonopodal aperture (Fig. 4M).

Gonopods (Fig. 5) with anterior and posterior pair equal in height. Promere spoon-shaped, relatively narrow, constricted in basal third; with two ridges: mesal ridge prominent all along; lateral ridge short, located only in apical part of promere (Fig. 5B, H). Mesomeral process simple, flattened, ribbon-shaped, with a small membranous lobe on top (Fig. 5A, C, G, I). Opisthomere tripartite (Fig. 5A, C, E, F). Solenomere long, slightly curved, with a caudomesal lamella notched apically; apical part with a fovea and short filament-like processes (Fig. 5A, C, G, I). Solenomere sometimes with an additional filiform process apically (see Remarks under Syrioiulus taliscius). Anterior process notched apically (Fig. 5A, C, E). Rod of solenomere relatively long, consisting of filament-like structures, lateral in position (Fig. 5A, D–G, I).

Figure 5. 

Amblyiulus hirtus sp. nov., paratype ♂ from Bash-Layski, Azerbaijan (ZMUM) A, G gonopod, mesal and lateral views, respectively B, H promere, subcaudal and caudal views, respectively C, I posterior gonopod, lateral and mesal views, respectively D, E end of solenomere, mesal and submesal views, respectively F end of opisthomere, mesocaudal view. Abbreviations: ap anterior process cl caudomesal lamella ff filiform process fo fovea lr lateral ridge mr mesal ridge ms mesomeral process op opisthomere pr promere r rod sl solenomere. Scale bars: 0.1 mm (A–C); 0.01 mm (D–F); 0.2 mm (G–I).

Female. First two leg pairs unmodified. Vulva rounded, operculum higher than bursa (Fig. 15B) and bilobed apically. Bursa asymmetric, lateral valve higher than mesal one. Each valve with two rows of long setae. Median field of bursa very short, narrow; emargination of median field suboval.

Remarks

This species seems to be endemic to the eastern part of the Caucasus Major within both northeastern Azerbaijan and the Republic of Dagestan, Russia (Fig. 16).

It is the presence of a laterally positioned rod that brings both A. georgicus and A. hirtus sp. nov. particularly close together. However, the rod in these two species is located laterally, whereas that in A. barroisi anteromesally (Enghoff 1992: fig. 11; Golovatch 2018: fig. 10C). These differences seem to be quite important, but because those three species share not only the presence of a rod, but also a small, but discernible fovea on top of the solenomere, for the time being it seems best to regard the trio as members of Amblyiulus.

Syrioiulus Verhoeff, 1914

Type species

Dolichoiulus polyzonus Attems, 1910, by subsequent designation of Jeekel (1971).

Diagnosis

All characters as in Amblyiulus, except as follows. Promere usually with two denticles in apical part. Head with or without frontal setae. Eyes present or absent. Opisthomere of posterior gonopod bipartite: a solenomere (with a distinct fovea on top) and an anterior process, vs. tripartite in Amblyiulus.

Species included

Syrioiulus adsharicus (Lohmander, 1936), Syrioiulus andreevi Mauriès, 1984, Syrioiulus aharonii (Verhoeff, 1914), Syrioiulus armeniacus sp. nov., Syrioiulus continentalis (Attems, 1903), Syrioiulus discolor (Lohmander, 1932), Syrioiulus incarnatus (Lohmander, 1932), Syrioiulus lohmanderi Vagalinski, 2020, Syrioiulus persicus (Golovatch, 1983), Syrioiulus polyzonus (Attems, 1910), Syrioiulus taliscius (Attems, 1927), and several others provisionally listed by Vagalinski (2020).

Syrioiulus adsharicus (Lohmander, 1936)

Figs 1C, 6, 7, 15C, 16

Amblyiulus (Heteropachyiulus) adsharicus Lohmander, 1936: 156–159, figs 131, 132 (D).

Amblyiulus adsharicusKobakhidze 1964: 191 (M); Lokšina and Golovatch 1979: 385 (M); Talikadze 1984: 143 (M); Kokhia and Golovatch 2018: 40 (M).

Syrioiulus adsharicusVagalinski 2020: 89 (M); Kokhia and Golovatch 2020: 207 (M).

Material examined

Georgia: 10 ♂♂, 16 ♀♀, 5 juv. (ZMUM), 15 km W of Adigeni, Abies, Picea, Fagus, Acer, etc. forest, 1500–1700 m a.s.l., litter, logs, under stones, 14–15.V.1983, leg. S. Golovatch; 4 ♂♂, 5 ♀♀ (ZMUM), near Adigeni, 9.VI.1977, leg. V. Dolin.

Diagnosis

Differs from all congeners by the following combination of somatic and gonopodal characters. Head with frontal setae. Collum and each metazona of following body rings with a whorl of long setae at caudal margin. Ommatidia present, but only a few ommatidia, all unpigmented and very small. Solenomere with small denticles apically. Anterior process of opisthomere with small filament-like spines apically.

Redescription

Length of adults 17–30 mm (♂♂) or 18–31 mm (♀♀), width 1.2–1.4 mm (♂♂) or 1.3–1.7 mm (♀♀). Number of body rings in adults, 50–63+1–2+T (♂♂) or 52–60+1–2+T (♀♀). Body subcylindrical (typical of Julidae), metazonae brownish grey, prozonae violet grey (Figs 1C, 6A, C). Head, collum and body rings from yellow to greyish yellow. Antennae, mouthparts and first leg pairs yellow, other pairs brown (Fig. 6A, C–H). Eyes present, unpigmented, very small, composed of 3–7 ommatidia, unequal numbers on opposite sides of head (Fig. 6B). Metazonae with weak striations, 17–19 striae per quarter of metazonital surface, i.e., that between dorsal axial line and ozopore (Fig. 6E). Ozopores small, lying behind suture and touching it (Fig. 6I).

Figure 6. 

Syrioiulus adsharicus (Lohmander, 1936), ♂ from Adigeni, Georgia (ZMUM) A, C, D anterior part of body, lateral, dorsal and ventral views, respectively B Eye, lateral view E midbody part, lateral view F–H posterior part of body, lateral, dorsal and ventral views, respectively I ozopores on midbody rings, lateral view J gnathochilarium, ventral view K leg pair 1, caudal view L leg pair 2, caudal view M ventral edge of pleurotergum 7, lateral view. Scale bars: 0.5 mm (A, C–I); 0.1 mm (B); 0.05 mm (J–M).

Antennae relatively long, in situ reaching segment 4. Head with 1+1 frontal, 11+11–13+13 labral and 2+2 supralabral setae (Fig. 6C, D). Gnathochilarium with four long setae on each lamella lingualis, stipites with a medial group of 7–10 short setae, three long and two short setae at anterolateral margin (Fig. 6J). Collum and each metazona of following rings with a whorl of long setae at posterior margin (Fig. 6A, E, F–H). Epiproct undeveloped (Fig. 6F, G). Hypoproct subtriangular, with long setae (Fig. 6H). Anal valves densely setose, setae being long.

Male. Mandibular stipites expanded, slightly swollen (Fig. 6A). Leg pair 1 small, unciform, telopodites curved anteromesad (as in most other Julidae), with a group of setae on each coxa and at base of telopodite (Fig. 6K). Leg pair 2 with pads on postfemur and tibia (Fig. 6L). Penes short, bifurcate. Ventral edge of male pleurotergum 7 with relatively wide and apically rounded lamellae bordering the gonopodal aperture (Fig. 6M).

Gonopods (Fig. 7) with anterior and posterior pair equal in height. Promere spoon-shaped, constricted in basal third; a mesal ridge along basal 2/3 extent; two apical denticles well-developed, mesal one vertical and with a weakly bifurcate apex, lateral one short, wide and rounded apically (Fig. 7B, E). Mesomeral process simple, flattened, ribbon-shaped, bifurcate (Fig. 7C, F). Opisthomere bipartite (Fig. 7D, C). Solenomere long, erect, with small denticles apically, bearing a fovea at apex; caudomesal lamella wide with a notched apical margin (Fig. 7A, C, F). Anterior process apically with small filament-shaped spines (Fig. 7D).

Figure 7. 

Syrioiulus adsharicus (Lohmander, 1936), ♂ from Adigeni, Georgia (ZMUM) (A–D) or holotype ♂, after Lohmander (1936) (E, F) A, F gonopod, mesal view B, E promere, caudal view C posterior gonopod, lateral view D end of solenomere, lateral view. Abbreviations: ap anterior process cl caudomesal lamella fo fovea ld lateral denticle md mesal denticle mr mesal ridge ms mesomeral process op opisthomere pr promere sl solenomere. Scale bars: 0.1 mm (A–C); 0.03 mm (D); not to scale (E, F).

Female. First two leg pairs unmodified. Vulva elongated, covered with long setae (Fig. 15C). Operculum relatively low, deeply divided. Bursa asymmetric, lateral valve higher than mesal one. Median field of bursa narrow; emargination of median field narrow and elongated.

Remark

This species was originally described from Batumi, “Bortschacha” (Lohmander 1936). Our new record from near Adigeni is evidence of the species likely to be endemic to the southern part of the Colchidan biogeographic province, all within Georgia (Fig. 16).

Syrioiulus continentalis (Attems, 1903)

Figs 1D, 8A, B, 9, 10, 15D, 16

Pachyiulus (Dolichoiulus) continentalis Attems, 1903: 147, 148, figs 82–84 (D).

Amblyiulus continentalisLohmander 1932b: 40, 41, figs 33–35 (D); 1936: 156 (R); Rakhmanov 1971: 1412 (R); 1972: 116 (R); Samedov et al. 1972: 1245; Lokšina and Golovatch 1979: 385 (M); Bababekova 1996: 90 (M).

Syrioiulus continentalisMauriès 1982: 441 (M); 1984: 43 (M); Vagalinski 2020: 89 (M).

Material examined

Azerbaijan: 3 ♂♂, 1 ♀ (ZMUM), Talysh Mts, Zuvand, Joni, 1500 m a.s.l., 28–29.V.1976, leg. V.G. Dolin; 1 ♂ (ZMUM), Lenkoran, Hyrcan forest, Khan Bulan River near Alexeevka, 22.IV.1985, leg. E.B. Kupriyanova; 7 ♂♂, 5 ♀♀, 2 juv. (ZMUM), Lenkoran, Hyrcan Nature Reserve, litter, 26.I.–4.II.1985, leg. A. Druk; 2 ♂♂, 4 ♀♀ (ZMUM), same locality, 21.IX.1987, leg. S. Zonstein; 2 ♂♂, 2 juv. (IZB); 1 ♂, 1 ♀, 1 juv. (SMNG), Lǝnkǝran rayon, Hyrcan Nature Reserve, Daştatük 1.3 km Xanbulan Reservoir, Parrotia forest, diverse bushes, under leaves, 110 m a.s.l., 38.6747°N, 48.7622°E; 1 ♂ (IZB); 1 ♂ (SMNG), same locality, SW of Aşağı Apu, Quercus forest, within leaves and rotten wood, 180 m a.s.l., 38.6726°N, 48.7362°E, all leg. F. Walther, H. Reip, D. Antić; 3 ♂♂, 2 ♀♀, 1 juv. (IZB); 5 ♂♂, 3 ♀♀ (SMNG), Lerik rayon, Hyrcan Nature Reserve, road Lǝnkǝran–Lerik at km 32, small side valley, forest of Parrotia with some Quercus, thick leaf layer, 400 m a.s.l., 38.7638°N, 48.5819°E; 1 juv. (IZB), Astara rayon, Hyrcan Nature Reserve, SW of Zünqülǝş, beginning of a small valley, Parrotia and Alnus bushes, in leaves, 60 m a.s.l., 38.4493°N, 48.7623°E; 2 ♂♂, 1 juv. (IZB); 2 ♂♂, 1 ♀ (SMNG), same locality, end of small valley, steep slope, Parrotia, Quercus, Acer trees, under leaves and rotten tree trunks, 130 m a.s.l., 38.4480°N, 48.7597°E, all leg. F. Walther, H. Reip, D. Antić; 2 ♂♂ (ZMUM), Azfilial, 100 m a.s.l., 31.V.–1.VI.1996; 1 ♂ (ZMUM), Apo below Bilasar, 350 m a.s.l., 8–9.VI.1996; 2 ♂♂, 1 ♀, 1 juv. (ZMUM), Astara Distr., Istisu ca. 8 km WSW of Astara, Quercus, Acer, Carpinus, etc. forest, 10–30 m a.s.l., litter, under stones and bark, 10.X.1983; 2 ♀♀ (ZMUM), Istisu ca. 8 km SW of Masally, Quercus, Acer, Carpinus etc. forest, 80–140 m a.s.l., under bark and stones, 19–20.X.1983; 2 ♀♀ (ZMUM), Istisu W of Astara, 100 m a.s.l., 2–6.VI.1996, all leg. S. Golovatch.

Diagnosis

Differs from all congeners by the following combination of somatic and gonopodal characters. Head with frontal setae. Collum and each metazona of following body rings with a whorl of long setae at caudal margin. Eyes present. Solenomere with a group of small spines on top. Anterior process of opisthomere subtriangular apically. This species is clearly distinguished in the field from all other millipedes by its characteristic greyish yellow colouration with a yellow stripe dorsally, and its particularly strong odour clearly resembling that of Pachyiulus krivolutskyi from the western Caucasus (= Colchis).

Redescription

Length of adults 28–45 mm (♂♂) or 26–46 mm (♀♀), width 2.0–2.3 mm (♂♂) or 2.2–2.7 mm (♀♀). Number of body rings in adults, 46–66+1–2+T (♂♂) or 49–66+1–2+T (♀♀). Body subcylindrical, metazonae from greyish yellow to yellow, prozonae light yellow (Figs 1D, 8A, B); live specimens dorsally with a darker, vivid yellow stripe (Fig. 8B). Head, collum and telson slightly lighter than other body rings (Fig. 8A, B). Antennae grey, mouthparts and legs light yellow (Fig. 9A–G). Eyes present, black, oval, each composed of 19–23 ommatidia (Fig. 9A, C). Striations on metazonae deep, not reaching the caudal margin, 28–32 striae per quarter of metazonal surface, i.e., between dorsal axial line and ozopore (Fig. 9D). Ozopores large, lying behind suture without touching it (Fig. 9H).

Figure 8. 

Live colouration of two Syrioiulus species from Hyrcan Nature Reserve, Azerbaijan A, B S. continentalis (Attems, 1903) C, D S. taliscius (Attems, 1927). Pictures by D.Ž. Antić, not taken to scale.

Antennae relatively long, in situ reaching segment 3. Head with 1+1 frontal, 8+8–9+9 labral and 2+2–4+4 supralabral setae (Fig. 9A–C). Gnathochilarium with three long setae on each lamella lingualis, groups of several small setae in median part of stipites and six or seven long setae at anterolateral margin (Fig. 9I). Collum and each following metazona with a whorl of long and thick setae at posterior margin (Fig. 9A–F). Epiproct undeveloped (Fig. 9E, F). Hypoproct subtriangular, with long setae (Fig. 9G). Telson and anal valves densely setose, setae being long.

Figure 9. 

Syrioiulus continentalis (Attems, 1903), ♂ from Istisu, Azerbaijan (ZMUM) A–C anterior part of body, lateral, dorsal and ventral views, respectively D midbody part, lateral view E–G posterior part of body, lateral, dorsal and ventral views, respectively H ozopores on midbody rings, lateral view I gnathochilarium, ventral view J leg pair 1, caudal view K leg 2, caudal view L ventral edge of pleurotergum 7, lateral view. Scale bars: 1.0 mm (A–G); 0.1 mm (H–L).

Male. Mandibular stipites expanded, with swollen lobes (Fig. 9A). Leg pair 1 small, unciform, telopodites directed anteromesad, with a group of long setae on each coxa; telopodite setose in basal part (Fig. 9J). Leg pair 2 with pads on postfemur and tibia (Fig. 9K). Penes short, bifurcate. Ventral edge of male pleurotergum 7 with narrow elongated lamellae bordering the gonopodal aperture (Fig. 9L).

Gonopods (Fig. 10) with anterior (promere) part higher than posterior (opisthomere) one. Promere spoon-shaped, constricted in basal third; mesal ridge wide along 2/3 extent; with denticles in apical part: mesal denticle small and broadly rounded, lateral one well-expressed and long (Fig. 10B, E). Mesomeral process simple, slightly curved, flattened apically (Fig. 10A, C, F). Opisthomere bipartite (Fig. 10D). Solenomere elongated, with an apical membranous lobe, subtriangular at apex, with a fovea and a group of small spines on top; caudomesal lamella wide (Fig. 10A, C, D, F). Anterior process of opisthomere subtriangular apically (Fig. 10D).

Figure 10. 

Syrioiulus continentalis (Attems, 1903), ♂ from Hyrcan Nature Reserve (A–C) (SMNG), from Istisu (D) (ZMUM) or after Lohmander (1932b) (E, F) A gonopod, mesal view B, E promere, caudal view C, F posterior gonopod, lateral view D end of solenomere, lateral view. Abbreviations: ap anterior process cl caudomesal lamella fo fovea ld lateral denticle md mesal denticle mr mesal ridge ms mesomeral process op opisthomere pr promere sl solenomere. Scale bars: 0.1 mm (A–C); 0.01 mm (D); not to scale (E, F).

Female. First two leg pairs unmodified. Operculum of vulva without setae on caudal surface, apical margin relatively flat (Fig. 15D). Bursa subsymmetrical, lateral valve slightly larger than mesal one. Each valve with two rows of long setae. Median field of bursa narrow; emargination of median field suboval.

Remarks

Probably one of the most common and apparently the largest species of the genus. The unusually strong odour and the chemical composition of the repugnatorial secretion are similar to those of Pachyiulus krivolutskyi (Makarov et al., pers. obs.). This species inhabits various deciduous forests in Azerbaijan, also occurring in northern Iran (Lohmander 1932b), endemic to the Hyrcanian biogeographic province (Fig. 16).

Syrioiulus taliscius (Attems, 1927)

Figs 1E, 8C, D, 11, 12, 15E, 16

Amblyiulus taliscius Attems, 1927: 243, 244, figs 336–338 (D).

Amblyiulus talisciusLohmander 1932b: 182 (M); 1936: 170 (M); Rakhmanov 1972: 116 (R); Lokšina and Golovatch 1979: 385 (M); Bababekova 1996: 90 (M).

Syrioiulus talisciusMauriès 1982: 441 (M); 1984: 43 (M); Vagalinski 2020: 92 (M).

Material examined

Azerbaijan: 4 ♂♂, 14 ♀♀, 1 juv. (ZMUM), Talysh, Joni, 1500 m a.s.l., 28–29.V.1976, leg. V.G. Dolin; 2 ♂♂ (ZMUM), Lenkoran, Hyrcan Nature Reserve, Telman, 28.IV.1984; 1 ♂, 1 ♀ (ZMUM), same locality, Gaftoni, 9.V.1984, all leg. H. Aliev; 1 ♀ (ZMUM), Hyrcan forest, Khan Bulan River near Alexeevka, 22.IV.1985, leg. E.B. Kupriyanova; 3 ♂♂, 12 ♀♀ (ZMUM), same locality, Avrora, Moscow-Forest, 50 m a.s.l., 1.VI.1996; 3 ♂♂, 3 ♀♀, 1 juv. (ZMUM), same locality, Apo below Bilasar, 350 m a.s.l., 8–9.VI.1996, all leg. S. Golovatch; 1 ♂ (SMNG), Lǝnkǝran rayon, Siyablı, Parrotia, Zelkova, Quercus coppice, steep slope, 110 m a.s.l., 38.7170°N, 48.7253°E; 1 ♂ (IZB), Lerik rayon, Hyrcan Nature Reserve, road Lǝnkǝran–Lerik at km 32, small side valley, forest of Parrotia with some Quercus, thick leaf litter, 400 m a.s.l., 38.7638°N, 48.5819°E; 7 ♀♀ (IZB), 7 ♀♀ (SMNG), Astara rayon, Hyrcan Nature Reserve, SW of Zünqüləş, end of small valley, steep slope, Parrotia, Quercus, Acer trees, under leaves and rotten tree trunks, 130 m a.s.l., 38.4480°N, 48.7597°E; 2 ♂♂, 1 ♀, 1 juv. (IZB); 2 ♂♂, 2 ♀♀, 1 juv. (SMNG), Lǝnkǝran rayon, Hyrcan Nature Reserve, Daştatük 1.3 km Xanbulan Reservoir, Parrotia forest, divers bushes, under leaves, 110 m a.s.l., 38.6747°N, 48.7622°E; 3 ♂♂, 2 ♀♀, 1 juv. (IZB); 5 ♂♂ (1 SEM), 3 ♀♀ (1 SEM), 1 juv. (SMNG) Lǝnkǝran rayon, Hyrcan Nature Reserve, SW of Aşağı Apu, Quercus forest, within leaves and rotten wood, 180 m a.s.l., 38.6726°N, 48.7362°E, all leg. F. Walther, H. Reip, D. Antić; 1 ♂ without gonopods (ZMUM), Shemakha Distr., farm Guseinzade, foothills, Vitis, summer 1982, leg. A. Ismailov; 1 ♀ (ZMUM), Zakataly Distr., Geyam, cornfield, 17.IV.1986, leg.?; 7 ♂♂, 5 ♀♀ (ZMUM), Baku, City parks, 18–21.V.1981; 1 ♂ (ZMUM), Belokani near Zakatali, 600 m a.s.l., village garden, 24.V.1981; 1 ♀ (ZMUM), above Akhsu 120 km W Baku, 900 m a.s.l., Quercus shrub, 22.V.1981, all leg. S. Golovatch and J. Martens; 1 ♀ (ZMUM), ca. 14 km W of Ismailly, Galyhjakh, under bark, 1.V.1987; 1 ♀ (ZMUM), Altyagach, 1050–1100 m a.s.l., Quercus, Fagus, Carpinus, etc. forest, litter, 20 and 26.IV.1987, all leg. S.Golovatch and K. Eskov; 3 ♂♂, 5 ♀♀, 1 juv. (IZB); 3 ♂♂, 5 ♀♀ (SMNG), İsmayıllı rayon, Topçu 7.8 km towards Vəndam, flat area with old Fagus forest, under leaves and dead wood, 630 m a.s.l., 40.9193°N, 48.0027°E; 1 ♂ (SMNG), İsmayıllı rayon, Xanəgah 2 rkm towards İsmayıllı, flat area with old Fagus forest, with channels, under leaves, 650 m a.s.l., 40.8233°N, 48.1518°E; 1 ♀ (SMNG), İsmayıllı rayon, S of Zərgəran, slope with Corylus, Clematis and some Prunus trees, stone heaps overgrown by moss, mainly in thick leaf litter and under stones, 880 m a.s.l., 40.7310°N, 48.3680°E, all leg. F. Walther, H. Reip, D. Antić.

Diagnosis

Differs from all congeners by the following combination of somatic and gonopodal characters. Head without frontal setae. Collum and metazonae of following body rings without setae. Eyes absent. Solenomere in apical part with a group of small spines. Anterior process of opisthomere subtriangular apically.

Redescription

Length of adults 26–33 mm (♂♂) or 26–34 mm (♀♀), width 1.2–1.3 mm (♂♂) or 1.2–1.4 mm (♀♀). Number of body rings in adults, 50–65+1–2+T (♂♂) or 49–70+1–2+T (♀♀). Body subcylindrical (typical of Julidae), live specimens with brownish grey pro- and metazonae (Fig. 8C, D), after storage in alcohol prozonae grey, metazonae yellow (Fig. 1E). Head, collum, a few postcollum rings, last body rings, telson and anal valves yellow (Figs 1E, 8C, D). Antennae, mouthparts, and legs light yellow (Fig. 11A–G). Eyes absent. Striations of metazonae deep, not reaching the caudal margin, 23–25 striae per quarter of metazonital surface, i.e., that between dorsal axial line and ozopore (Fig. 11D). Ozopores small, lying behind suture without touching it (Fig. 11H).

Figure 11. 

Syrioiulus taliscius (Attems, 1927), ♂ from Avrora, Azerbaijan (ZMUM) A–C anterior part of body, lateral, dorsal and ventral views, respectively D midbody part, lateral view E–G posterior part of body, lateral, dorsal and ventral views, respectively H ozopores on midbody rings, lateral view I gnathochilarium, ventral view J leg pair 1, caudal view K leg 2, caudal view L ventral edge of pleurotergum 7, lateral view. Scale bars: 0.5 mm (A–G); 0.1 mm (H–L).

Antennae relatively long, in situ reaching segment 3. Head without frontal setae, 9+9–12+12 labral and 2+2 supralabral setae (Fig. 11A–C). Collum and metazonae without setae (Fig. 11A–G). Gnathochilarium with 3–4 long setae on each lamella lingualis, stipites with a medial curved row of 4–5 thick setae and three long setae at anterolateral margin (Fig. 11I). Epiproct undeveloped (Fig. 11E, F). Hypoproct rounded, with several setae (Fig. 11G). Telson covered with long setae, anal valves densely setose.

Male. Mandibular stipites unmodified (Fig. 11A). Leg pair 1 small, unciform, telopodites directed anteromesad (as in most Julidae), with long setae on each coxa and in basal part of telopodite (Fig. 11J). Leg pair 2 with pads on postfemur and tibia (Fig. 11K). Penes short, bifurcate. Ventral edge of male pleurotergum 7 with wide curved lamellae bordering the gonopodal aperture (Fig. 11L).

Gonopods (Fig. 12) with anterior and posterior parts both equal in height. Promere spoon-shaped, constricted in basal third; mesal ridge relatively narrow along 2/3 extent; with denticles in apical part: mesal denticle small and broadly rounded, lateral one well-expressed and long (Fig. 12B, H). Mesomeral process simple, slightly curved, with a wide subquadrate lamella apically (Fig. 12A, C, G, I). Opisthomere bipartite. Solenomere elongated, with a caudomesal, notched, membranous lobe, in apical part with a fovea and a group of small spines (Fig. 12A, C–F). Fovea may be equipped with a filiform process (Fig. 12E). Anterior process subtriangular apically.

Figure 12. 

Syrioiulus taliscius (Attems, 1927), ♂ from Hyrcan Nature Reserve (A–C) (SMNG), from Avrora (D, G, H) (ZMUM) or from Apo, Azerbaijan (E, F) (ZMUM) A, G gonopod, mesal and lateral views, respectively B, H promere, caudal view C, I posterior gonopod, caudolateral and mesal views, respectively D–F end of solenomere, mesal views. Abbreviations: ap anterior process cl caudomesal lamella fo fovea ld lateral denticle md mesal denticle mr mesal ridge ms mesomeral process op opisthomere pr promere sl solenomere. Scale bars: 0.1 mm (A–C); 0.02 mm (D–F); 0.2 mm (G–I).

Female. First two leg pairs unmodified. Operculum of vulva without setae on caudal surface, apical margin poorly divided (Fig. 15E). Bursa mostly symmetric, lateral valve slightly larger than mesal one. Each valve with two rows of long setae. Median field of bursa narrow; emargination of median field suboval.

Remarks

This species was described from the Talysh Mts, Lenkoran, Azerbaijan (Attems 1927). In the Caucasus, this is probably one of the most common and widespread congeners. Like S. continentalis, it inhabits various deciduous forests, but it can only be considered as subendemic to the Hyrcanian biogeographic province (Fig. 16).

Syrioiulus armeniacus sp. nov.

Figs 1F, 13, 14, 15F, 16

Material examined

Holotype ♂ (ZMUM), Armenia, Kafan Distr., Shikahoh Nature Reserve, Shikahoh, 900–950 m a.s.l., Quercus, Fagus, Carpinus forest by spring, litter, logs and under stones, 28.IV.1983, leg. S. Golovatch. Paratypes: 14 ♂♂, 24 ♀♀ (ZMUM), same collection data as holotype.

Non-type material

Armenia: 4 ♂♂, 3 ♀♀ (ZMUM), Shikahoh Nature Reserve, Nerkin And, old Platanus stand along river, litter, in rotten wood, under stones, 30.IV.1983; 1 ♂, 1 ♀ (ZMUM), near Kajaran, Megri Mt. Ridge, N of Tashtun Pass, 2000 m a.s.l., Quercus forest on steep slope, litter, logs, 27.IV.1983; 5 ♂♂, 3 ♀♀, 2 juv. (ZMUM), Megri Distr., SSE of Lichk, Megri River valley, Quercus forest, litter, under stones and in rotten wood, 25.IV.1983; 2 ♂♂, 16 ♀♀, 2 juv. (ZMUM), above Kuris, 1500 m a.s.l., Quercus and Carpinus forest, litter, under bark and stones along spring, 26.IV.1983; 12 ♂♂, 11 ♀♀, 1 juv. (ZMUM), ca. 4 km NNW of Megri, Legvaz, Juglans and Quercus shrub with Paliurus and Rosa, litter and under stones, 1000 m a.s.l., 24–25.IV.1983; 5 ♂♂, 2 ♀♀ (ZMUM), 6 km N of Shvanidzor, sparse Quercus forest, 1200–1300 m a.s.l., litter, under stones and bark, 24.IV.1983; 5 ♂♂, 6 ♀♀, 2 juv. (ZMUM), environs of Megri, xeriphytous bare canyon, under stones, sparse Juniperus and Paliurus, ca. 1000 m a.s.l., 24.IV.1983, all leg. S. Golovatch; 2 ♀♀ (ZMUM), Odzun W of Alaverdi, 1500–1550 m a.s.l., Quercus, Fagus, Carpinus etc. forest, litter and under stones with ants, 23–24.V.1987, leg. S. Golovatch and K. Eskov; 1 ♀ (ZMUM), Nurkus, 7.VII.1985, leg. V.A. Zakharyan.

Diagnosis

This new species belongs to the genus Syrioiulus because of the presence of only two apices on the opisthomere. Differs from all regional congeners by the following combination of somatic and gonopodal characters. Head with frontal setae. Collum and metazonae of following body rings without setae. Eyes absent. Solenomere with a pointed process apically. Anterior process rounded on top.

Name

The new species is named after its terra typica; adjective.

Description

Holotype: length 25 mm, width 1.2 mm, number of body rings 50+2+T. Paratypes and non-type material: length 17–33 mm, width 1.2–1.6 mm, number of body rings, 50–68+1–2+T (♂♂); or length 20–29 mm, width 1.2–1.6 mm, number of body rings, 46–55+2–3+T (♀♀). Body subcylindrical, metazonae brownish yellow, prozonae brownish grey (Figs 1F, 13A–G). Head, collum and telson slightly lighter than body rings (Fig. 1F). Antennae, mouthparts, and legs yellow (Fig. 13A, C–E, G). Eyes absent. Metazonae with weakly developed, dense, and regular striations, 20–23 striae per quarter of metazonital surface, i.e., that between dorsal axial line and ozopore (Fig. 13D). Ozopores relatively large, situated between striae in touch with one of them, lying behind suture without touching it (Fig. 13H).

Figure 13. 

Syrioiulus armeniacus sp. nov., paratype ♂ from Shikahoh, Armenia (ZMUM) A–C anterior part of body, lateral, dorsal and ventral views, respectively D midbody part, lateral view E–G posterior part of body, lateral, dorsal and ventral views, respectively H ozopores on midbody rings, lateral view I gnathochilarium, ventral view J leg pair 1, caudal view K leg 2, caudal view L ventral edge of pleurotergum 7, lateral view. Scale bars: 0.5 mm (A–G); 0.1 mm (H–L).

Antennae relatively long, in situ reaching segment 3. Head with 1+1 frontal, 9+9–10+10 labral and 2+2 supralabral setae (Fig. 13A–C). Gnathochilarium with three thick setae on each lamella lingualis; stipites with a group of 6–9 setae in medial part and three long setae in anterolateral part (Fig. 13I). Collum and metazonae without setae (Fig. 13A, B). Epiproct poorly developed, triangular, with several long setae (Fig. 13E, F). Hypoproct subtriangular, covered with long setae (Fig. 13G). Telson and anal valves densely setose.

Male. Mandibular stipites modified, slightly swollen in distal part (Fig. 13A). Leg pair 1 small, unciform, telopodites directed anteromesad (typical of Julidae), with a group of setae on coxa and telopodite (Fig. 13J). Leg pair 2 with small pads on postfemur and tibia (Fig. 13K). Penes short, bifurcate. Ventral edge of male pleurotergum 7 with wide curved lamellae bordering the gonopodal aperture (Fig. 13L).

Gonopods (Fig. 14) with anterior and posterior parts both equal in height. Promere spoon-shaped, bowl being relatively wide, constricted in basal third; mesal ridge well-developed all along, with a small mesal denticle in apical part; lateral denticle large, rounded on top (Fig. 14B, F). Mesomeral process simple, flattened, ribbon-shaped, widened apically, with a group of small teeth (Fig. 14A, C, E, G). Opisthomere bipartite. Solenomere long, slightly curved; its apical part with a fovea and a pointed process (Fig. 14A, C, D, G). Anterior process as high as solenomere, rounded at tip (Fig. 14D).

Figure 14. 

Syrioiulus armeniacus sp. nov., paratype ♂ from Shikahoh, Armenia (ZMUM) A, E gonopod, mesocaudal and mesal views, respectively B, E promere, caudal views C, G posterior gonopod, mesal views D end of solenomere, mesal view. Abbreviations: ap anterior process cl caudomesal lamella fo fovea ld lateral denticle md mesal denticle mr mesal ridge ms mesomeral process op opisthomere pr promere sl solenomere. Scale bars: 0.1 mm (A–C); 0.01 mm (D); 0.2 mm (E–G).

Female. First two leg pairs unmodified. Vulva rounded, operculum higher than bursa (Fig. 15F). Operculum slightly divided at apical margin. Bursa asymmetric, lateral valve higher than mesal one. Each valve with two rows of long setae. Median field of bursa narrow; emargination of median field elongated and suboval.

Figure 15. 

Vulvae of Amblyiulus and Syrioiulus species from the Caucasus, caudal views A A. georgicus Lohmander, 1932 from Shnokh, Armenia (ZMUM) B A. hirtus sp. nov., paratype from Bash-Layski, Azerbaijan (ZMUM) C S. adsharicus (Lohmander, 1936) from Adigeni, Georgia (ZMUM) D S. continentalis (Attems, 1903) from Istisu, Azerbaijan (ZMUM) E S. taliscius (Attems, 1927) from Avrora, Azerbaijan (ZMUM) F S. armeniacus sp. nov., paratype from Shikahoh, Armenia (ZMUM). Abbreviations: em emargination of median field lv lateral valve mf median field mv median valve op operculum. Scale bars: 0.03 mm.

Remark

This species seems to be endemic to the Caucasus Minor within Armenia, but most likely it also occurs in the adjacent parts of eastern Azerbaijan and northwestern Iran (Fig. 16).

Figure 16. 

Distributions of Amblyiulus and Syrioiulus species in the Caucasus: red circle A. georgicus Lohmander, 1932; green circle A. hirtus sp. nov.; white circle S. adsharicus (Lohmander, 1936); blue circle S. continentalis (Attems, 1903); pink circle S. taliscius (Attems, 1927); brown circle S. armeniacus sp. nov.; yellow square Pachyiulini gen. sp. 1; purple square Pachyiulini gen. sp. 2.

Pachyiulini gen. sp. 1

Fig. 16

Material examined

1 ♀ (ZMUM) Russia, Stavropol Area, Kochubeevsky Distr., Nevinnomyssk, mouth of Bolshoy Zelenchuk River, floodplain forest, 17.IV.2016, leg. R.B. Zuev, A.V. Aulova.

Brief description

Body grey, head and collum dark yellow, antennae and legs yellow. The following somatic characteristics seem to be the most important: absence of eyes, presence of frontal setae, presence of caudal whorls of setae on metazonae, absence of an epiproct, and setose anal valves.

Remark

Unfortunately, the only female does not allow a closer generic allocation.

Pachyiulini gen. sp. 2

Fig. 16

Material examined

Georgia: 1 ♀ (ZMUM), Ajaria, Kintrish Nature Reserve, Zeraboseli, 450–600 m a.s.l., 1–3.VI.1981, leg. S. Golovatch, J. Martens; 1 ♀ (ZMUM), same locality, 800 m a.s.l., Rhododendron thicket, litter, 13.X.1981, leg. S. Golovatch; 1 ♀ (ZMUM), same locality, valley of Khekpara River, 12.X.1984, leg. E. Kvavadze.

Brief description

Body greyish yellow. Head, collum, a few postcollum rings and telson slightly lighter than other body rings. Ommatidia and frontal setae absent. Collum and each metazona of following rings with a whorl of long setae at posterior margin. Epiproct undeveloped. Anal valves densely setose.

Remarks

These specimens differ from Syrioiulus adsharicus in the absence of frontal setae and ommatidia. The absence of males makes it impossible to definitively identify the above samples. At least the taxonomic significance of frontal setae must not be overestimated, as they may be present or absent even within the same species, e.g., S. aharonii (see Golovatch 2018).

Since the two unidentified species may well prove to represent Amblyiulus or Syrioiulus, we map their records in Figure 16, but omit them from the key below.

Key to Amblyiulus and Syrioiulus species known to occur in the Caucasus (based on males)

1 Opisthomere of gonopod with a rod (Fig. 3D, F, G, I) (genus Amblyiulus) 2
Opisthomere of gonopod without rod (genus Syrioiulus) 3
2 Head without frontal setae, collum and metazonae of body rings without setae (Fig. 2A–C). Promere relatively wide, with two apical denticles (Fig. 3B, E) A. georgicus
Head with 1+1 frontal setae, collum and metazonae of body rings with whorls of setae at caudal margin (Fig. 4A–C). Promere narrow, without apical denticles (Fig. 5B, H) A. hirtus sp. nov.
3 Eyes present 4
Eyes absent 5
4 Larger: width > 2.0 mm. Eyes well-developed, black, oval, each composed of 19–23 ommatidia (Fig. 9A) S. continentalis
Smaller: width < 1.3 mm. Eyes present, but very small and unpigmented, each composed of 3–7 ommatidia (Fig. 6B) S. adsharicus
5 Head without frontal setae (Fig. 11A–C) S. taliscius
Head with 1+1 frontal setae (Fig. 13A–C) S. armeniacus sp. nov.

Discussion

Two allopatric species of Amblyiulus, both likely endemic, are found to populate the Caucasus. Amblyiulus georgicus inhabits western and central Transcaucasia, while A. hirtus sp. nov. seems to be confined to northeastern Transcaucasia and the eastern Caucasus, i.e., occurring on both macro slopes of the Caucasus Major (Fig. 16).

The genus Syrioiulus is more diverse and widespread, but presently it seems to be restricted to Transcaucasia. Thus, S. adsharicus has a rather narrow distribution in the southwestern parts of the Colchidan biogeographic province. Two most widespread species, S. continentalis and S. taliscius, are often sympatric to even syntopic within the Hyrcanian biogeographic province, but the latter species also occurs in the Caucasus Minor and the eastern part of the Caucasus Major. Syrioiulus armeniacus sp. nov. inhabits the Caucasus Minor within southern and central Transcaucasia (Fig. 16).

As regards the vertical distributions, most species of Amblyiulus and Syrioiulus in the Caucasus are confined to montane forests, as are probably all Syrioiulus species known from Hyrcania, including the Iranian S. astrabadensis (Lohmander, 1932b), S. discolor (Lohmander, 1932b), S. incarnatus (Lohmander, 1932b), S. lohmanderi Vagalinski, 2020, S. persicus (Golovatch, 1983) and S. zarudnyi (Lohmander, 1932b) (Enghoff and Moravvej 2005; Vagalinski 2020). Within the Republic of Azerbaijan, however, S. taliscius inhabits not only lowland to foothill woodlands (50–1100 m a.s.l.), but also drier steppe- or bush-clad slopes, being inclined to synanthropisation as well (parks in Baku City). The same generally applies to both S. continentalis and S. armeniacus sp. nov., as they also populate xerophytic environments. Amblyiulus hirtus sp. nov. has been encountered in mid-montane deciduous forests, as well as subalpine and alpine meadows up to 2550 m a.s.l.

Only two species of Syrioiulus, S. continentalis and S. taliscius, are endemic or subendemic, respectively, to the Hyrcanian biogeographic province within the Republic of Azerbaijan and Iran, while the remaining Amblyiulus and Syrioiulus spp., however provisionally, are formally strictly endemic to the Caucasus sensu lato, including Hyrcania (Fig. 16). Generally, the problem concerning the origins of the Amblyiulus and Syrioiulus species could be approached through analysing the distribution areas of both these genera. However, because their species compositions are far from settled, including new congeners and records certainly ahead, their exact ranges cannot be outlined at the moment. Based on the highest species diversity estimates, one of the main centres of Amblyiulus and Syrioiulus speciation could have lain in the Middle East, whence members of both genera might have reached the Caucasus sensu lato. The roles that both Colchis and, especially, Hyrcania, two major, relictual, meso- to hygrophilous biogeographic provinces of, and refugia in, the region concerned, could have played in the evolution and secondary speciation seem paramount. It is hardly random that most pachyiuline species in the Caucasus are encountered in the Caucasus Minor, the immediate northern peninsular continuation of Asia Minor.

Both unidentified species seem to be endemic to the Caucasus, with Pachyiulini gen. sp. 1 being confined to Ciscaucasia, and Pachyiulini gen. sp. 2 to deciduous forests in southern Colchis, occurring sympatrically with Syrioiulus adsharicus (Fig. 16).

The above picture is definitely very far from final, but it agrees well with the biogeography of the Caucasus (e.g., Abdurakhmanov 2017). Given that the Pachyiulini in the faunas of Turkey (Enghoff 2006) and Iran (Vagalinski 2020) are likewise quite poorly known, each amounting to only a handful of species, there can hardly be any doubt that future progress in the study of pachyiulines in the Caucasus region and adjacent countries will reveal numerous novelties. In addition, given the presence in Crimea of a troglobitic species, Syrioiulus kovali (Golovatch, 2008), finding cave species of Pachyiulini in the Caucasus could also be expected (Golovatch 2008; Golovatch et al. 2017; Turbanov et al. 2018). The present taxonomy and distributions as outlined above must be clarified and refined through future research, both morphology- and molecular-based.

Acknowledgements

We are most grateful to all collectors who rendered us their material for treatment. We are also greatly obliged to both Igor Zabiyaka (Don StateTechnical University, Rostov-on-Don, Russia) for taking SEM pictures. DA is grateful to Hans Reip and Frank Walther (both Germany) for their help during the field trip in Azerbaijan in March 2015, as well as again to Hans Reip and Karin Voigtländer (Germany) for their help in the preparation of some samples used here for SEM. AE was supported by RFBR, Project No. 20-54-18008, SG by the Presidium of the Russian Academy of Sciences, Programme No. 41 “Biodiversity of Natural Systems and Biological Resources of Russia”, while DA by the project “Biogeography of the land molluscs of the Caucasus region” funded by VolkswagenStiftung, and the Serbian Ministry of Education, Science, and Technology (Grant 173038) for making his field trip to Azerbaijan possible, as well as by the SYNTHESYS Project (DE-TAF-5619) financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program which allowed him short visits to the Senckenberg Museum of Natural History in Görlitz (Germany) in 2016. We are most thankful to Henrik Enghoff (Copenhagen, Denmark) and Boyan Vagalinski (Sofia, Bulgaria), the reviewers who provided constructive criticism and thus considerably improved our paper. Pavel Stoev (Sofia, Bulgaria) very helpfully served as the managing editor.

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