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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by A. Henrard: 4 Aug. 2022; published: 25 Aug. 2022 161
Zootaxa 5178 (2): 161–177
https://www.mapress.com/zt/
Copyright © 2022 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5178.2.3
http://zoobank.org/urn:lsid:zoobank.org:pub:B466BB45-91F1-456E-971A-E3FAA38CBCA9
New taxonomic considerations in Zodariellum Andreeva & Tyshchenko, 1968
(Araneae: Zodariidae), with notes on the presence of cymbial diverticulum in
different zodariid genera
ALIREZA ZAMANI1,4* & YURI M. MARUSIK2,3,5
1Zoological Museum, Biodiversity Unit, University of Turku, FI-20014, Finland.
2Institute for Biological Problems of the North RAS, Portovaya Str.18, Magadan, Russia.
3Department of Zoology & Entomology, University of the Free State, Bloemfontein 9300, South Africa.
�
yurmar@mail.ru; https://orcid.org/0000-0002-4499-5148
*Corresponding author.
�
zamani.alireza5@gmail.com; https://orcid.org/0000-0002-8084-9666
Abstract
New taxonomic considerations in the zodariid spider genus Zodariellum Andreeva & Tyshchenko, 1968 are proposed.
On the basis of male specimens collected in Turkmenistan, Z. turanicum sp. n. and Z. turkmenicum sp. n. are described
as new to science. Six species, namely Z. hunanense (Yin, 2012) comb. n., Z. planum (Zhang & Zhang, 2019) comb. n.,
Z. spasskyi (Charitonov, 1946) comb. n., Z. tadzhikum (Andreeva & Tyshchenko, 1968) comb. n., Z. testaceofasciatum
(Spassky, 1941) comb. n. and Z. zebra (Charitonov, 1946) comb. n., are newly transferred from Zodarion Walckenaer,
1826 to Zodariellum. Two new synonymies are proposed: Zodarion apertum Zhang & Zhang, 2019 syn. n. is synonymized
with Zodariellum mongolicum Marusik & Koponen, 2001, and Zodarion martynovae Andreeva & Tyshchenko, 1968 syn.
n. is synonymized with Z. continentale (Andreeva & Tyshchenko, 1968). Additionally, illustrations are provided for three
poorly known Central Asian species, namely Z. asiaticum (Tyshchenko, 1970), Z. bactrianum (Kroneberg, 1875) and
Z. nenilini (Eskov, 1995). A deep and conical diverticulum of unknown function, located on the retrodorsal side of the
cymbium is documented in Zodariellum for the first time; the presence of similar structures in different zodariid genera
is discussed.
Key words: Aranei, Central Asia, new combination, new species, new synonymy
Introduction
Zodariellum Andreeva & Tyshchenko, 1968 is a genus of the spider family Zodariidae Thorell, 1881, currently
comprising 13 species distributed from the Volga River’s delta to northeastern China, with their highest diversity in
the Central Asia (WSC 2022). The genus, described as monotypic, has had a somewhat confusing taxonomic history
starting with several inconsistencies in the paper (Andreeva & Tyshchenko 1968) in which the name was designated
(e.g. change of authorship through the text, using “Lodariellum” in the heading instead of Zodariellum). Jocqué
(1991) considered Zodariellum to be a junior synonym of Acanthinozodium Denis, 1966. Marusik & Koponen
(2001) refuted this and considered Zodariellum be the valid name, as no type species was designated by Denis for
Acanthinozodium. Marusik & Koponen (2001) also provided the descriptions of two new species and proposed
eight new combinations, which when added to the two species later described by Ponomarev (2007), raised the
total number of known Zodariellum species to 13. Jocqué & Henrard (2015) considered both Acanthinozodium and
Zodariellum as valid, with Zodariellum comprising only the type species, Z. surprisum Andreeva & Tyshchenko,
1968, and transferred the remaining species to Zodarion Walckenaer, 1826. Finally, Zamani & Marusik (2021)
redefined the genus, proposed one new combination, and except for Zodarion inderense (Ponomarev, 2007),
transferred all the other species back into Zodariellum. Of the currently 13 nominal species considered in the genus,
three are known only by males and three (including the type) have no proper figures illustrating the male palp.
Recently, we had the opportunity to examine some material of this genus collected in the Central Asia, including
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162 · Zootaxa 5178 (2) © 2022 Magnolia Press
two new species from Turkmenistan which are described in this paper. Furthermore, six new combinations and
two new synonymies are proposed, raising the total number of species known in the genus to 21. Finally, a deep
and conical diverticulum of unknown function, located on the retrodorsal side of the male palp’s basal area is
documented in Zodariellum for the first time; the presence of similar structures in other zodariids is discussed.
Material and methods
Specimens were photographed using a Canon EOS 7D camera attached to an Olympus SZX16 stereomicroscope,
and a JEOL JSM-5200 scanning electron microscope at the Zoological Museum of the University of Turku. Digital
images were montaged using CombineZP and edited using Corel Draw graphic design software. Measurements of
legs are listed as: total length (femur, patella, tibia, metatarsus, tarsus). All measurements are given in millimeters.
Abbreviations used in text: Eyes: ALE—anterior lateral eye, AME—anterior median eye, PLE—posterior
lateral eye, PME—posterior median eye. Legs: Fe—femur, Pa—patella, Ti—tibia, Mt—metatarsus. Male palp:
RTA—retrolateral tibial apophysis. Abbreviations used in the figure plates are given in the legends and within the
descriptions and diagnoses.
Depositories: ISEA—Institute of Systematics and Ecology of Animals, Novosibirsk, Russia (Galina N.
Azarkina); ZMMU—Zoological Museum of Moscow State University, Moscow, Russia (Kirill G. Mikhailov).
Family Zodariidae Thorell, 1881
Subfamily Zodariinae Thorell, 1881
Genus Zodariellum Andreeva & Tyshchenko, 1968
Zodariellum Andreeva & Tystshenko, 1968: 688.
Zodariellum: Marusik & Koponen 2001: 40 (removed from the synonymy of Zodarion).—Zamani & Marusik 2021: 184.
Type species. Zodariellum surprisum Andreeva & Tyshchenko, 1968 from Tajikistan, by original designation.
Emended diagnosis. Males of Zodariellum differ from that of Zodarion nitidum (Audouin, 1826) (Fig. 11A–
C), the generotype of Zodarion, by having: 1) RTA (Ra) longer than wide, with solid base and either bifurcated or
solid tip, with branch or branches (Rb) bent retrolaterally at more than right angle (Figs 4A–D, 5A–D) (vs. wider
than long and divided into 3 parts basally; Fig. 11B), 2) lacking ventral tibial apophysis (vs. present, Va in Fig.
11A–B), 3) cymbium with short tip (i.e. as long as wide, shorter than RTA) and prominent tutaculum (Tu) (vs. tip
longer than wide and longer than RTA, tutaculum absent), 4) large tegular apophysis (Tg) longer that bulb, with
prolateral spur (Sp) located basally and directed postero-ectally, and long retrolateral arm (Tr) with tip resting on
tutaculum (vs. tegular apophysis shorter than bulb), 5) very long conductor (Cn), longer than bulb, with prolateral
arm (Cp) terminating at a 6 o’clock position or earlier (vs. conductor lacking prolateral arm and shorter than bulb),
and 6) longer filamentous embolus originating at a 6 o’clock position or earlier (vs. 6:30). Females of Zodariellum
differ from those of Zodarion sensu lato by the long, twisted (i.e. similar to a screw), converging copulatory ducts
(Fig. 10A–D) (vs. not twisted or diverging if twisted).
Furthermore, males of Zodariellum differ from those of all other zodariid genera by having a very deep (=long)
diverticulum of cymbium (Di, Fig. 2F) (about 0.4 time of cymbium length and 0.75 of cymbium width). The species
of Acanthinozodium also have a cymbial diverticulum, although in the form of a short conical (i.e. depth about one
radius) round pit (Jocqué & Henrard 2015: fig. 9A, B; Zamani & Marusik, 2021: figs 13C, 14G‒H, 15C, F, 21F,
22H‒I, 23C, 24C). Furthermore, the male palps of Zodariellum species have serrated processes of RTA and serrated
spur of the tegular apophysis (Sp), which have not been recorded in other Zodariinae genera.
Composition. 21 species (including those described here and the new combinations):
Z. asiaticum (Tyshchenko, 1970) (♂♀), Z. bactrianum (Kroneberg, 1875) (♂♀), Z. bekuzini (Nenilin, 1985)
(♂♀), Z. chaoyangense (Zhu & Zhu, 1983) (♂♀), Z. continentale (Andreeva & Tyshchenko, 1968) (♂♀), Z. furcum
(Zhu, 1988) (♂♀), Z. hunanense (Yin, 2012) comb. n. (♀), Z. mongolicum Marusik & Koponen, 2001 (♂♀), Z.
nenilini (Eskov, 1995) (♂♀), Z. planum (Zhang & Zhang, 2019) comb. n. (♂), Z. proszynskii (Nenilin & Fet, 1985)
(♂♀), Z. schmidti Marusik & Koponen, 2001 (♂♀), Z. spasskyi (Charitonov, 1946) comb. n. (♀), Z. surprisum
NEW TAXONOMIC CONSIDERATIONS IN ZODARIELLUM Zootaxa 5178 (2) © 2022 Magnolia Press · 163
(Andreeva & Tyshchenko, 1968) (♂), Z. sytchevskajae (Nenilin & Fet, 1985) (♂♀), Z. tadzhikum (Andreeva &
Tyshchenko, 1968) comb. n. (♀), Z. testaceofasciatum (Spassky, 1941) comb. n. (♀), Z. turanicum sp. n. (♂), Z.
turkmenicum sp. n. (♂), Z. volgouralense Ponomarev, 2007 (♂♀) and Z. zebra (Charitonov, 1946) comb. n. (♀).
According to the shape of the RTA, the genus can be split into three species-groups:
1) furcum species-group, comprising Z. furcum and Z. planum; in both species, RTA has a process arising from
its mid part, and a pointed tip (Zhang & Zhang 2019: figs 4E, 12C).
2) bactrianum species-group, comprising Z. bactrianum and Z. turanicum sp. n.; in both species, RTA has two
processes originating posteriorly, and an abrupt tip (Figs 2B, D).
3) surprisum species-group, comprising the rest of the species with known males; in all species of this group,
RTA has one or two processes, and a roundly or almost roundly bent tip (Figs 2A, C, E).
Distribution. From Volga River’s delta to northeastern China, south to northeastern Iran and south China (ca
25ºN) (Fig. 12).
FIGURE 1. Habitus of Zodariellum asiaticum (A), Z. bactrianum (B), Z. turanicum sp. n. (C), Z. nenilini (D–E) and Z.
turkmenicum sp. n. (F), dorsal view. A–D, F males; E female. Scale bars = 0.5 mm.
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Zodariellum asiaticum (Tyshchenko, 1970)
Figs 1A, 2A, 3A, 4A, 5A, 6A, 9A, 12A
Zodarion asiaticum Tyshchenko, in Marikovski & Tyshchenko 1970: 199, fig. 1 (♂♀).
Material examined. KAZAKHSTAN: Jambyl Region: 1♂ (ZMMU), Korday Dist., 10 km SE of Korday Vil.,
Yeradzhy well, [43°02'N 74°42'E], 20.04.1997 (A.V. Gromov).
FIGURE 2. Male palps of Zodariellum asiaticum (A), Z. bactrianum (B, F), Z. nenilini (C), Z. turanicum sp. n. (D) and Z.
turkmenicum sp. n. (E). A–E ventral view; F dorsal view of cymbium, showing cymbial diverticulum. Scale bars = 0.2 mm.
Abbreviations: Cn—conductor, Cp—prolateral arm of conductor, Eb—base of embolus, Di—diverticulum, Ra—retrolateral tibial
apophysis, Rb—branch(es) of retrolateral tibial apophysis, Sp—spur of tegular apophysis, Tg—tegular apophysis, To—triangular
outgrowth, Tr—retrolateral arm of tegular apophysis, Tu—tutaculum.
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FIGURE 3. SEM images of the male palps of Zodariellum asiaticum (A), Z. bactrianum (B), Z. nenilini (C), Z. turanicum sp.
n. (D) and Z. turkmenicum sp. n. (E), ventral view. Scale bars = 0.1 mm. Abbreviations: Cn—conductor, Cp—prolateral arm of
conductor, Eb—base of embolus, Ra—retrolateral tibial apophysis, Rb—branches of retrolateral tibial apophysis, Sp—spur of
tegular apophysis, Tg—tegular apophysis, Tr—retrolateral arm of tegular apophysis, Tu—tutaculum.
Description. See Marikovski & Tyshchenko (1970). Habitus of male as in Fig. 1A. Male palp as in Figs 2A,
3A, 4A, 5A, 6A, 9A.
Comments. Previously illustrated only in the original description. The type material was collected from the
nest of Polyergus rufescens (Latreille, 1798) (Hymenoptera: Formicidae). Marikovski & Tyshchenko (1970) and
Pekár (2009) described its hunting and courtship behaviours.
Distribution. Kazakhstan (type locality; Fig. 12A), Kyrgyzstan (Zonstein et al. 1996) and Tajikistan (Andreeva
1976). The record from Tajikistan is based on a singleton female that is most likely misidentified, and therefore not
included in the map.
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FIGURE 4. Male palps of Zodariellum asiaticum (A), Z. bactrianum (B), Z. nenilini (C), Z. turanicum sp. n. (D) and Z.
turkmenicum sp. n. (E), retrolateral view. Scale bars = 0.2 mm. Abbreviations: Ap—anterodorsal part of retrolateral tibial
apophysis, Cn—conductor, Di—diverticulum, Ra—retrolateral tibial apophysis, Rb—branches of retrolateral tibial apophysis,
Tr—retrolateral arm of tegular apophysis, Tu—tutaculum.
NEW TAXONOMIC CONSIDERATIONS IN ZODARIELLUM Zootaxa 5178 (2) © 2022 Magnolia Press · 167
FIGURE 5. SEM images of the male palps of Zodariellum asiaticum (A), Z. bactrianum (B), Z. nenilini (C), Z. turanicum
sp. n. (D) and Z. turkmenicum sp. n. (E), retrolateral view. Scale bars = 0.1 mm. Abbreviations: Cn—conductor, Cp—prolateral
arm of conductor, Di—diverticulum, Eb—base of embolus, Ra—retrolateral tibial apophysis, Rb—branches of retrolateral tibial
apophysis, Tr—retrolateral arm of tegular apophysis, Tu—tutaculum.
Zodariellum bactrianum (Kroneberg, 1875)
Figs 1B, 2B, 2F, 3B, 4B, 5B, 6B, 7A–C, 8A–C, 9B–C, 12A
Zodarion bactrianum Kroneberg, 1875: 12, plate 1, fig. 5 (♂♀).
Zodarion bactrianum: Andreeva & Tyshchenko 1968: 687, fig. 4 (♂).—Andreeva 1976: 35, fig. 42 (♂; figure from Andreeva
& Tyshchenko 1968).
Zodariellum bactrianum: Zamani & Marusik 2021: 185, fig. 28D–F (♂).
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Material examined. TAJIKISTAN: Khatlon Region: 1♂ (ZMMU), Shaartuz Dist., Babatag Mt. ridge, 37°04'32.9"N
68°01'10.6"E, 427 m, cay cliffs, 20.04.2015 (Y.M. Marusik); Sughd Region: 1♂ (ZMMU), Buston, ca. 40°31'N
69°20'E, 18.05.2002 (S.L. Zonstein).
Description. See Andreeva & Tyshchenko (1968). Habitus of male as in Fig. 1B. Male palp as in Figs 2B, 2F,
3B, 4B, 5B, 6B, 7A–C, 8A–C, 9B–C.
Distribution. This species was described from eastern Uzbekistan, and later reported from Tajikistan (Spassky
& Luppova 1945; Andreeva & Tyshchenko 1968; Andreeva 1976; Zamani & Marusik 2021) (Fig. 12A). According
to Mikhailov (2021), this species also occurs in Kazakhstan and Kyrgyzstan.
FIGURE 6. Male palps of Zodariellum asiaticum (A), Z. bactrianum (B), Z. nenilini (C), Z. turanicum sp. n. (D) and Z.
turkmenicum sp. n. (E), dorsal view. Scale bars = 0.2 mm.
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FIGURE 7. SEM images of the retrolateral tibial apophyses of Zodariellum bactrianum (A–C) and Z. nenilini (D–F). Scale bars = 0.1 mm.
Zodariellum continentale (Andreeva & Tyshchenko, 1968)
Fig. 12A
Zodarion continentalis Andreeva & Tyshchenko, 1968: 687, fig. 5 (♂).
Zodarion martynovae Andreeva & Tyshchenko, 1968: 684, figs 1–2 (♀). Syn. n.
Zodarion martynovae: Andreeva 1976: 37, figs 45–46 (♀).
Zodarion continentalis: Andreeva 1976: 35, fig. 43 (♂).
Zodarion continentale: Brignoli 1983: 212.—Jocqué & Henrard 2015: 21.
Zodariellum continentale: Marusik & Koponen 2001: 41.
Zodariellum continentalis: Zamani & Marusik 2021: 185.
Comments. The proposed synonymy is due to the fact that the type materials of Z. martynovae (described on the basis of
female) and Z. continentale (described on the basis of male) were collected in the same locality. The holotype female of Z.
martynovae is in the Institute of Zoology of the Polsih Academy of Sciences (W. Wawer, pers. comm.).
Distribution. Uzbekistan, Tajikistan (Fig. 12A).
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FIGURE 8. SEM images of the cymbia of Zodariellum bactrianum (A–C) and Z. nenilini (D–F). Scale bars = 0.1 mm, unless
stated otherwise. Abbreviations: Cp—prolateral arm of conductor, Di—diverticulum, Tr—retrolateral arm of tegular apophysis,
Tu—tutaculum.
Zodariellum hunanense (Yin, 2012) comb. n.
Fig. 12B
Zodarion hunanensis Yin, in Yin et al. 2012: 1148, figs 611a–e (♀).
Comments. The southernmost species of the genus, currently known only from the original description based on
material from southern China (Fig. 12B). Here, it is transferred to Zodariellum due to its screw-like copulatory
ducts.
Distribution. Southern China (Hunan) (Fig. 12B).
Zodariellum mongolicum Marusik & Koponen, 2001
Fig. 12B
Zodariellum mongolicum Marusik & Koponen, 2001: 45, figs 3–5, 8–9, 17–18, 26 (♂).
Zodarion mongolicum: Jocqué & Henrard 2015: 21.
Zodariellum mongolicum: Zamani & Marusik 2021: 185.
Zodarion apertum Zhang & Zhang, 2019: 69, figs 1A–D, 2A–E, 3A–E (♂♀). Syn. n.
NEW TAXONOMIC CONSIDERATIONS IN ZODARIELLUM Zootaxa 5178 (2) © 2022 Magnolia Press · 171
Comments. Accounting the original descriptions, we hereby synonymize Z. apertum Zhang & Zhang, 2019 syn.
n. (described from Xinjiang, northwestern China) with Z. mongolicum (described from Mongolia), as no noticeable
differences can be detected between the illustrated male palps (cf. Zhang & Zhang 2019: fig. 2C and Marusik &
Koponen 2001: figs 17–18).
Distribution. Mongolia and northwestern China (Xinjiang) (Fig. 12B).
FIGURE 9. SEM images of the male palps of Zodariellum asiaticum (A), Z. bactrianum (B–C) and Z. turanicum sp. n. (D). A
anteroretrolateral view; B ventroretrolateral view; C anteroventral view; D anterior view. Scale bars = 0.1 mm. Abbreviations:
Cn—conductor, Eb—base of embolus, Ra—retrolateral tibial apophysis, Rb—branches of retrolateral tibial apophysis, Tr—
retrolateral arm of tegular apophysis, Tu—tutaculum.
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FIGURE 10. Epigynes of Zodariellum nenilini (A) and Z. proszynskii (B–D). A–B intact, ventral view; C–D macerated, dorsal
and ventral views. Scale bars = 0.2 mm. Abbreviations: Cd—copulatory duct, Fo—fovea, Re—receptacle.
FIGURE 11. SEM images of the male palp of Zodarion nitidum. A ventral view; B retrolateral view; C dorsal view. Scale
bars = 0.1 mm. Abbreviations: Cn—conductor, Eb—base of embolus, Ra—retrolateral tibial apophysis, Tg—tegular apophysis,
Va—ventral tibial apophysis.
Zodariellum nenilini (Eskov, 1995)
Figs 1D–E, 2C, 3C, 4C, 5C, 6C, 7D–F, 8D–F, 10A, 12A–B
Zodarion nenilini Eskov, in Eskov & Marusik 1995: 62, figs 27–29 (♂♀).
Zodariellum nenilini: Marusik & Koponen 2001: 42, figs 1, 6, 10–13, 19–24 (♂♀).—Esyunin et al. 2007: 57, figs 21–23
(♂).—Marusik & Kovblyuk 2011: 277, figs 44.2. 4–5 (♂♀).
Material examined. KAZAKHSTAN: East Kazakhstan Region: 2♂ (ISEA), W outreaches of Narym Mt. Range,
Kaindy River Valley, 10–15 km SE of Slavyanka, [43°N 78°E], steppes, 6–7.05.1999 (R. Dudko, I. Lyubechanski);
NEW TAXONOMIC CONSIDERATIONS IN ZODARIELLUM Zootaxa 5178 (2) © 2022 Magnolia Press · 173
1♀ (ISEA), northern Cis-Zaisan (=N cosat of Zaisan Lake), N part of Kuludzhun sands, [48°40'N 83°20'E], 8–
10.05.1999 (R. Dudko, I. Lyubechanski).
Description. See Eskov & Marusik (1995) and Marusik & Koponen (2001). Habitus of male and female as in
Fig. 1D, E. Male palp as in Figs 2C, 3C, 4C, 5C, 6C, 7D–F, 8D–F. Epigyne as in Fig. 10A: fovea (Fo) wider than
long; copulatory ducts (Cd) contiguous and forming angle >90º.
Distribution. Known from South Urals (51°54'N, 55°20'E; Esyunin et al. 2007), east Kazakhstan (type locality)
and Gobi-Altai and Bayankhongor Aimags (ca 101ºE) of Mongolia (Marusik & Koponen 2001) (Fig. 12A–B). It is
possible that the records from South Ural belong to Z. volgouralense Ponomarev, 2007.
FIGURE 12. Distribution records of Zodariellum spp. A: 1) Z. asiaticum, 2) Z. bactrianum, 3) Z. bekuzini, 4) Z. continentale,
5) Z. proszynskii, 6) Z. spasskyi and Z. zebra, 7) Z. surprisum, 8) Z. sytchevskajae, 9) Z. turanicum sp. n., 10) Z. turkmenicum sp.
n., 11) Z. tadzhikum, 12) Z. testaceofasciatum, 13) Z. volgouralense, 14) Z. nenilini (in part, excluding records from Mongolia).
B: 1) Z. nenilini (in part, records from Mongolia), 2) Z. schmidti, 3) Z. mongolicum, 4) Z. chaoyangense, 5) Z. furcum, 6) Z.
hunanense, 7) Z. planum. A single point may refer to several closely located records.
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Zodariellum planum (Zhang & Zhang, 2019) comb. n.
Fig. 12B
Zodarion planum Zhang & Zhang, 2019: 73, figs 4A–E, 5A–C (♂).
Comments. Currently known only from the original description, on the basis of material from central China
(Fig. 12B). Here, it is transferred to Zodariellum due to the conformation of the male palp with large RTA, long
filamentous embolus and long conductor, as well as the presence of a tutalculum and cymbial diverticulum (Zhang
& Zhang 2019: fig. 4E).
Distribution. Central China (Shaanxi) (Fig. 12B).
Zodariellum spasskyi (Charitonov, 1946) comb. n.
Fig. 12A
Zodarion spasskyi Charitonov, 1946: 22, fig. 11 (♀).
Zodarion spasskyi: Charitonov 1969: 72.
Comments. This species is transferred to Zodariellum due to its screw-like copulatory ducts, a character unknown
in other Zodariinae.
Distribution. According to WSC (2022), this species is known from Uzbekistan and Kazakhstan. Accounting
that the only figure provided with the original description is not very informative, it is very much possible that the
record from Kazakhstan belongs to another species. We have only marked the type locality on the map (Fig. 12A).
Zodariellum tadzhikum (Andreeva & Tyshchenko, 1968) comb. n.
Fig. 12A
Zodarion tadzhikum Andreeva & Tyshchenko, 1968: 686, fig. 3 (♀).
Zodarion tadzhikum: Andreeva 1976: 39, fig. 47 (♀).
Comments. There are no proper figures of the epigyne available for this species. We have transferred it to Zodariellum
considering that Andreeva (1976) compared it with Z. bactrianum and Z. zebra. The holotype is in the Zoological
Institute of the Polish Academy of Sciences (W. Wawer, pers. comm.).
Distribution. Tajikistan (Fig. 12A).
Zodariellum testaceofasciatum (Spassky, 1941) comb. n.
Fig. 12A
Zodarion testaceo-fasciatum Spassky, 1941: 14, fig. 3 (♀).
Zodarion testaceo-fasciatum Spassky & Luppova, 1945: 45 (♀).
Zodarion testaceofasciatum: Brignoli 1983: 213.
Comments. This species was described twice, in two different publications and under the same name. Apparently,
the original manuscript was sent to the publisher in Riga prior to World War II, and Spassky was unaware that his
paper was published during the war (in 1941) and resubmitted the manuscript again, this time in coauthorship with
E.P. Luppova to the Soviet journal Entomologicheskoe Obozrenie. The second manuscript was published in 1945,
without any figures.
The species is known only from the type locality, and only one figure showing the abdominal pattern has been
published for it. It is very likely that Z. zebra, a species with similar abdominal pattern and type locality not far
from that of Z. testaceofasciatum, is a junior synonym of it. Although the epigyne of this species is not described,
we decided to transfer it to Zodariellum considering its similarity to Z. zebra and the fact that only Zodariellum and
“Zodarion” denisi Spassky, 1938 are known to occur in the region.
Distribution. Tajikistan (Fig. 12A).
NEW TAXONOMIC CONSIDERATIONS IN ZODARIELLUM Zootaxa 5178 (2) © 2022 Magnolia Press · 175
Zodariellum turanicum sp. n.
Figs 1C, 2D, 3D, 4D, 5D, 6D, 9D, 12A
Type material. Holotype ♂ (ZMMU), TURKMENISTAN: Lebap Region: Kugitang-Tau Mnt. range, environs of
Khodzhapil, [37°56'N 66°37'E], ca 1100 m, on the ground, 20.05.1985 (A.V. Tanasevitch).
Etymology. The specific epithet refers to Turan, a historical region in Central Asia.
Diagnosis. The new species has abdominal pattern similar to those of Z. zebra and Z. testaceofasciatum, two
species described not far from each other (eastern Uzbekistan and western Tajikistan) and currently known only
from females. It differs from the males of its congeners by having two transverse light stripes on the abdomen (vs.
lacking in other species), and a V-shaped dark marking on a light carapace (vs. carapace uniformly coloured or
with small longitudinal spots along the fovea). The male palp of Z. turanicum sp. n. is most similar to that of Z.
bactrianum by having two processes in the distal half of RTA, but can be differentiated by the angled anterodorsal
part of RTA (Ap) (vs. rounded), and branches located at the same level (vs. mesal one [=closer to cymbium] located
more posteriorly).
Description. Male. Habitus as in Fig. 1C. Total length 3.55. Carapace 1.60 long, 1.50 wide. Eye sizes: AME:
0.18, ALE: 0.10, PME: 0.10, PLE: 0.08. Carapace, sternum, chelicerae, labium and maxillae yellowish. Carapace
light coloured with irregular V-shaped marking. Legs slightly lighter than carapace. Abdomen dorsally dark maroon,
with 2 pale transverse stripes in anterior half and 3 obtuse triangular spots in anterior half; venter light beige.
Measurements of legs: I: 6.78 (1.82, 0.62, 1.59, 1.77, 0.98), II: 6.23 (1.67, 0.55, 1.37, 1.78, 0.86), III: 6.57 (1.77,
0.61, 1.38, 1.91, 0.90), IV: 8.22 (2.34, 0.63, 1.88, 2.44, 0.93).
Palp as in Figs 2D, 3D, 4D, 5D, 6D, 9D. Tibial apophysis longer than tibia, 2.3 times longer than wide in
proximal part, tip inclined, with anterodorsal part (Ap) angled; dorsal part with 2 processes originated at anterior 1/3,
processes of subequal length and shape and located at the same level (one behind the other), ectal margin smoothly
rounded in ventral view; cymbium length/width ratio 1.8; embolus originating at ca. 4:30 o’clock position.
Female. Unknown.
Distribution. Known only from the type locality in Lebap Region, northeastern Turkmenistan (Fig. 12A)
Zodariellum turkmenicum sp. n.
Figs 1F, 2E, 3E, 4E, 5E, 6E, 12A
Type material. Holotype ♂ (ZMMU), TURKMENISTAN: Lebap Region: 90 km WNW of Repetek, [38°46'N
62°21'E], saline land, 16.05.1978 (V. Kaplin).
Etymology. The specific epithet refers to Turkmenistan, from where the holotype was collected.
Diagnosis. The male of this species differs from those of its congeners known in the region (except for Z.
surprisum) by the uniformly coloured carapace and abdomen. The palp of the new species is most similar to those of
Z. surprisum and Z. continentale by having unbranched and gradually tapering process of RTA with sharply pointed
tip, but can be differentiated from Z. surprisum by the process of RTA directed proximally (vs. anterolaterally). The
palp of Z. turkmenicum sp. n. is similar to that very schematicly illustrated by Andreeva & Tyshchenko (1968) for
Z. continentale, but differs by having a triangular outgrowth at the tip of RTA (To, Fig. 2E).
Description. Male. Habitus as in Fig. 1F. Total length 4.65. Carapace 2.35 long, 1.97 wide. Eye sizes: AME:
0.26, ALE: 0.11, PME: 0.12, PLE: 0.11. Carapace, sternum, chelicerae, labium and maxillae uniformly dark brown.
Legs colored as carapace on the proximal segments, yellowish on the distal ones. Abdomen dark maroon, lacking
any pattern, lighter ventrally. Spinnerets uniformly yellowish brown. Measurements of legs (unlisted segments
missing): Fe I: 2.46, II: Fe 2.71, Pa 0.93, Ti 2.31, Mt 3.44, III: 12.02 (2.95, 0.96, 2.70, 3.85, 1.56), IV: 13.86 (3.72,
0.96, 3.29, 4.24, 1.65).
Palp as in Figs 2E, 3E, 4E, 5E, 6E. RTA 2 times longer than tibia, and almost 3 times thinner than tibia, tip
roundly bent, with only one process; cymbium 2.2 times longer than tibia together with RTA, 1.9 times longer than
wide; bulb as in congeners with exception to long spur (Sp) of tegular apophysis, about as long as RTA; embolus
originating at about 5:30 o’clock position.
Female. Unknown.
Distribution. Known only from the type locality in Lebap Region, northeastern Turkmenistan (Fig. 12A).
ZAMANI & MARUSIK
176 · Zootaxa 5178 (2) © 2022 Magnolia Press
Zodariellum zebra (Charitonov, 1946) comb. n.
Fig. 12A
Zodarion zebra Charitonov, 1946: 21, fig. 10 (♀).
Zodarion zebra: Charitonov 1969: 71.
Comments. This species is hereby transferred to Zodariellum due to its screw-like copulatory ducts, a character
unknown in other Zodariinae. Judging from the name and the detail verbal description by Charitonov (1969), it
is very likely that Z. zebra is a junior synonym of Z. testaceofasciatum, a species with striped abdomen and type
locality not far from that of Z. zebra.
Distribution. Uzbekistan (Fig. 12A).
Discussion
In this paper, we provided an emended diagnosis and the first species-group classification for the genus Zodariellum.
Furthermore, we described two new species from Turkmenistan, and proposed six new combinations and two new
synonymies, raising the total number of species known in this genus to 21.
One particularly important finding reported here is the presence of a deep and conical diverticulum on the
retrodorsal side of the male palp’s basal area in all Zodariellum species with known males. Jocqué & Henrard
(2015) reported a similar structure in the form of a short conical round pit in Acanthinozodium (termed ‘cymbial
pit’, assumed to represent the outlet of a gland that produces a mating plug), and considered it a diagnostic character
for the entire genus (it is noteworthy that the male of the type species, A. spinulosum Denis, 1966, remains
undescribed). Following this, Zamani & Marusik (2021) described 12 species in Acanthinozodium, all bearing a
cymbial diverticulum (or ‘cymbial groove’) but at the same time showing differences in the overall configuration of
the palpal structures. The finding of this diverticulum in Zodariellum proves that this structure is more widespread
than once thought, and on its own cannot be used as a diagnostic generic character.
Besides species of Acanthinozodium and Zodariellum, such diverticulum is present in at least one more species,
Zodarion ovatum Zhang & Zhang, 2019 (see: Zhang & Zhang 2019: fig. 7C). Although the male palp of this
species is somewhat similar to that of Z. nitidum by having short and broad RTA and filamentous embolus, it has
large tegular apophysis with much longer conductor and small base of embolus. The endogyne of Z. ovatum is very
different from that of Z. nitidum, which has twisted and diverging copulatory ducts. Zodarion ovatum has large
bursae unknown in other Zodariinae (see: Zhang & Zhang 2019: figs 7E and 8E, marked as ‘copulatory duct’ in the
latter). Accounting the presence of a deep cymbial diverticulum and several differences in the epigyne and the male
palp of Z. ovatum and Z. nitidum, we believe that the Chinese species should be placed in a separate genus.
Furthermore, considering that the cymbial diverticulum occurs not only in Acanthinozodium as previously
thought, a revision on the species currently assigned to this genus seems necessary; most likely, some of the
species described in Acanthinozodium by Zamani & Marusik (2021) are misplaced and may represent undescribed
genera. Finally, species of the spinibarbe group of Zodarion show great similarity to many of those described in
Acanthinozodium, and it is possible that at least some of them are misplaced and should perhaps be transferred to
that genus.
Acknowledgments
We are grateful to Wioletta Wawer (Museum and Institute of Zoology, Polish Academy of Sciences, Poland) for
providing information on the depository of the material in their collection, and Feng Zhang (Hebei University,
Baoding, China) for confirming the presence of cymbial diverticulum in Z. ovatum. Yuri Marusik is thankful to
Seppo Koponen (Turku, Finland) for arranging his stay in Turku. The comments and suggestions by the subject
editor Arnaud Henrard (Tervuren, Belgium) and the reviewer Tarık Danışman (Kırıkkale, Turkey) are greatly
appreciated.
NEW TAXONOMIC CONSIDERATIONS IN ZODARIELLUM Zootaxa 5178 (2) © 2022 Magnolia Press · 177
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