Zootaxa 3846 (3): 398410

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Copyright 2014 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

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http://dx.doi.org/10.11646/zootaxa.3846.3.4
http://zoobank.org/urn:lsid:zoobank.org:pub:47DD5009-C837-4A1D-B800-DB8BE098F960

A re-description of rare alpheid shrimp Amphibetaeus jousseaumei (Coutire,

1896) and description of a new species of the genus Athanopsis Coutire, 1897
(Crustacea: Decapoda: Alpheidae) from Iranian coast of the Persian Gulf
IVAN MARIN1), RAHELEH SHEIBANI2) & ALIREZA SARI3)
1)

A. N. Severtzov Institute of Ecology and Evolution of RAS, Moscow, Russia

Email: coralliodecapoda@mail.ru, vanomarin@yahoo.com
2)
Austrian Institute of Technology Gmbh (AIT), Konrad-Lorenz-Strasse 24, 3430 Tulln, Austria
Email: raheleh.sheibani@ugent.be, rahel14350@yahoo.com
3)
School of Biology and Centre of Excellence in Phylogeny of Living Organisms, College of Science, University of Tehran, Tehran, Iran
Email: sari@ut.ac.ir
Corresponding authorcoralliodecapoda@mail.ruIvan Marin

Abstract
A new record of rare alpheid shrimp Amphibetaeus jousseaumei (Coutire, 1896) (Decapoda: Caridea: Alpheidae) is presented based on material collected from Qeshm Island, Iran, the Persian Gulf. This is the first record of the species for the
Iranian waters as well as the second record and re-description of A. jousseaumei from its original description given by
Coutire (1896). Besides, a new alpheid shrimp species Athanopsis tarahomii sp. nov. is described from the same locality.
The new species can be clearly separated from all remaining species of the genus Athanopsis by deep rostrum curved
downward and the presence of well marked blunt extra-cornea tooth on carapace.
Key words: Crustacea, Decapoda, Alpheidae, Amphibetaeus, Athanopsis, new records, new species, Persian Gulf, Iran

INTRODUCTION
The invertebrate fauna of the Northwest Indian Ocean including the Persian Gulf and the Gulf of Oman are purely
studied and crustaceans are not the exception in this regard. Several new species of caridean shrimp were recently
described from Iranian waters (Marin, 2009; Anker et al, 2010). During the course of study on caridean shrimps of
the Iranian intertidal zone (Sheibani, 2008) at Qeshm Island, the largest Iranian Island of the Persian Gulf located at
the entrance of the Persian Gulf at the Hormoz Strait (26 51' N, 56 08' E), at January 2008, representatives of two
rare genera of the family Alpheidae were found. Below, the first re-description of Amphibetaeus jousseaumei
(Coutire, 1896), since its original description (Coutire, 1896) based on freshly collected material is given. Three
Coutires syntypes from Djibouti and southern Yemen, Gulf of Aden were deposited in the Museum National
dHistoire Naturelle, Paris (MNHN). Presently, only the major cheliped of one of these specimens is designated as
the lectotype of A. jousseaumei in MNHN (MNHN-Na 13677). The other record belongs to undescribed species of
the relatively rare alpheid genus Athanopsis Coutire, 1897, which is described here in as a new species.
All shrimps were caught by small hand net at subtidal zone under rocks and coral boulders and were
immediately preserved in 70% solution of ethanol. Carapace length (cl., in mm), the length from the tip of rostrum
to the posterior margin of the carapace, and total length (tl., in mm), the length from the tip of rostrum to the distal
margin of the telson, are used as the standard measurement of size. The material of Amphibetaeus jousseaumei and
Athanopsis tarahomii sp. nov. is deposited at Senckenberg Museum (SMF), Frankfurt am Main, Germany and
Zoological Museum, University of Tehran (ZUTC), Tehran, Iran.

398 Accepted by J. Goy: 10 Jul. 2014; published: 4 Aug. 2014

SYSTEMATIC PART
Order Decapoda Latreille, 1802
Family Alpheidae Rafinesque, 1815
Genus Amphibetaeus Coutire, 1896
Amphibetaeus jousseaumei Coutire, 1896
(Figs. 13)
Betaeus jousseaumei Coutire, 1896a: 313, figs. 112; Coutire, 1896b: 236.
Amphibetaeus jousseaumei Coutire, 1896c: 384.

Material examined.Neotype, male (cl. 11.2 mm, tl. 30.0 mm) (SMF-43232) and female (cl. 12.0 mm, tl. 33.0
mm, female material get lost after drawing)Northwest Indian Ocean, Qeshm Island, the Persian Gulf, Iran,
2651N, 5608E, subtidal, under stone, 10Jan2008.
Description.Rather large shrimp with robust body. Carapace glabrous, smooth, with well marked cardiac
notch (Fig.1a), no any teeth, orbital hood completely overlapping eyes (Fig. 1b, c); dorsal or lateral carina absent;
pterygostomial angle rounded, slightly produced. Eyes large, with well pigmented cornea, eyestalk robust, distally
bluntly produced (Fig. 1b).
Antennular peduncle (Fig. 1c) robust; basal segment with large, strong tooth ventromedially, stylocerite robust,
triangular, slightly overreaching basal antennular segment; second segment subequal, about as long as wide; distal
segment as long as wide; upper flagellum short, biramous, with 5 proximal robust segments, shorter ramus with
34 segments and each with group of 23 aesthetascs, longer ramus with more than 10 segments. Lower flagellum
longer than upper and well developed.
Antenna (Fig. 1a, h) with robust basicerite with acute robust ventral tooth; carpocerite slightly overreaching to
just beyond the tip of scaphocerite, about twice longer than wide; scaphocerite well developed, subequal to length
of antennular peduncle, lamina about 1.3 times longer than wide, blade slightly overreaching distolateral tooth,
lateral border slightly convex, distolateral tooth small and blunt.
Abdominal somites smooth, ventrolateral posterior border of fourth and fifth somites pubescent, sixth somite
with fused triangular plate at posterolateral edge (Fig. 1e). Telson (Fig. 1d, e) about 1.5 times longer than proximal
width, tapering distally, lateral margins slightly convex, dorsal surface with 2 pairs of developed submarginal
dorsal spines, about 0.2 of telson length, subequal in slight depressions; distal margin of telson (Fig. 1f) rounded,
with submarginal row of short setae and longer terminal posterior setae, distolateral corner with 2 pairs of spines,
lateral spines about half length of mesial spines, between posterior spines and longer terminal posterior setae.
Uropods overreaching telson (Fig. 1d); exopod of uropod (Fig. 1g) with feebly convex lateral margin covered with
small setae, dieresis with subdistal movable acute spine, distolateral corner subzngulated, unarmed, midpart
bearing deep rounded depression.
Mandible (Fig. 2a) robust, with strong molar process, incisor process with 5 teeth and palp 2segmented,
reaching about to end of incisor process. Maxillula (Fig. 2b) with bilobed palp, upper lobe having 2 slender setae
distally and lower lobe with single longer and thicker setae distally; upper lacinia stout, with numerous setae on
mesial margin, lower lacinia slender with 510 long spines and numerous small setae distally. Maxilla (Fig. 2c)
with well developed tapering palp, distal endite feebly bilobed with small setae on mesial margin, proximal endite
short, thumb-like, with scaphognathite broad. First maxilliped (Fig. 2d) with well developed 2-segmented palp;
epipod large and broad, bilobed; exopod very long, slender, with small caridean lobe fringed with plumose setae;
distal endite broad with numerous setae medially, separated by notch from broadly rounded and less setose
proximal endite. Second maxilliped (Fig. 2e) normal, with large ear-shaped epipod; exopod large and slender, with
several long setae distally; coxa rounded with few setae mesially; ischiobasis robust, propodus stout, greatly
convex distodorsally; dactylus about 2.5 times longer than wide, with numerous simple spines along outer margin.
Third maxilliped (Fig. 2f) exceeding antennular peduncle by tip of ultimate segment; with ear-shaped epipod,
arthrobranch and setobranch; exopod slender, equal to the length of antepenultimate segment, antepenultimate
segment about 5 times longer than broadest width; penultimate segment about 0.4 times as long as antepenultimate

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FIGURE 1. Amphibetaeus jousseaumei Coutire, 1896, a-j female , k male (SMF-43232) : acarapace, lateral view; bfront
of carapace, lateral view; csame, dorsal view; dlast abdominal segments, telson and uropods, dorsal view; esame, lateral
view; fdistal margin of telson, dorsal view; gexopod of uropods, dorsal view; hantenna; isecond pereiopod; jthird
pereiopod; kendopod of second pleopod.

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FIGURE 2. Amphibetaeus jousseaumei Coutire, 1896, female : amandible; bmaxillula; cmaxilla; dfirst maxilliped ;
esecond maxilliped ; fthird maxilliped .

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FIGURE 3. Amphibetaeus jousseaumei Coutire, 1896, a-g female, h male (SMF-43232): acarapace and appendages; bc
first major pereiopod , different views; e, fsame, fingers; gminor first pereiopod ; hsame, chela.

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segment, about twice longer than wide; distal segment slender, about 2 times longer than penultimate and about 5
times longer than wide, tapering distally, with more than 15 rows of stout setae and spines distally.
First pereiopods unequal in size and shape, both in male and female, carried flexed. Total length of major chela
equal to postorbital length of carapace (Fig. 3a). Major cheliped (Fig. 3ad) with slender proximal segments and
stout chela; ischium about 1.5 times as long as wide, unarmed; merus slender, ventrally depressed, about 4 times
longer than wide, widening distally, with straight smooth margins; carpus stout, cup-shaped, unarmed. Palm (Fig. 3
b-d) stout, smooth, subcylindrical about 2 times as long as proximal width, subequal to fingers in length; dorsal
side depressed proximally; lateral margin near dactyl-propodal articulation transversely constricted, adhesive
p;aque large. Fingers (Fig. 3bd, e, f) strong, compressed surfaces smooth, tips strongly curved, crossing, fixed
finger with pointed tip, cutting edgewith 2 proximal triangular teeth and small tvercle-like tooth in distal third,
midpart with large deep fossa; movable finger slightly shorter than fixed finger, cutting edge with 2 triangular
proximal teeth and large flattened tooth fitting into fossa on the fixed finger. Minor cheliped (Fig. 3g) significantly
smaller than major; ischium small, about 1.5 times as long as wide; merus slender, about 7 times as long as wide,
ventral margin depressed, unarmed; carpus (Fig. 3h) stout, cup-shaped; palm stout, swollen, surfaces smooth,
slightly shorter than fingers; fingers slender, compressed, tips sharply pointed, crossing, cutting edges forming thin
plate with small triangular teeth on both cutting edges at distal third.
Second pereiopod (Fig. 1i) slender; coxa with setobranch and hook-like epipod; basis slightly longer than
wide, unarmed; ischium and merus subequal, about 8 times longer than wide, unarmed; carpus longer than merus,
subdivided into 5 segments, with ratio about 5:1:1:1:2, with distal segment subequal to length of chela; chela small,
palm subequal to the length of fingers, about 1.7 times longer than wide, fingers simple, about twice as long as
wide, tips sharply pointed.
Ambulatory pereiopods robust, with slender simple dactylus. Third pereiopod (Fig. 1j) with robust basal
segments; coxa with setobranch and hook-like epipod; ischium as long as wide, unarmed; merus with convex
smooth margins, about 3.5 times as long as maximum width, unarmed; carpus relatively slender, subequal to
propodus, about 4 times as long as maximal width, expanded distally, with small distal spine on flexor margin;
propodus about 4 times longer than wide; flexor margin with 2 spines and one pair of small distal spines; dactylus
simple, sharply pointed, slightly curved, about 0.3 times length of propodus and 3 times longer than proximal
width. Fourth and fifth pereiopods similar to third.
Second pleopod with appendix masculina (Fig. 1k) twice shorter than appendix interna, 0.15 times length of
endopod.
Remarks.Originally, the species was well described by Coutire (1896a, b) and our specimens are identical
to its original description. Unfortunately, the bodies and most appendages of the all syntypes designated by
Coutire and deposited in the collection of MNHN were lost and only one large chela is still deposited. This chela
indicated as a lectotype was re-described by Anker & Jeng (2006) with presented diagnosis of the genus. At the
same time, new record of this rare species which has been never found since its original description in 1896 as well
as the absence of the type specimen are the reasons for presenting this re-description from the Persian Gulf and
designation of neotype.
Coloration.Not recorded. The species is probably translucent with some reddish tiny chromatophores.
Habitats.The described specimens were found on sand-muddy flat bottom, under large boulder, together
with large burrowing ghost shrimp Neocallichirus jousseaumei (Nobili, 1904) (Callianasidae) and two echiuran
individuals. It was already mentioned that the species could be symbiotic with burrowing ghost-shrimps,
sipunculans or echiurans (Coutire 1899). Association with large burrowing callianassid shrimps is characteristic
for some alpheids (for example, Anker & Marin, 2009).
Distribution.Known only from the northwest part of the Indian Ocean: from Qeshm Island, Iran; Bab al
Mandab Strait between the Red Sea and the Gulf of Aden; from Tadjourah (the Republic of Djibouti) and Perim
(off southwestern coast of the Republic of Yemen).

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Genus Athanopsis Coutire, 1897

Athanopsis tarahomii sp. nov.
(Figs. 47)
Material examined.Holotype, male (cl. 8.0 mm, tl. 22.0 mm) (SMF-43233); paratype, female (cl. 9.0 mm, tl.
23.0 mm) (ZUTC Nat. 1118)Northwest Indian Ocean, the Persian Gulf, Iran, Qeshm Island, 2651N, 5608E,
subtidal, under stone, 11Jan2008.
Description.Medium-sized robust shrimp. Carapace (Fig. 4ac) glabrous, smooth, unarmed; orbital hoods
partly covered eyes, extra-cornea tooth blunt, well marked, adrostral furrow deep, rounded. Rostrum well
developed, in lateral view, deep, compressed, curved downward, about 1.5 times as long as wide at base, dorsal
margin with small rounded tooth at distal third; median side with laminal plate reaching to distal third of rostrum;
tip rounded, reaching to the middle of the second antennular segment (Fig. 4a, b, d, e). Rostrum in dorsal view,
broad triangular with median ridge, as long as proximal width, tip bluntly pointed (Fig. 4c). Pterygostomial angle
slightly produced, bluntly rounded (Fig. 4a, d, e).
Antennular peduncle (Fig. 4h) robust; first segment stout, about 1.5 times longer than distal width, stylocerite
large, with convex lateral margin, tip bluntly pointed, reaching to the midlength of second segment, mesial margin
with blunt small tooth; second segment subequal to third segment, about as long as wide, upper flagellum short,
biramous, with 34 proximal robust and partly fused segments, shorter ramus with 9 segments and each with group
of 3 aesthetascs, longer ramus with more than 30 segments, lower flagellum longer than upper.
Antenna (Fig. 4a) robust, with robust basicerite bearing large sharp dorsolateral tooth and acute ventral tooth;
carpocerite overreaching scaphocerite, scaphocerite well developed, about 2.5 times longer than maximum width,
with small distolateral tooth reaching beyond the blade. Flagellum longer than postorbital carapace length.
Antennal gland opening on small process.
Abdomen smooth, ventrolateral posterior border of fourth and fifth somites pubescent, sixth somite with
movable bluntly rounded triangular plate at posterolateral edge (Fig. 4g). Telson (Fig. 4f, g) with slightly convex
lateral margins, about 1.4 times as long as maximaum width, about 1.3 length of sixth abdominal somite, with
posteriorly rounded margin about half of proximal margin, dorsal surface with 2 pairs of small submarginal spines,
anterior part situated in midlength of telson; distal margin with numerous long setae and submarginal row of short
setae, lateral corner with one pair of spines, mesial spine about twice longer than lateral spines.
Uropods with sympodite greatly produced distally, sharp pointed, tip reaching little less than half of exopod
(Fig. 4f, g); exopod shorter than endopod and telson, lateralmargin almost straight, dieresis with small acute
distolateral tooth and long sublateral spine (Fig. 4i); endopod normal.
Mouthparts typical of genus. Mandible (Fig. 5a) robust, with strong molar process, incisor process with 67
triangular teeth along distal margin, palp 2-segmented, tip reaching to the midlength on molar process. Maxillula
(Fig. 5b) with bilobed palp, distal lobe bearing one row of small setae distally, proximal lobe with single long seta;
distal lacinia with setae and short blunt spines distally, proximal lacinia slender with several long spines and
numerous setae distally. Maxilla (Fig. 5c) with tapering palp, proximal endite short and thumb-like, distal endite
bilobed, distal lobe with setae mesially, proximal lobe small; scaphognathite broad, with marginal setae. First
maxilliped (Fig. 5d) with epipod large and broad; palp long, rather broad, bluntly pointed with plumose apical seta;
mesial margin with setae; exopod long and slender, caridean lobe feebly developed, with plumose setae laterally;
distal endite broad, bilobed, proximal small, rounded. Second maxilliped (Fig. 5e) normal, with ear-shaped epipod
and well developed long slender exopod. Endopod robust; ischiobasis, merus longer than carpus, carpus as long as
wide; propodus with rounded distal lobe fringed with long setae; dactylar narrow, about 3 times wider than long,
with numerous small teeth and simple setae along mesial margin. Third maxilliped (Fig. 5f) robust, exceeding
antennular peduncle by tip of ultimate segment; epipod with terminal hook; exopod rather broad, subequal to the
length of antepenultimate segment; antepenultimate segment about 3 times longer than maximum width,
significantly broadening distally; penultimate quadrate in shape, as long as wide; ultimate segment slender, about 3
times longer than penultimate, and 4 times longer than wide, tapering distally, mesial margin with 1520 rows of
setae distal corner with tufts of long simple setae (Fig. 31).
First pereiopods unequal (Fig. 6ad) in both size and shape, in male and female, heavier and wider in males,
carried flexed beneath the body. Total length of major chela equal to postorbital length of carapace. Major cheliped

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FIGURE 4. Athanopsis tarahomii sp. n., ad, fi paratype female (ZUTC Nat. 1118), e holotype male (SMF-43233) a,
bfront of carapace, lateral view; csame, dorsal view; d, erostrum; f, gsixth abdominal segment, telson and uropods;
hantennula; idistolateral margin of uropodal exopod.

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FIGURE 5. Athanopsis tarahomii sp. n., paratype female ( ZUTC Nat. 1118): amandible; bmaxillula; cmaxilla;
dfirst maxilliped ; esecond maxilliped ; fthird maxilliped.

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FIGURE 6. Athanopsis tarahomii sp. n., paratype female (ZUTC Nat. 1118): a, bmajor first pereiopod ; cminor first
pereiopod ; dsame, chela; esecond pereiopod ; fthird pereiopod; gfourth pereiopod ; hfifth pereiopod.

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FIGURE 7. Alive coloration of paratype female (ZUTC Nat. 1118) of Athanopsis tarahomii sp. n.

(Fig. 6a, b) with basis stout, unarmed; ischium as long as wide, bearing 3 teeth on dorsal margin, ventral margin
with well marked blunt triangular tubercle (Fig. 6b) merus unarmed, expanded distally, about 2.5 longer than
maximum width, ventral margin excavated, ventrolateral margin strongly convex distally; carpus cup-shaped,
about as long as wide, with sharply produced distoventral margin; palm unarmed, about 3.5 times longer than
fingers, about 3 times as long as maximum width, mesial face strongly convex, ventral face excavated; fingers
compressed with flattened tips, tips bearing 7-8 small triangular teeth, fixed finger about twice longer than wide,
cutting edge unarmed; movable finger about 2.5 times as long as wide, curved downward, cutting edge with 2 large
proximal teeth and 8 small triangular teeth along distal margin. Minor cheliped (Fig. 6c-d) rather slender; coxa
about as long as wide, unarmed; ischium about 1.5 times as long as wide, unarmed, with small tubercle at ventral
margin; merus about 3.5 times as long as wide, with straight margins, unarmed; carpus cup-shaped, slightly longer
than wide, unarmed; palm (Fig. 6d) cylindrical, about 1.5 times longer than fingers, about 2.5 times as long as
wide, surfaces smooth; fingers curved downward, about 3 times longer than wide, tips pointed, cutting edges
bearing numerous small teeth.
Second pereiopod (Fig. 6e) slender; coxa with setobranch and stick-like epipod; basis about slightly longer
than wide, unarmed; ischium about twice longer than wide and 2 times longer than basis, unarmed; carpus long,
subdivided into 5 segments, with ratio about 8:1:1:1:2, with distal article subequal to length of palm; chela small,
palm subequal to the length of fingers, about 1.2 times longer than wide, fingers simple, about 2.5 times as long as
wide, tips sharply pointed.
Ambulatory pereiopods robust, similar in size and shape. Third pereiopod (Fig. 6f) with ischium armed with 2

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spines on ventral margin, merus about 3 times as long as wide, dorsal margin slightly convex, unarmed,
distoventral corner obtusely angulated; carpus about 2.3 times as long as wide, slightly expanded distally, ventral
margin with 4 single spines and one pair of distal spines; dactylus stout, simple, slightly curved, about 2.5 times as
long as propodus, about 2.5 times longer than proximal width, tip sharply pointed. Fourth pereiopod (Fig. 6g) with
ischium armed with 2 spines on ventral margin, merus about 2.5 times as long as wide, with slight convex dorsal
margin, dorsoventral corner obtusely angulated, unarmed; carpus about 2 times as long as wide, slightly expanded
distally, distoventral margin with single spine, propodus about 3 times as long as wide, ventral margin with 3 single
spines and one pair of distal spines; dactylus stout slightly curved, about 2 times as long as propodus, about 2.5
times longer than proximal width, tip sharply pointed. Fifth pereiopod (Fig. 6h) with ischium armed with 2 spines
on ventral margin, merus about 2.5 times as long as wide, with straight dorsal margin, unarmed, distoventral corner
obtusely angulated; carpus about 2 times as long as wide, slightly expanded distally, ventral margin with one pair
of distal spines; propodus rather slender, about 4.5 times as long as wide, ventral margin with 2 spines and one pair
of distal spines, distal margin with row of simple setae, dactylus stout, simple, slightly curved, about 4 times as
long as propodus, about 2.5 times longer than proximal width, tip sharply pointed.
Endopod of male first pleopod about 4 times longer than width, tapering distally with 2 terminal plumose
setae, medial border with 6 plumose setae and lateral border with 3 simple setae. Second pleopod with appendix
masculina slightly shorter than appendix interna, about 5 times longer than wide, with 6-7 distal spines.
Differential diagnosis.Recently the genus Athanopsis includes 6 described species such as a type species of
the genus, A. platyrhynchus Coutire, 1897 from Djibouti; A. dentipes Miya, 1980 from sandy mud of Amakusa
and Nagasaki, Kyushu, Japan; A. brevirostris Banner & Banner, 1981 from southern Red Sea, among corals; A.
australis Banner & Banner, 1982 from sandy sediment of Port Phillip Bay, A. rubricinctuta Berggren, 1991 from
Mozambique and A. saurus Anker, 2011 from Australia. The new species is the first record of the genus from the
Persian Gulf. The new species can be clearly separated from other species of the genus by deep, curved downward
rostrum and the presence of well marked blunt extra-cornea teeth. The new species can be clearly separated from A.
platyrhynchus by deeper rostrum and expanded palm nd merus of the major cheliped; from A. dentipes by deeper,
curved downward rostrum and well developed endites of maxilla and first maxilliped; from A. brevirostris by deep
well developed rostrum, blunt extra-corneal teeth, shorter stylocerite of antenulla and; from A. australis by deeper
rostrum, absence of meral ventral spine of third pereiopod, the presence of two spines on ischium of ambulatory
pereiopods and broader telson; from A. rubricinctuta by deeper rostrum, the presence of well marked extra-corneal
teeth, smooth lateral margin of propodus of major first pereiopod, shorter finger of minor first pereiopod and
absence of meral ventral spine of ambulatory pereiopod. The most morphologically similar species is A. dentipes
Miya, 1980 known from Japan.
Coloration.General body and appendages are yellowish translucent; antennula, distal part of scaphocerite,
dorsal margin of carapace and abdominal somites, telson and uropods covered with tiny red chromatophores (Fig.
7).
Etymology.This species named after the late PhD student M. Tarahomi who tragically passed away in Iran
during field collection in October 2009.
Habitat.The specimens were found on sandy soft bottom in cobble/rocky area under large boulder in
association with echiurans similar to other species of the genus.
Distribution.Presently the species is known from type locality, Qeshm Island, Iran, the Persian Gulf, only.

ACKNOWLEDGMENTS
The authors would like to thank Office of Research Affairs, University of Tehran for providing financial support of
the field collecting. The second author (RS) thanks Dr A. Anker (University of Florida) for his critical comments
on the identity of these specimens and Professor M. Trkay (Senckenberg Museum) for the hospitality and
providing research facility at Senckenberg Museum, Frankfurt am Main, Germany. We thank A. Kazemi and Dr. R.
Naderloo for their invaluable help during sampling. This study was supported by Grant of the President of Russian
Federation -4481.2014.4 (to IM) and Russian Foundation of Fundamental Researches 12-04-00540- (to IM).

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