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A new species of Pseudocerastes with elaborate tail ornamentation from western Iran (Squamata: Viperidae)

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Abstract

A new species of viper, Pseudocerastes urarachnoides, is described from the Zagros Mountains in western Iran. The new species has a short tail, few pairs of subcaudals (15 in the known specimens), the distal pairs forming an oval knob-like structure; lat-eral dorsal caudal scales projected to form elongate "appendages" along the sides of the terminal knob. Several rows of lateral dorsal scales are weakly keeled and outer rows are only faintly keeled. We speculate that the caudal appendage may serve as a lure for prey in an ambush predator.
A New Species of Pseudocerastes with Elaborate Tail
Ornamentation from Western Iran (Squamata: Viperidae)
Hamid Bostanchi1, Steven C. Anderson2,5,6, Haji Gholi Kami3, and Theodore J. Papenfuss4,5
1P.O.Box 31375-378, Mehrvilla, Karaj, Iran;Email:hamid_bostanchi2005@yahoo.com;
2Department of Biological Sciences, University of the Pacific, Stockton, California, USA 95211;
Email:asaccus@aol.com;3Department of Biology, Faculty of Sciences, Agriculture Sciences and
Natural Resources, Gorgan University, Gorgan, Golestan Province, Iran;Email:hgkami2000@
yahoo.com;4Museum of Vertebrate Zoology, University of California, Berkeley, California,
USA 94720;Email:asiaherp@calmail.berkeley.edu.
A new species of viper, Pseudocerastes urarachnoides, is described from the Zagros
Mountains in western Iran. The new species has a short tail, few pairs of subcaudals
(15 in the known specimens), the distal pairs forming an oval knob-like structure; lat-
eral dorsal caudal scales projected to form elongate “appendages” along the sides of
the terminal knob. Several rows of lateral dorsal scales are weakly keeled and outer
rows are only faintly keeled. We speculate that the caudal appendage may serve as a
lure for prey in an ambush predator.
KEYWORDS: Pseudocerastes urarachnoides, Pseudocerastes persicus, Pseudocerastes fieldi,
new taxon, Iran, caudal lure, caudal ornamentation, Viperidae
The Second Street Expedition to Iran (1968) collected amphibians and reptiles incidental to the
mammals that were the primary objectives. These specimens were deposited in the Field Museum
of Natural History, where one of us (SCA) examined and identified them in 1970. The first impres-
sion of FMNH 170292, seen through the bottle in the preparations room, was that a small solpugid
was clinging to its tail. Subsequent examination revealed that the snake was a specimen of
Pseudocerastes with a peculiar growth at the tail tip. It was identified as P. persicus, with which it
agreed in most particulars. As there was only the single specimen, it was not possible to say whether
the pecular growth of the tip of the tail had a genetic origin or was, perhaps, some sort of tumor or
caused by some parasite. Thus, the specimen languished, but was not forgotten, for nearly four
decades. Then, in 2003, one of us (HB) collected a second specimen with identical tail ornamenta-
tion and similar scale counts and morphology. It seems likely that there has been genetic continu-
ity in this character over the past 35 years. Were the caudal appendage the result of a characteristic
tumor or due to the action of a parasite, we might expect it to have been observed elsewhere and,
perhaps, in other species. We believe that these two specimens represent an undescribed species,
related closely to Pseudocerastes fieldi and P. persicus. This appears to be the most elaborate mor-
phological caudal ornamentation yet reported in a snake, with the possible exception of the rattles
of Crotalus and Sistrurus.
PROCEEDINGS OF THE CALIFORNIAACADEMY OF SCIENCES
Fourth Series
Volume 57, No. 14, pp. 443–450, 13 figs., 1 table. September 15, 2006
5Research Associate, California Academy of Sciences.
6Address for correspondence.
443
Reprinted from the PCAS, ser. 4, vol. 57 (15 September 2006)
Pseudocerastes Boulenger, 1896
TYPE SPECIES: Cerastes persicus Duméril, Bibron, and Duméril, 1854, by monotypy
DEFINITION.— Head distinct from neck, covered with small scales; pupil of eye vertical; nos-
tril directed outwards and upwards, in large undivided nasal shield (pierced between two small
scales, a larger crescentic anterior and a smaller scale-like posterior [Gasperetti 1988:350]) (nasal
aperture in a large circular or crescentic shield, the upper part of the aperture leading into the
supranasal sac [Smith 1943:490, fig. 155A]); supralabials with serrated lower margin and with
inner groove to receive lower lip (The structure of the lips, to provide complete closure of the
mouth, and the valvular prominence within the nasal aperture, are typical desert modifications
against the ingress of blown sand. They are found also in Eristocophis [Smith 1943:490]); body
scales in 21–25 longitudinal rows, none obliquely disposed; keels on body scales do not reach pos-
terior edge of scale but end in swollen knob before outer edge, keels not serrated; ventrals round-
ed, without lateral keels; tail short, subcaudals paired (Leviton et al. 1992:114–115).
DISTRIBUTION.The North Arabian Desert from Sinai and southern Israel, Jordan, Iraq,
southwestern Iran east to Afghanistan and Pakistan west of the Indus River, outlying population in
northern Oman. (Fig. 13).
We include here brief descriptions of the previously recognized taxa of Pseudocerastes for
comparison with the new species. Some authors have considered P. fieldi a subspecies of P. persi-
cus. There appears to be a geographic hiatus in the distribution of the genus, P. persicus and P. fiel-
di nowhere known to be parapatric, their ranges separated by the Zagros Mountains. For this rea-
son and the differences in venom properties (see below), we prefer to recognize them at the species
level.
Pseudocerastes persicus (Duméril, Bibron, and Duméril, 1854)
Cerastes persicus Duméril, Bibron, and Duméril, 1854:1443, pl. 78b.
DESCRIPTION (From Smith 1943:490–492, fig. 155).Head depressed, snout short and broad-
ly rounded; diameter of the eye less than its distance from the mouth; nostril very large, pierced in
a large circular or crescentic nasal, bounded above by a supranasal which may be broken up; two
scales between the nasal and the rostral; scales on top of the head small, imbricate, smooth on the
snout, keeled behind in the young, tuberculate and more strongly keeled in the adult; an erect horn-
like scale above the eye surrounded by small scales; 9–12 scales on a line between the horns; 16–20
scales round the eye; 3–4 scales on a line between the eye and the nasal; temporal scales small,
keeled; 13–14 supralabials, 4 series of scales between them and the eye, 1st pair of infralabials larg-
er than the others; a pair of large anterior genials, the scales posterior to them being much smaller.
Scales in 23 or 25:23 or 25: 19 rows, striated and strongly keeled, the outermost scales strong-
ly overlapping the ventral scales. V 144–158; C 34–49, paired.
Hemipenis short, extending to the 8th caudal plate, deeply forked; the distal end is calyculate,
the remainder spinose, the largest spines being at the proximal end; sulcus lips also spinose.
Snout-vent length 688–690 mm, tail 80–85 mm.
Grayish-brown above, with squarish, dark brown, black-edged spots, which alternate with one
another on either side of the vertebral line, or are confluent to form cross-bars; sides of the body
with rounded, less distinct spots; top of the head pale grey, upper lip and side of the head darker,
the two colors meeting in a sharply defined line which extends from the eye to the angle of the
mouth; whitish beneath, spotted with brown. In the adult the markings are much less distinct and
may be almost entirely absent. In large adults, tip of tail often dark. An adult from Kacha,
444 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Baluchistan, is heavily marked and mottled with black and cream.
Total length: 890 mm; tail 110 mm.
DISTRIBUTION.Southeastern Anatolia, Turkey and Iran east of the Zagros Mountains
through the Iranian Plateau to central Afghanistan and western Pakistan (Fig. 13). Gasperetti
(1988:353, fig. 127) shows four localities in northern Oman and the Musandam Peninsula.
NOTE.— Wall (1913:64) described Pseudocerastes bicornis from Khajuri Kach, Waziristan,
Pakistan, based on a single specimen now in the Natural History Museum, London. The specimen
consists of the head and anterior one-fourth of the body (Smith 1943:492). Subsequent authors
(Minton 1966; Mertens 1969; Gasperetti 1988) have placed it in the synonymy of P. persicus,
although Khan (2002:190) regards it as distinct. It must be noted that were the two specimens here
described as a new species missing the tails, they would be identified as P. persicus.
Pseudocerastes fieldi Schmidt, 1930
Pseudocerastes fieldi Schmidt, 1930:227, fig. 2 (Type locality: Ayn Bair, Jordan).
DESCRIPTION.Head depressed, snout is very short and broadly rounded; diameter of the eye
less than its distance from the mouth; nostril very large, pierced in a large circular or crescentic
nasal, bounded above by a supranasal which may be broken up; two scales between the nasal and
the rostral; the upper head scales are small, imbricate, keeled; there is an erect horn-like tubercle
above each eye, covered with several imbricate scales; 9–12 scales on a line between the horns; 15
scales around the eyes; one or two series of scales between the nasal and the rostral; scales on top
of the head small, imbricate, smooth on the snout, keeled behind; scales on top of the head small,
imbricate, smooth on the snout, keeled behind; 13 supralabials, three series of scales between the
eye and the labials; four infralabials in contact with the chin shields (genials). 1st pair of infralabi-
als larger than the others; a pair of large anterior genials, the scales posterior to them being much
smaller. Dorsal scales are strongly keeled in 21–23 rows; ventrals 134–138; the anal is entire, sub-
caudals, 35–38, divided (Schmidt's type had 21 dorsal scale rows, 134 ventrals, 35 subcaudals);
total length 890 mm, tail 110 mm, TL/T = 6.1–8.6.
Colors are grayish or brownish above with four series of large dark spots, the two median rows
sometimes confluent and forming cross bars; a dark streak on each side of the head from the eyes
to behind the gape; the under parts are whitish, dotted with dark and a lateral series of dark spots.
(Schmidt 1930:227–229; Gasperetti 1988:352–354, Table15).
DISTRIBUTION.Sinai, Israel, Jordan, Syria, Iraq, southwestern Iran (Fig. 13). The specimen
cited by Anderson (1963:472) from Binak, Iran, upon reexamination, is identified here as P. fieldi
(table 1).
NOTE.— Probably the main external morphological difference between P. persicus and P. fiel-
di is that the scale rows of P. persicus are all strongly keeled, whereas in P. fieldi several lateral
rows are nearly smooth and the outer rows are entirely without keels. Pseudocerastes fieldi also has
a significantly shorter tail. Sexual dimorphism has not been studied adequately in either taxon.
Bdolah (1986) demonstrated that the composition and properties of the venoms of P. fieldi and
persicus differ greatly. The venom of P. persicus showed the typical complexities of most viperid
snake venoms with potent hemorrhagic activity whereas that of P. fieldi demonstrated none. The
venom of P. persicus has a yellow pigment typical of most snake venoms and that of P. fieldi is
completely lacking in yellow flavin pigment and its isoelectric focusing profile is a simple one with
very few protein bands. Pseudocerastes persicus, on the other hand, had close to 30 protein bands
spanning a wide pH range. It was concluded that these differences indicate a long genetic separa-
tion of the two forms (Bdolah 1986:726) (Gasperetti 1988:352–353).
BOSTANCHI ET AL.: NEW SPECIES OF PSEUDOCERASTES FROM IRAN 445
The tail tip is often dark in adults of both taxa, suggesting the possibility of caudal luring.
(Gasperetti 1988:354, Table 15).
Pseudocerastes urarachnoides Bostanchi, Anderson, Kami, and Papenfuss, sp. nov.
Figures 1–12.
MATERIAL EXAMINED.HOLOTYPE: FMNH 170929 [], Iran: Ilam Province: 70 km SW Ilam [prob-
ably on road to Amirabad and Mehran], collected by Daniel R. Womochel and Anthony F. DeBlase, Second
Street Expedition to Iran, 27 August 1968 (see map, Fig. 13). PARATYPE: ZMGU [], 1300, Iran: Kermanshah
Province: 25 km south of Qasr-e-Shirin on road to Gilan-e Gharb, open level area in agricultural region, ca.
200 meters elevation, collected by Hamid Bostanchi, May 15, 2001 at about 0800 hrs (see map, Fig. 13).
DIAGNOSIS.— A Pseudocerastes with a short tail (TL/T = 9.65), few pairs of subcaudals (15 in
the known specimens), the distal pairs forming an oval knob-like structure; lateral dorsal caudal
scales projected to form elongate "appendages" alongside terminal knob. Several rows of lateral
dorsal scales are weakly keeled.
DESCRIPTION OF HOLOTYPE (FMNH 170929 , [Figs. 1–2, 8, 10]; head scales based on
paratype, ZMGU 1300 [?] [Figs. 5–7, 9]).The head of the holotype is severely damaged and
accurate counts of many of the head scales are not possible; consequently the description of miss-
446 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Fourth Series, Volume 57, No. 14
FIGURES 1–4. (1) Holotype, FMNH 170929, dorsal view (SCA photo). (2) Holotype, FMNH 170929, ventral view (SCA
photo). (3) Paratype, ZMGU 1300, dorsal view (James Parham photo). (4) Paratype, ZMGU 1300, ventral view (HGK
photo).
12
34
ing head characters is based on the paratype. Head depressed, snout
short and broadly rounded; the upper head scales small, imbricate,
keeled; there is an erect horn-like scale above the eye surrounded by
small imbricate scales; 16 scales on a line between the horns; (Fig. 6);
17 scales around the eyes; three series of scales between the eye and
the labials; two series of scales between the nasal and the rostral;
11/12 upper labials; 13/12 lower labials, 3–5 in contact with the chin
shields; anteriormost lower labials enlarged, in contact behind mental
and in contact with one pair of genials (fig. 7). Dorsal scales strongly
keeled, in 21 rows at level of 6th ventral, 23 rows at level of 73rd ven-
tral, 17 rows at level of 145th ventral; several lateral rows are weakly
keeled but the outer row is faintly, but distinctly keeled at midbody
(Fig. 10); ventrals 145; the anal is entire, subcaudals 15 pairs; total
length 531 mm, tail 55 mm, TL/T = 9.65.
Tail short, with only 15 pairs of subcaudals; dorsolateral caudal
scales adjoining subcaudals have elongated keels, progressively
longer from proximal to distal end of tail, keels of distal-most becom-
ing entire scale, the longest measuring 11.2 mm; tail terminates in an
elongate, oval, bulb-like structure, measuring 10.4 mm, apparently
formed of last pair of subcaudals, much enlarged, and a single
enlarged dorsal scale; lateral scales elongated to the extent that they
give the impression of arthropod legs (Figs. 8–9). The caudal verte-
brae extend well into this structure and are not deformed or modified
(Fig. 12).
Colors are grayish and brownish above with four series of large
dark spots, the two median rows sometimes confluent and forming
cross bars; a dark streak on each side of the head from the eyes to
behind the gape; the under parts are cream, with a lateral series of
dark spots (Figs. 1–4).
REMARKS ON PARATYPE.— ZMGU 1300 is a juvenile, apparent-
ly male (see Table 1 for counts and measurements). The caudal orna-
mentation is less well developed (Fig. 9), possibly a factor of age.
The new species most closely resembles Pseudocerastes persicus
in the dorsal scale characters which distinguish that taxon from P. fiel-
di, apart from the greatly shortened tail and the elaborate caudal
appendage, which set it apart from both. To the human observer this
caudal appendage greatly resembles an arthropod clinging to the tail
tip.
Unfortunately, neither specimen is in ideal condition, the head of
FMNH 170929 having been severely damaged and ZMGU 1300 hav-
ing partially rotted prior to proper preservation.
BOSTANCHI ET AL.: NEW SPECIES OF PSEUDOCERASTES FROM IRAN 447
FIGURES 5–10. (5) Lateral view of head of Paratype, ZMGU 1300 (James Parham
photo). (6) Dorsal view of head of paratype, ZMGU 1300 (HGK photo). (7) Ventral view
of head of paratype, ZMGU 1300 (James Parham photo). (8) Tail ornamentation of
Holotype, FMNH 170929 (SCA photo). (9) Tail ornamentation of paratype, ZMGU 1300
(HGK photo). (10) Ventrolateral scale rows keeled (holotype, FMNH 170929) (SCA
photo).
5
6
7
8
9
10
DERIVATION OF NAME.— From the Greek: ura = tail;
arachno = spider; ides = similar to.
DISCUSSION AND SPECULATION.This unusual snake
occupies a region that lies between the recorded ranges of P.
persicus and P. fieldii. Its discovery raises a number of inter-
esting questions and speculations. Only one of us (HB) has
seen this species alive, and while we speculate that the caudal
appendage functions as a caudal lure, this remains to be veri-
fied. While anthropomorphic interpretation should be avoided
where documentation is lacking, caudal luring is known for
several species of snakes, e.g., Bitis caudalis, Crotalus
cerastes, Sistrurus catenatus, Agkistrodon contortrix,
Acanthophis antarcticus, Acanthophis praelongus, Morelia
viridis, and others, and will probably be verified in many
species with differentially colored tails. Gasperetti (1988:354)
suggested that this might be the case for the previously known
species of Pseudocerastes. Neill (1980) reviewed the litera-
ture on caudal luring in juvenile snakes, especially crotalids
and boids, and suggested that the loss of contrasting tail color
with age marked the transition from ectothermic prey (e.g.,
amphibians, lizards, scorpions, centipedes) to mammals.
Greene (1992:111–112) added additional species to the list of
snakes exhibiting caudal luring and suggested that contrasting
tail coloring and luring behavior might be a synapomorphy in the Crotalinae, Viperidae, or larger
clade. This raises the question of the elaborate and sophisticated appearance of the caudal
appendage in our new species, as the waving or wriggling motion of a distinctively colored tail tip
seems perfectly adequate to attract lizard and anuran prey. We can only speculate that in the case of
the present species, the caudal lure serves to deceive a more specific kind of prey, such as shrews
or birds. Indeed, ZMGU 1300 contains an undigested, unidentified passerine bird in the stomach
(the feet protruding through the body wall). When sufficient specimens become available, addition-
al stomach contents should be examined and observations made of the behavior of captive individ-
448 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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11
FIGURES 11–12. Radiographs of holo-
type showing undeformed caudal vertebrae
extending into bulbus area of tail indicating
that the tail tip was neither damaged nor
regenerated. (Radiographs by Mark Zim-
merman, FMNH.)
12
FIGURE 13. Map showing distribution of the genus Pseudocerastes (lim-
its of distribution uncertain). (Map prepared by Karen Klitz.)
uals. Latifi (1991:131) states that the diet of P. persicus consists of lizards and mice; Khan (2002)
gives deserticolous lizards and arthropods as prey items.
As both available specimens were initially preserved in formalin, no tissue samples suitable for
molecular analysis are available. Should such become available in the future, by comparing DNA
from the three known species of Pseudocerastes, it may be possible to obtain a molecular clock esti-
mate of the time required to evolve such a structure.
The venom of Pseudocerastes persicus and P. fieldi differ in their chemical composition and in
their physiological action (Bdolah 1986:726), and it will be interesting to compare these with the
venom properties of the new species, if and when the opportunity arises.
ACKNOWLEDGMENTS
We thank Professor Martha Bowsky, University of the Pacific, who assisted with the etymolo-
gy of the species name; Karen Klitz, Museum of Vertebrate Zoology, University of California,
Berkeley, who prepared the map (Fig. 13); James Parham, University of California Museum of
Paleontology, who took photos of the paratype; Professor Richard Tenaza, University of the Pacific,
who helped in preparing two of the illustrations; and Mark Zimmerman, DVM, who took the radi-
ographs.
LITERATURE CITED
ANDERSON, STEVEN C. 1963. Amphibians and reptiles from Iran. Proceedings of the California Academy of
Sciences, ser. 4, 31(16):417–498, figs. 1–15.
BDOLAH, AVNER. 1986. Comparison of venoms from two subspecies of the false horned viper (Pseudocerastes
persicus). Toxicon 24(7):726–729, figs. 1–2.
BOULENGER, GEORGE A. 1896. Catalogue of the Snakes in the British Museum (Natural History). Volume III.,
BOSTANCHI ET AL.: NEW SPECIES OF PSEUDOCERASTES FROM IRAN 449
TABLE 1. Counts and measurements for specimens examined.7
P. urarachnoides P. persicus P. fieldi
FMNH ZMGU Latifi Latifi CAS CAS
170929 1300 (2000) (2000) 86633 159051
Sex ♀♂[?] ♀♀
Scale counts
Ventrals 145 146 144–163 142 153 158
Subcaudals (pairs) 15 15 38–50 46 41 38
Anal plate single single single single single single
Dorsal scales at midbody 21 23 23–25 21 23 23
Scales around eye 818/17 15–20 16 20 18
Interorbital scales ~9–10 16 10–15 10 15 12
Scales between eyes and
upper labials 3 3 2–4 3 4 4
Upper labials ~8 11/12 11–14 14/13 11/13
Lower labials 13/12 15/14 13–17 18/17 13/13
Measurements
Snout-vent (mm) 531 386 1160 750 515 279
Tail (mm) 55 46 130 80 75 39
7Additional counts and measurements for P. persicus and P. fieldi are given by Gasperetti (1988:354, table
15).
8Head damaged
Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae and Viperidae.
Trustees of the British Museum (Natural History), London, UK. xiv + 727 + 19 pp., pls. 1–25.
DUMÉRIL, ANDRÉ MARIE CONSTANT, GABRIEL BIBRON, AND AUGUSTE HENRI ANDRÉ DUMÉRIL. 1854. Erpétologie
Général ou Histoire Naturelle Complète des Reptiles, vol. 7 (part 2). Librairie Encyclopédique de Roret,
Paris (1), (1), xii, 781–1536 pp., 2 folding tables, pls.83–84, pls. 75–84.
GASPERETTI, JOHN. 1988. Snakes of Arabia. Fauna of Saudi Arabia 9:169–450, 135 figs., 17 tables.
GREENE, HARRY W. 1992. The ecological and behavioral context for pitviper evolution. Pages 107–118 in J.
A. Campbell and E. D. Brodie, Jr. eds. Biology of the Pitvipers. Selva Press, Tyler, Texas, USA.
KHAN, MUHAMMAD S. 2002. A Guide to the Snakes of Pakistan. English Edition. Edition Chimaira, Frankfurt
am Main, Germany. 265 pp., 155 pls., 20 distribution maps.
LATIFI, MAHMUD. 1991. The Snakes of Iran, English ed. Contributions in Herpetology, 7. Society for the Study
of Amphibians and Reptiles, Oxford, Ohio, USA. viii + 159 pp. 24 text-figs., 25 col. pls., 3 tables.
LATIFI, MAHMUD. 2000. The Snakes of Iran. 3rd Persian ed. Tehran, Iran. 478 pp. (In Farsi)
LEVITON, ALAN E., STEVEN C. ANDERSON, KRAIG K. ADLER, AND SHERMAN A. MINTON, JR. 1992. Handbook to
Middle East Amphibians and Reptiles. Society for the Study of Amphibians and Reptiles, Oxford, Ohio,
USA. 252 pp., unnumbered text-figs., 21 figs., 32 col. pls. 1 b&w pl., 1 table.
MINTON, SHERMAN A., JR. 1966. AContribution to the Herpetology of West Pakistan. Bulletin of the American
Museum of Natural History 134(2):29–184, pls. 9–36.
NEILL, WILFRED T. 1960. The caudal lure of various juvenile snakes. Quarterly Journal of the Florida
Academy of Sciences 23(3):173–200.
SCHMIDT, KARL P. 1930. Reptiles of the Marshall Field North Arabian Desert Expedition, 1927–1928. Field
Museum of Natural History, Zoological Series 12(13):221–230, figs. 1 [map] 2, pl. 2., figs. 1–2.
SMITH, MALCOLM A. 1943. Reptilia and Amphibia, vol. 3, Serpentes. The Fauna of British India, Ceylon and
Burma, including the Whole of the Indo-Chinese Subregion. Taylor and Francis, London, UK. xii + 583 +
5 pp., 166 figs., 1 fold-out map.
450 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Copyright © 2006 by the California Academy of Sciences
San Francisco, California, U.S.A.
... These gaps are particularly associated with developing regions such as the Middle East, which has been subject to extended periods of political instability and ongoing tension. The data-deficient nature of the Middle Eastern herpeto-faunal literature is exemplified by the relatively recent description of a third species of viperid snake from the genus Pseudocerastes (Boulenger, 1896), the spider-tailed viper Pseudocerastes urarachnoides (Viperidae: Viperinae) (Bostanchi, Anderson, Kami, and Papenfuss, 2006) from western Iran [2,3]. ...
... These gaps are particularly associated with developing regions such as the Middle East, which has been subject to extended periods of political instability and ongoing tension. The data-deficient nature of the Middle Eastern herpeto-faunal literature is exemplified by the relatively recent description of a third species of viperid snake from the genus Pseudocerastes (Boulenger, 1896), the spider-tailed viper Pseudocerastes urarachnoides (Viperidae: Viperinae) (Bostanchi, Anderson, Kami, and Papenfuss, 2006) from western Iran [2,3]. ...
... persicus and P. urarachnoides) [4,7]. Distribution data on these species is incomplete, though they are mostly found throughout the Middle East: E. macmahonii in the Baluchistan (covering the Iran-Afghanistan-Pakistan border) and the Thar (northwest India) deserts [8]; P. urarachnoides in western Iran and eastern Iraq [9][10][11]; P. persicus from northeast Iraq to western Pakistan (including southern Afghanistan and Iran), as well as some isolated populations in Oman and the United Arab Emirates (UAE) [12,13]; and P. fieldi from northern Saudi Arabia, through the south of Israel, Jordan, and Syria, and across Iraq and southern Iran [3,14]. There appears to be limited range overlap between these species, though sympatry exists between P. fieldi and P. urarachnoides and between P. urarachnoides and P. persicus in the northern and southern edges of P. urarachnoides' range in western Iran, respectively [13,15]. ...
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Snakes of the genera Pseudocerastes and Eristicophis (Viperidae: Viperinae) are known as the desert vipers due to their association with the arid environments of the Middle East. These species have received limited research attention and little is known about their venom or ecology. In this study, a comprehensive analysis of desert viper venoms was conducted by visualising the venom proteomes via gel electrophoresis and assessing the crude venoms for their cytotoxic, haemotoxic, and neurotoxic properties. Plasmas sourced from human, toad, and chicken were used as models to assess possible prey-linked venom activity. The venoms demonstrated substantial divergence in composition and bioactivity across all experiments. Pseudocerastes urarachnoides venom activated human coagulation factors X and prothrombin and demonstrated potent procoagulant activity in human, toad, and chicken plasmas, in stark contrast to the potent neurotoxic venom of P. fieldi. The venom of E. macmahonii also induced coagulation, though this did not appear to be via the activation of factor X or prothrombin. The coagulant properties of P. fieldi and P. persicus venoms varied among plasmas, demonstrating strong anticoagulant activity in the amphibian and human plasmas but no significant effect in that of bird. This is conjectured to reflect prey-specific toxin activity, though further ecological studies are required to confirm any dietary associations. This study reinforces the notion that phylogenetic relatedness of snakes cannot readily predict venom protein composition or function. The significant venom variation between these species raises serious concerns regarding antivenom paraspecificity. Future assessment of antivenom is crucial.
... Such a list would provide a starting point for understanding the evolution of sidewinding as well as its biomechanical underpinnings. Here, I review the current knowledge 50 of sidewinding, provide an extensive list of species known to sidewind to varying degrees, and use this list to draw some inferences about sidewinding. ...
... He later used electromyography to demonstrate that both forms of locomotion involve bilateral activity of the spinalis muscle (Jayne 1988). Finally, he emphasized the continuous propagation of waves in these two types of 50 locomotion, in contrast to concertina locomotion. Overall, the body of evidence favors the possibility that sidewinding derives from lateral undulation (Gray 1946(Gray , 1968Brain 1960;Jayne 1986Jayne , 1988. ...
... Like many other 45 aspects of sidewinding, frequency relates to the conditions eliciting the behavior: a snake trying to make a quick escape from a predator needs to use a higher frequency than does a snake on an unhurried, longdistance trek (personal observation). Although indi- 50 vidual snakes do not always use the same frequency, different sidewinding species (both specialized and facultative) could tend toward higher or lower frequencies. Muscular ability could set a maximum limit on frequency, but higher frequency would not 55 necessarily indicate more proficient sidewinding. ...
Article
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Specialist species often possess adaptations that strongly distinguish them from their relatives, obscuring the transitional steps leading to specialization. Sidewinding snakes represent an example of locomotor specialization in an elongate, limbless terrestrial vertebrate. We typically think of sidewinding as a gait that only a handful of very specialized snake species perform, mostly vipers from sandy desert environments. Some of these desert-dwelling vipers are so specialized that they only rarely use more common types of locomotion. However, some non-viper species sidewind facultatively in particular circumstances, and a few may regularly sidewind under natural conditions. Numerous accounts report facultative sidewinding in species that more typically perform other types of locomotion. I have compiled these accounts, uncovering evidence that dozens of species perform sidewinding with varying proficiency under a variety of conditions. These facultative sidewinders can reveal insight into the evolution and biomechanics of sidewinding, and they provide ample opportunities for future study.
... There are two other species in this genus, Field's horned viper (P. fieldi) which is distributed in Egypt, Sinai, Israel, Jordan, Saudi Arabia, Syria and Iraq (Bostanchi et al., 2006;Mallow et al., 2003); and the Iranian spider-tailed viper (P. urarachnoides) which occurs in Western Iran (Bostanchi et al., 2006;Fathinia et al., 2018Fathinia et al., , 2020, and in Wasit Province, in the extreme East of Iraq (Al-Sheikhly et al., 2019). ...
... fieldi) which is distributed in Egypt, Sinai, Israel, Jordan, Saudi Arabia, Syria and Iraq (Bostanchi et al., 2006;Mallow et al., 2003); and the Iranian spider-tailed viper (P. urarachnoides) which occurs in Western Iran (Bostanchi et al., 2006;Fathinia et al., 2018Fathinia et al., , 2020, and in Wasit Province, in the extreme East of Iraq (Al-Sheikhly et al., 2019). ...
Article
Despite the wide distribution of the Persian false-horned viper (Pseudocerastes persicus) in the Middle East, few identified bites have been reported. A 33-year-old herpetologist bitten on the hand by Pseudocerastes persicus in Kerman Province, Southeastern Iran, developed local pain and extensive swelling with mild non-specific systemic symptoms and minimal laboratory evidence of systemic envenoming.
... Yet, other forms of mimicry involve apparent action, such as the hornet-like body movement and buzzing in the hornet moth, the apparent struggle of the nonexistent "ant" with the "caterpillar" in lobster moth larvae or the illusion of ant-to-ant helpfulness in the jumping spider. Similarly, the spider-tailed horned viper (Pseudocerastes urarachnoides) of Iran enacts a sequence of movements of its tail tip shaped like a spider as if a careless spider was crawling on a rock (caudal luring-compare Bostanchi et al. 2006;Fathinia, Rastegar-Pouyani 2010;Fathinia et al. 2015). This attracts the spider's bird predator who flies in to catch the spider (i.e., to become its predator). ...
... This is a case of "THE PREY IS THE PREDATOR," a biometaphor that protects the moths by making them look like the object of their fear, as the spiders are less likely to attack members of their own species. "THE PRE-DATOR IS FOOD" (or "THE PREDATOR IS THE PREY") is the metaphor embodied by the Iranian spider tailed viper (Pseudocerastes urarachnoides) whose tail ending looks and moves like a spider, attracting bird predators who, in turn, become the viper's prey (Bostanchi et al. 2006). Thus, not only does the actual predator mimic prey with its tail (caudal lure, including movement, re-enacting a miniature event that is not what it seems to be) but by attacking the bird attacker it turns the other predator into its prey. ...
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Can Cognitive Metaphor Theory (CMT) be applied productively to the study of mimicry in zoosemiotics and ethology? In this theoretical comparison of selected case studies, I would like to propose that biological mimicry is a type of biosemiotic metaphor. At least two major parallels between cognitive metaphors in human cognition and mimicry among animals justify viewing the two phenomena as isomorphic. First—from the semiotic point of view—the argument is that both metaphor and mimicry are cases of semiotic transfer (etymologically: metaphor) of the identity / sign of the source onto the perceived identity / sign of the target. This identity transfer, in turn, triggers appropriate changes in the response (behavior) of the surrounding (human or animal) interpreters (e.g. predators). Semiotically, the mimicry turns the body of its bearer into a sign of something else, resulting in the interpreters’ (e.g. predators’) perception of species X as species Y—hence, a type of embodied sign and cognitive metaphor. Second, ecologically, a species occupying one niche (e.g. a moth: non-venomous, herbivorous primary consumer) is perceived and identified as an occupant of a different niche (e.g. a hornet: venomous, omnivorous predator). Thus, a potential predator’s Umwelt is affected by its perceiving a hornet moth as “a hornet” where there is, in fact, a moth, and its response to this stimulus will not be predation but avoidance. In terms of CMT, we could call this a biosemiotic metaphor (bio-metaphor), e.g. “A MOTH IS A HORNET” or “PREY IS A PREDATOR”. Further correspondences between mimicry and metaphor include the fact that this bio-metaphorical identification by mimicry does not typically require a “perfect” resemblance between the source and the target sign (or species); this seems to correspond to the prototype categorization in CMT where categories are “open-ended” and only a partial similarity is sufficient for metaphorical identification (compare Lakoff, Johnson 1980; Rosch 1983). Such an identification of mimicry as metaphor could be based on Prodi’s argument that “hermeneutics is not a late product of culture, but the same elementary movement of life that is born because something obscurely interprets something else” (Eco 2018: 350; Kull 2018, 352—364). Inasmuch as animal Umwelten are interconnected inter alia by this natural hermeneutics, the trans-disciplinary approach to the study eco-zoosemiotic interpretants on the basis of metaphor-mimicry isomorphism could open new opportunities in comparative studies of semiosis in human and animal cognition and interactions.
... To date, at least 72 endemic reptilian species have been reported from Iran (Chefaoui, Hosseinzadeh, Mashayekhi, Safaei-Mahroo, & Kazemi, 2018;Fathinia, Rastegar-Pouyani, & Shafaeipour, 2019;Gholamifard, Rastegar-Pouyani, & Rastegar-Pouyani, 2019;Torki, 2019). As one of the restricted-range species, the Spider-tailed Viper, Pseudocerastes urarachnoides Bostanchi, Anderson, Kami & Papenfuss, 2006, is reported to have a limited distribution along the Zagros mountains in western areas of Iran (Fathinia, Anderson, Rastegar-Pouyani, Jahani, & Mohamadi, 2009) and the eastern parts of Iraq (Al-Sheikhly, Al-Barazengy, & Al-Haideri, 2019). ...
... Study area. A total of 99 occurrence records of P. urarachnoides were obtained from our numerous fieldwork conducted between 2008 to 2019 and from the literature (Al-Sheikhly et al., 2019;Bostanchi et al., 2006;Fathinia et al., 2009;Fathinia & Rastegar-Pouyani, 2010;Fathinia, Rastegar-Pouyani, Rastegar-Pouyani, Todehdehghan, & Mansouri 2017). Climatic variables. ...
Article
To date, at least 72 endemic reptilian species have reported from Iran including the Spider-tailed Viper (Pseudocerastes urarachnoides), which has a very limited, narrow distribution, and occurs in areas of western Iran and eastern Iraq. The potential distribution of Pseudocerastes urarachnoides in Iran for the present, the past (mid- Holocene and last glacial maximum), and the future (2100) was predicted by Maximum Entropy (MaxEnt) modelling using 99 occurrence records as well as 19 environmental variables derived from climate databases. For all projected potential distributions, the principal components 1 (explained primarily by thermal variables) and 2 (explained primarily by precipitation variables) contributed more than 80% collectively in all MaxEnt models. The extreme eastern distribution range of P. urarachnoides corresponds to the western slopes of the Zagros Mountains within the Iraq territories. According to the projections, the current distribution area is smaller than in the mid- Holocene but larger than the last glacial maximum and three out of four scenarios of the future (2100). Future projections dramatically displace the suitable habitat, leading to a mismatch between the current and future habitat range of the Spider-tailed Viper.
... Although the most common recent ancestor (MRCA) for P. persicus and P. urarachnoides dates back to˜8 mya (Fathinia et al., 2018), P. persicus is morphologically more similar to P. fieldi (MRCA;˜12 mya) than to P. urarachnoides . Some morphological characteristics that can be regarded as autapomorphies for P. urarachnoides are the unique caudal structure and rugosity of scales in comparison to its congeners, which deeply separate it from its two congeners (Bostanchi et al., 2006;Fathinia and Rastegar-Pouyani, 2010). Scale microornamentation revealed that the microstructure of scales in P. fieldi and P. persicus are more similar to each other than to P. urarachnoides , coincides with morphological characteristics, but contrary to molecular relationships within this genus. ...
Preprint
Ecomorphological studies are aimed to find out the relation between the morphology of organisms and their ecology. Many studies on reptile scale microornamentation indicate that it has important functional values. In this study microornamentation and light reflection of scales’ surface among six viperid and two colubrid snake species in relation to their habitat were examined. To compare microornamentation and light reflection analyses, skin specimens were prepared and analyzed using scanning electron microscopy and FluoVision Imaging System and spectrometer, respectively. The results showed that snake species inhabiting similar habitats had different microornamentation and vice versa. Likely the scale microstructures are more influenced by phylogenetic relationships than by the environment. In examining the scales’ reflection, different species and different body parts reflect various wavelengths of visible light, which relates to ecological condition of them.
... However, research on the direct link between tail use and prey capture is limited. Previous studies found tail movements are essential to the rapid high-speed maneuvers used by cheetahs to capture prey (Wilson et al. 2013;Patel et al. 2016), directly used to capture insects mid-flight in bats (Kalko 1995), and used as lures to attract or distract prey in a variety of snake species (Neill 1960), most dramatically the spider-tailed viper, Pseudocerastes urarachnoides (Bostanchi et al. 2006). However, these specialized uses do little to elucidate the mechanical role that the tail plays in prey capture strikes. ...
Article
Tails are versatile structures with diverse forms and functions across vertebrates. They are involved in almost all behaviors critical to survival including locomotion, feeding, and predator avoidance. Although the tail’s role in locomotion and stability has been widely studied, its role in prey capture is relatively unknown. Lizards are an ideal system to examine the tail’s impact on prey capture as most are capable of autotomizing, or dropping, their tail in response to predation and intraspecific competition. Tail autotomy can lower reproduction, decrease locomotor performance, impart instability during jumping, and decrease social status. Desert banded geckos (Coleonyx variegatus) frequently capture evasive prey in nature and appear to use their tail during strikes. However, it is unclear if these tail movements are important for the strike itself, or if they simply draw attention to that part of the body. We used high-speed 3D videography to quantify prey capture performance and kinematics of C. variegatus striking at crickets before and after total caudal autotomy. Trials were conducted within two hours of autotomy and then repeatedly over a two-week period. Overall prey capture success was unaffected by caudal autotomy. However, maximum strike velocity decreased significantly after autotomy, highlighting the importance of the tail during prey capture. Strike kinematics were altered after autotomy in several ways, including geckos adopting a more sprawled posture. Maximum pectoral girdle and mid-back height were significantly lower during post-autotomy strikes, whereas maximum pelvic girdle height was unaffected. However, individual variation was considerable. This downward pitching of the body after tail loss suggests that the tail is necessary for counterbalancing the anterior portion of the body and resisting the rotational inertia incurred after pushing off with the hindlimbs. Utilizing autotomy to test tail function in prey capture can provide valuable insight into how the tail is used in terrestrial predation across a wide variety of species and ecological niches.
... It was the impression of the observers that the distinctive black, wormlike tip of this viper's tail was very likely to be used as a lure to attract prey, as has been speculated by others (e.g., Cunningham 2002). In addition to the closely similar appearance of the tail tip to images of other species known to use tail luring (e.g., Dressler, showing Peringuey's Adder), that inference is further supported both by the coiling posture adopted by the snake, which left the tail positioned to the fore, and by the recent recognition of an even more elaborate caudal lure in a close Iranian congener, the Spider-Tailed Horned Viper P. urarachnoides (Bostanchi et al. 2006 The only other snake so far confirmed from the Olive Highlands is the Sand Snake Psammophis schokari, which was recorded once at 700-800 metres and suspected at ca. 1000 metres on Jebel Qitab, among vegetated boulders below cliffs along the summit ridge. A "racer" was also recorded on the summit ridge of Jebel Sfai, which was probably either P. schokari or Platyceps rhodorachis. ...
Article
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The area that is here called the Olive Highlands is a remote and little known region of high ridges (ca. 1000 metres) within the Hajar Mountains of the United Arab Emirates, south-west of Fujairah city. It represents an ecologically unique 'island' of biodiversity, characterised by a distinctive high elevation flora and fauna including a number of plant species (e.g., the wild olive tree Olea europaea, the large shrubs Ehretia obtusifolia and Grewia tenax, and the cactus-like, yellow-flowering milkweed Desmidorchis flavus) as well as at least one animal species (the Arabian Grizzled Skipper butterfly Spialia mangana) that are not found elsewhere in the UAE, and other plant and animal species not otherwise found outside the Ru'us al-Jibal (the mountains of the Musandam peninsula). On the basis of what was known about their flora, the Olive Highlands were identified as a discrete biological habitat and were highlighted for attention by a recent biodiversity assessment for national conservation planning. Subsequently, the area has been confirmed to be a refuge for rare fauna as well, including the Arabian Grizzled Skipper and the Persian Horned Viper Pseudocerastes persicus, for each of which there are no prior records from the Hajar Mountains of the UAE. The Olive Highlands also host an unusual colour morph of the Blue Pansy butterfly Junonia orythia and a large population of the Bar-Tailed Semaphore Gecko Pristurus celerrimus. Most if not all of these distinctive plant and animal species probably represent relicts of more widespread populations that flourished in the more mesic climate of an earlier time, perhaps as recently as ca. 6,000-10,000 years ago during the so-called climatic optimum. Protection of the Olive Highlands from a conservation standpoint is obviously desirable in order to preserve their unique contribution to UAE and regional biodiversity. Protection should be straightforward as a technical matter. The area remains largely unknown and unvisited, relatively inaccessible and largely undisturbed. It does not require active intervention or management; it needs only to continue to be left alone – to be earmarked for preservation and closed to development activities and other molestation. The area in question encompasses parts of three of the seven Emirates – Fujairah, Ras al-Khaimah and Sharjah. This means that effective preservation ultimately requires cooperation among the concerned authorities in the several Emirates, still an unproven model in the UAE, so the Olive Highlands present an opportunity that is unique not only in terms of biodiversity but in a political sense as well.
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Amphibia: Anura: Bufonidae: Duttaphrynus arabicus (Heyden, 1827); Reptilia: Squamata: Sauria: Agamidae: Pseudotrapelus sinaitus sinaitus (Heyden, 1827); Gekkonidae: Asaccus montanus Gardner, 1994, Bunopus spatalurus hajarensis Arnold, 1980, Hemidactylus n. sp. aff. H. persicus, Pristurus celerrimus Arnold, 1977, P. gasperetti gallagheri Arnold, 1986, P. rupestris rupestris Blanford, 1874; Lacertidae: Omanosaura cyanura (Arnold, 1972) O. jayakari (Boulenger, 1887a); Scincidae: Ablepharus pannonicus Fitzinger, 1823; Trachylepis tessellata Anderson, 1895; Serpentes: Colubridae: Platyceps rhodorachis (Jan, 1863), Psammophis schokari Forskål, 1775; Arabian Peninsula: Sultanate of Oman: Hajar al-Gharbi: Jebel Akhdar: Saiq Plateau: new records; range extensions; field observations.
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A systematic list and identification keys to the families, genera and species are given, with a taxonomic description and discussion of size, habits and distribution. Sketch maps showing the distribution in Arabia as well as general distribution are included. Some of the species are illustrated in colour from live animals, some of which show considerable colour variation within a species. A physiographic sketch of the peninsula and the zoogeography in the context of the Arabian Peninsula and its neighbouring countries is outlined. A separate chapter deals with species conservation. -from Author
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Typescript. Thesis (M.A.)--San Francisco State College. Includes bibliographical references (leaves 110-129).
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Gel-isoelectric focusing of Pseudocerastes persicus fieldi venom shows a simple pattern with very few protein bands, while the venom of P. p. persicus reveals close to 30 protein bands. Molecular sieve chromatography of P. p. persicus venom shows a typical elution profile of a viperid snake, with hemorrhagic activity and L-amino acid oxidase activity confined to the high molecular weight peak of proteins. In the venom of P. p. fieldi, which has no hemorrhagin or L-amino acid oxidase, the high molecular weight fraction of proteins is practically missing.
The Snakes of Iran. 3rdPersian ed. Tehran, Iran. 478 pp Handbook to Middle East Amphibians and Reptiles pp., unnumbered text-figs., 21 figs
  • Mahmud Leviton Latifi
  • C Anderson
  • Sherman A Minton
LATIFI, MAHMUD. 2000. The Snakes of Iran. 3rdPersian ed. Tehran, Iran. 478 pp. (In Farsi) LEVITON, ALAN E., STEVEN C. ANDERSON, KRAIG K. ADLER, AND SHERMAN A. MINTON, JR. 1992. Handbook to Middle East Amphibians and Reptiles. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, USA. 252 pp., unnumbered text-figs., 21 figs., 32 col. pls. 1 b&w pl., 1 table
The Snakes of Iran viii + 159 pp. 24 text-figs., 25 col. pls
  • Mahmud Latifi
LATIFI, MAHMUD. 1991. The Snakes of Iran, English ed. Contributions in Herpetology, 7. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, USA. viii + 159 pp. 24 text-figs., 25 col. pls., 3 tables
Handbook to Middle East Amphibians and Reptiles
  • Alan E Leviton
  • Steven C Anderson
  • Kraig K Adler
  • Sherman A
  • J R Minton
LEVITON, ALAN E., STEVEN C. ANDERSON, KRAIG K. ADLER, AND SHERMAN A. MINTON, JR. 1992. Handbook to Middle East Amphibians and Reptiles. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, USA. 252 pp., unnumbered text-figs., 21 figs., 32 col. pls. 1 b&w pl., 1 table.