• TABLE OF CONTENTS
HIDE
 Front Cover
 Copyright
 Abstract
 Table of Contents
 Introduction
 Material
 Characters
 Sample areas
 Variation
 Sexual variation
 Summary and discussion
 Zoogeographical implications
 Possible selective forces
 Literature cited
 Tables
 Appendix 1
 Back Cover






Group Title: Bulletin of the Florida Museum of Natural History
Title: Studies on Pakistan reptiles, part 1 : the genus echus ( viperidae )
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00095827/00001
 Material Information
Title: Studies on Pakistan reptiles, part 1 : the genus echus ( viperidae )
Series Title: Bulletin - Florida Museum of Natural History ; volume 35, number 5
Physical Description: v. : ill., maps ; 23 cm.
Language: English
Creator: Auffenberg, Walter
Rehman, Hafeezur
Donor: unknown ( endowment )
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1991
Copyright Date: 1991
 Subjects
Subject: Reptiles -- Geographical distribution -- Pakistan   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Spatial Coverage: Pakistan
 Notes
Bibliography: Includes bibliographical references (pt. 1., p. 308-309).
General Note: Cover title.
General Note: Abstracts in English and Spanish.
Statement of Responsibility: Walter Auffenberg and Hafeezur Rehman.
 Record Information
Bibliographic ID: UF00095827
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 24638589

Table of Contents
    Front Cover
        Page 261
    Copyright
        Page 262
    Abstract
        Page 263
    Table of Contents
        Page 264
    Introduction
        Page 265
    Material
        Page 266
        Page 267
    Characters
        Page 268
        Page 269
        Page 270
        Page 271
    Sample areas
        Page 272
        Page 273
        Page 274
        Page 275
        Page 276
    Variation
        Page 277
        Page 278
        Page 279
        Page 280
        Page 281
        Page 282
        Page 283
        Page 284
        Page 285
        Page 286
        Page 287
        Page 288
        Page 289
        Page 290
        Page 291
        Page 292
        Page 293
    Sexual variation
        Page 294
    Summary and discussion
        Page 295
        Page 296
        Page 297
        Page 298
    Zoogeographical implications
        Page 299
        Page 300
        Page 301
        Page 302
        Page 303
    Possible selective forces
        Page 304
        Page 305
        Page 306
        Page 307
    Literature cited
        Page 308
    Tables
        Page 309
        Page 310
        Page 311
    Appendix 1
        Page 312
        Page 313
        Page 314
        Page 315
    Back Cover
        Page 316
Full Text





of the



FLOR1EA
MUSEUM OF
NATURAL HISTORY


STUDIES ON PAKISTAN REPTILES
PT. 1. THE GENUS ECHIS (VIPERIDAE)
Walter Auffenberg and Hafeezur Rehman



Biological Sciences, Volume 35, Number 5, pp. 263-314 1991


UNIVERSITY OF FLORIDA


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STUDIES ON PAKISTAN REPTILES
PT. 1. THE GENUS ECHIS (VIPERIDAE)



Walter Auffenberg and Hafeezur Rehman*




ABSTRACT



This is the first of a series of studies intended to analyze the geographic patterns in the
meristic characters of Pakistan reptiles. This part concerns the saw-scaled vipers, Echis carinatus.
The scales analyzed are numbers of ventrals, subcaudals, dorsal rows, rows of oblique dorsals,
gulars, suboculars and supraoculars; color patterns include number of dorsal spots, ventral
pattern, infralabials, lateral body and head. Maps of geographic variation in the means of each of
these characters are provided and the patterns of variation within Pakistan are discussed.
Each of the characters studied tends to have a different geographic distribution of mean
values. However, one recurrent pattern is a rapid change in character state between Iranian
Plateau populations (Chagai area, Pakistan) and those of the Quetta highland area. Coastal
populations are also rather distinctive, but are generally similar to those in the Indus River Plain.
Himalayan foothill populations are often similar to those of coastal areas, suggesting that these
geographical peripheral populations are also peripheral in regard to genetic adaptation. In the
Indus River Delta, populations can be subdivided into several distinct subgroups. The resulting
mosaic of character states is attributed to historic factors concerned with changing positions of
the deltaic distributaries during geologic time. Finally, the population in the Cholistan-Thar
Desert at the Indo-Pakistan border is distinctive in both color and scalation. These differences
are probably adaptive responses to local arid conditions. Several character states of this
population and that in the Chagai Desert of northwest Pakistan are convergent.
Of those climatic factors believed important within the adaptive mileau of this species,
mean daily maximum and minimum temperatures during June and July (when they are most
active) are considered most important. Some of the population differences probably reflect
adaptations to water and heat loss.
Contrary to recently published opinion, we conclude there is only one species of saw-scaled
viper in Pakistan; its scientific name is Echis carinatus. Within Pakistan we recognize three
distinct geographic races--E.c. multisquamatus Cherlin, E. c. sochureki Stemmler and E. c. astolae
Mertens.
The geographic distribution of characters analyzed shows that the major barrier to gene
flow between E. c. sochureki and E. c. multisquamatus is the central north-south massif (Sulaiman
Range, et al.). For E. c. sochureki and E. c. astolae it is the Arabian Sea, and for E. c. sochureki
and E. c. carinatus (Indian Peninsula) it is largely a matter of overall distance. In general, the
race E. c. sochureki is intermediate between E. c. multisquamatus and E. c. carinatus.

* Dr. Walter Auffenberg is Distinguished Service Curator, Florida Museum of Natural History, University of Florida,
Gainesville, FL, 32611, USA. Mr. Hafeezur Rehman is Biologist, Herpetology Section, Zoological Survey Department, BI. 61,
Pakistan Secretariat, Karachi 1, Pakistan.


Auffenberg, W., and H. Rehman. 1991. Studies on Pakistan Reptiles. Pt. 1. The Genus Echis
(Viperidae). Bull. Florida Mus. Nat. Hist., Biol. Sci. 35(5):263-314.







264 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)

RESUME

Este es el primero de una series de studios que pretenden analizar los patrons geogrAficos
en los caractees meristicos de los reptiles de Pakistan. Esta parte concierne a las viboras de
escamas aserradas (saw-scaled) Echis carinatus. Se analiza el nimero de escamas ventrales,
subcaudales, filas de escamas dorsales, filas de oblicuas dorsales, gulares, suboculares y
supraoculares; los patrons de color incluyen nilmero de manchas dorsales, patr6n ventral,
infralabiales, lados del cuerpo y cabeza. Mapas de variaci6n geografica en los promedios de cada
uno de estos caracteres son provistos y se discuten los patrons de variaci6n dentro de Pakistin.
Cada uno de los caracteres estudiados tiende a mostrar una distinta distribuci6n geogrAfica
de sus valores promedio. Sin embargo, un patr6n recurrente es un cambio ripido en la condici6n
de un character entire las poblaciones de la placa irani (Chagai, Pakistan) y aquellas de la regi6n
alta de Quetta. Las poblaciones de la costa son tambien distintas, pero generalmente similares a
las que se encuentran en la planicie del rio Indo (Indus River). Las poblaciones que ocupan el
pie de montafa de los Himalaya son a menudo similares a las costeras, lo cual sugiere que estas
poblaciones geograficas perifdricas son tambidn perifdricas en cuanto a sus adaptaciones
gendticas. En el delta del rio Indo, las poblaciones pueden subdividirse en various subgrupos. El
mosaico de condiciones de caracteres resultante es atribuido a factors hist6ricos relacionados al
cambio de posci6n de los distributarios del delta en el tiempo geol6gico. Finalmente, la
poblaci6n del desierto de Cholistan-Tar en el limited indo-pakistani es distinta en color y
escamaci6n. Estas diferencias son probablemente respuestas adaptativas a condiciones aridas
locales. Varias de estas condiciones de character de esta poblaci6n y la del desierto Chagai en el
Noroeste de Pakistan son convergentes.
Entre los factors climAticos que se creen importantes dentro el medio ambiente de esta
especie, la temperatures medias maxima y minima durante Junio y Julio (cuando la especie es
mas active) se consideran los mas importantes. Algunas de las diferencias entire poblaciones
probablemente reflejen adaptaciones ala perdida de agua y calor.
En contrast con opinions recientemente publicadas, nosotros concluimos que s6lo hay
una especie de vibora de escamas aserradas en Pakistan; su nombre cientifico es Echis carinatus.
Dentro Pakistan reconocemos tres razas geogrAficas--E. c. multisquamatus Cherlin, E. c.
sochureki Stemmler y E. c. astolae Mertens.
La distribuci6n geografica de los caracteres analizados muestra que la mayor barrera para
el flujo gen6tico entire E. c. sochureki y E. c. multisquamatus es el macizo central Norte-Sur
(Sulaiman Range, et al.). Para E. c. sochureki y E. c. astolae es el Mar ArBbigo, y para E. c.
sochureki and E. c. carinatus (Peninsula de la India) es por demis una cuesti6n de distancia. En
general, la raza E. c. sochureki es intermedia entire E. c. multisquamatus yE. c. carinatus.


TABLE OF CONTENTS


Introduction........................................................................................................................................... 265
M aterial.................................................................................................................................................. 266
Sources........................................................................................................................................ 266
L o calities..................................................................................................................................... 266
C haracters.............................................................................................................................................. 268
Characters used previously............................................................................................... 268
Characters used here ......................................................................................................... 268
Sam ple A areas ................................................................................................................................. 272
V ariatio n ................................................................................................................................................ 277
G geographic variation ......................................................................................................... 277
Sexual variation ......................................................................................................................... 294
Sum m ary and D discussion ............................................................................................................. 295
System atics................................................................................................................................. 295
Zoo geographical implications ................................................................................................. 299
Possible selective forces ........................................................................................................... 304
L literature C ited .................................................................................................................................... 308
T ab les..................................................................................................................................................... 309
A o end ix ................................................................................................................................................ 312






AUFFENBERG & REHMAN: THE GENUS ECHIS


INTRODUCTION


For many years all Pakistan Echis populations were considered a single
species (i.e., E. carinatus (Schneider) 1801; type loc. nr. Madras, India).
Constable (1949) suggested that the Pakistan (and other) populations should
be referred to E. carinatus pyramidium (Geoffroy 1827; type loc. Egypt).
Stemmler (1969) modified this arrangement by placing all Pakistan (and some
other) populations in E. carinatus sochureki (Stemmler 1969, type loc. Ban
Kushdil Khan, nr. Pishin, Baluchistan, Pakistan). He also demonstrated that
within the geographic confines of his new race, the populations from southern
Russia were larger and possessed a distinctive scalation when compared to
those from more southern parts of the range, but he did not describe them as a
separate subspecies. In 1969, Mertens described E. carinatus astolae from the
Mekkran Coast of Pakistan (type loc. Astola Island, Baluchistan). In 1981,
Cherlin described E. multisquamatus from Russian Turkistan (type loc.
Bayram Ali, Turkmen SSR). He referred some of his material from Chagai
District, northern Baluchistan to his new species, and stated that in this same
area it was sympatric with populations of E. carinatus. Following additional
analysis, Cherlin (1983a) raised Stemmler's sochureki to a full species and
restricted E. carinatus to peninsular India. All mainland Pakistan populations
(except those in northwestern Chagai) were to be called E. sochureki sochureki
Stemmler; the insular form was now E. sochureki astolae Mertens. Echis
carinatus was divided into two races, E. c. carinatus on the Indian peninsular
mainland, and E. c. sinhalensis Deraniyagala (1951) on Sri Lanka.
Being committed to produce a book-length compendium of the reptiles
and amphibians of Pakistan, we were anxious to determine the range and
character limits of these two species in Pakistan. To do this we examined all
available Echis material from Pakistan and neighboring areas. This was
particularly important because no geographic sympatry of the two species
believed to exist in Pakistan (Cherlin 1981) had actually been clearly
demonstrated. We initially presumed that the available material (in museums
and the several hundred specimens we had collected during field work for the
larger project on the herpetology of Pakistan) would establish such areas of
overlap. As shown below, we soon learned that such sympatry could not be
demonstrated and that the current taxonomic arrangement as it applies to
Pakistan is probably incorrect. The results of our studies are presented below.
Mr. Colin McCarthy, British Museum of Natural History, is presently
completing his broader analysis of all Echis "carinatus" material from southern
Asia and northern Africa. This important study will allow our own
geographically restricted one to be placed in proper perspective.
We believe our data demonstrate a pattern of variation in scalation and
color character states that is significant in zoogeographic interpretations






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


regarding Pakistan. This study is the first of several we have planned, each
covering one or more reptile taxa, through which we hope to develop a
zoogeographical overview of Pakistan reptiles based on analyses of more
specimens than has been available to earlier workers.


MATERIAL

Sources


This study is based on 658 specimens from the twenty one institutions
listed below. The museum source of those specimens specifically referred to
are identified by the abbreviations given. The numbers in parentheses indicate
the number of individuals examined in each institution.
American Museum of Natural History, New York (AMNH 9). Natural
History Museum, London (BMNH 11). Bombay Natural History Society
(BNHS 51). California Academy of Sciences, San Fransisco (CAS 5). Field
Museum of Natural History, Chicago (FMNH 21). University of Humboldt
Museum, Berlin (ZMB 1). Naturhistoriches Museum, Basel (NMBA 8).
Museum of Comparative Zoology, Harvard University, Cambridge (MCZ 9).
Natural History Museum of Vienna (NMW 2). Pakistan Museum of Natural
History, Islamabad (PMNH 2). Royal Scottish Museum, Edinburgh (RSM
17). Senckenberg Museum, Frankfurt (SMF 88). University of
Florida/Florida Museum of Natural History, Gainesville (UF 28). University
of Michigan Museum of Zoology, Ann Arbor (UMMZ 4). National Museum
of Natural History, Washington, D.C. (USNM 11). Zoological Survey
Department of Pakistan, Karachi (ZSD 328). Zoological Survey of India,
Calcutta (ZSI 55). National Institute of Zoology, Academy of Sciences,
Leningrad (ZIN 18), and Alexander Koenig Museum, Bonn (ZFMK 5).
All drawings were done by the senior author.


Localities


Figure 1 shows the localities from which specimens were examined.
Appendix 1 provides data on museum holdings of specimens examined from
different geographic locations. No specimens with general, questionable, or
erroneous locality data have been included; nor have we included any literature
records.





AUFFENBERG & REHMAN: THE GENUS ECHIS


0


I I
I---
C


Figure 1. Map showing the localities from which specimens of Echis were examined during this
study.


I






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


ACKNOWLEDGEMENTS


We particularly thank the United States Fish and Wildlife Service (Washington), the
Deutscher Akademischer Austauschdienst (Bonn, Germany), and the Office of Sponsored
Research (University of Florida) for making funds available for us to conduct this study. To all
the curators and collection managers of the institutions listed above, we extend our sincere
thanks for helping us locate material, putting specimens at our disposal for study, and for the
many other ways in which they have contributed to the success of this project. Finally we wish to
acknowledge the support offered by our respective institutions.


CHARACTERS

Characters Used Previously


Cherlin (1981) separated Echis multisquamatus from Echis carinatus
sochureki on the basis that the former possesses a higher number of ventral
scales (mean 188.1, vs. 173.4), a greater number of dorsal scale rows (mean
37.1, vs. 32.7), lateral serrated scales (both sides added together, mean 10.3, vs.
8.8), supraocular absent (vs. present), the light-colored lateral crescent
markings are connected to form a zigzag (vs. separated crescents), and a light-
colored, narrow, cross-like dorsal head marking (vs. a spear or broad "cross,"
see Cherlin 1983a, fig. 2).
In his studies of Indo-Pakistan Echis populations, Stemmler (1969)
separated his Echis carinatus sochureki from congeners on the bases of being
viviparous (vs. oviparous), dorsal color pattern (white spots, vs. other colors),
wide head with a short snout, relatively small eyes, short tail, higher dorsal
scale rows, enlarged supraoculars, head color pattern and lower number of
ventral and subcaudal scales. Of these, the most important characters from the
standpoint of the Pakistan material have also been studied by Cherlin.


Characters Used in This Study


The following characters were tabulated for all the specimens listed
above: (1) number of ventral scales, (2) number of subcaudals, (3) number of
dorsal scale rows, (4) number of serrated oblique dorsal scale rows, (5)
number of gular scale rows, (6) number of subocular scales, (7) presence of
supraoculars, (8) number of dorsal spots, (9) ventral color pattern, (10),
color of infralabials and chin, (11) lateral color pattern, (12) dorsal color
pattern, (13) head and nape pattern. Several of these have been used in
previous studies of Echis, others are used here for the first time. When


268






AUFFENBERG & REHMAN: THE GENUS ECHIS


A B


Figure 2. Variation in number of gular rows. (A) two rows between the enlarged chin shields
and the first ventral shield; (B) four rows.


pertinent, sex was determined by dissection. Definitions for characters
analyzed follow.

Number of ventrals.-- The first scale immediately posterior to the last
chin shield to (but not including) the anal scute. Partial scutes were counted if
they were larger than one third the width of a normal scute.

Number of subcaudals.-- Counts were made from the first normal-sized
subcaudal behind the cloaca to, but not including the terminal spine.
Specimens with a truncated tail were not included in the analyses.

Number of dorsal and oblique scale rows.-- In all specimens of Echis
examined, a number of lateral rows were reduced in size, inclined in
orientation (oblique), and distinctly serrated. This reduction in size and the
attendant close packing of these scales sometimes makes counting dorsal scale
rows difficult. If the change in orientation is not followed and the person
making the count continues the standard manner along the diagonal set by the
first pair of dorsals bordering the ventrals, the counts may regularly be two to
four scales too low.
In this paper all scale row counts were made at midbody. An acceptable
degree of reproducibility is attained by starting the counts caudad and
reversing in the standard manner at the vertebral ridge. The number of
oblique scale rows were counted on only one side and always at midbody. The
oblique, smaller, serrated scales begin on scale rows three or four and continue
caudo-dorsally for from two to six scale rows. The remaining dorsal scales are


269






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


normal in both size and position. Neonates from the same geographic area
have the same number of oblique scales as adults and they seem to be serrated
to the same degree.

Number of gulars.-- The gular rows are those scales extending from
immediately between the posterior pair of enlarged chin shields and the most
anterior ventral scale (Fig. 2). They are more or less arranged in pairs, one
following the other, each pair straddling the midline.

Subocular development.-- In the populations examined, a series of scales
always surrounds the orbit (circumorbital ring); those in the lower part of the
ring are termed suboculars. The subocular count was tabulated as the least
number of scales vertically in the ring below the eye and between it and the
supralabials below. This least number was usually above supralabial number
five, rarely four (Figs. 3, 4).

Supraocular shield.-- Within the circumorbital ring, the most dorsally
located scale above the eye is sometimes larger (longer and often wider) than
the remaining adjacent dorsal head scales. This condition was interpreted as
one in which the supraocular was well developed ("yes"). In other individuals
none of the scales in the dorsal section of the circumorbital series was larger
than the remaining dorsal head scales; this condition was interpreted as lacking
a supraocular ("no") (Fig. 4).

Dorsal body color pattern.-- Figure 5 is a diagrammatic view of the dorsal
scale area of Echis carinatus at mid-body. Both mid-dorsal and lateral color
pattern elements are indicated. The dorsal portion is comprised of a median
longitudinal series of darker dorsal blotches, separated by inter-blotch patches,
both of which are located on a ground color of intermediate density. Dorsal
blotches vary in the density of melanin, number of blotches, and their size.
Melanin density was subjectively noted. Some of the notes taken also indicated
whether the scales were completely or partially darkened and whether the
blotch was lighter in the center or not. Blotch size was approximated by
counting the scales in both the longitudinal and transverse directions. The
number of dark dorsal blotches was counted from the first one immediately
after the light-colored marking on the head and nape, continuing to the last
one over the cloaca (or immediately anterior to it if no spot above it). Due to
considerable bilateral asymmetry in the markings, counts were always made on
the right side of the vertebral ridge. If the blotches were dim their light
interspaces (always evident) were used as a guide.
The lateral color pattern is constructed around two longitudinal rows of
smaller spots, each spot usually covering only a few scales. The lower row of
smaller, usually dimmer spots is designated as I, the upper row of larger, often


270







AUFFENBERG & REHMAN: THE GENUS ECHIS


B C


Figure 3. Lateral view of head, showing typical variation in the minimum number of rows
beneath the eye. (A) minimum subocular rows two SMF 21054, Duschak, Transcaspia USSR;
(B) minimum subocular rows two BNHS 2448, Dhera Ismail Khan, Pakistan; (C) subocular rows
one BNHS 2425, Cumbun, Madura, Tamil Nadu, India. Head shape differences due mainly to
age and size; C smallest, B largest.


Figure 4. Above, lateral views of head. (A) supraocular shield differentiated from remaining
head scales (BNHS 2359, Chabar, Persian Gulf); (B) supraocular not differentiated (SMF 57356,
Astola Island, Baluchistan; holotype E. c. astolae). Below, top view of left side of head. (C)
supraocular not differentiated (SMF 57356, same data as above); (D) supraocular differentiated
(BNHS 2441, Mand, Baluchistan, Pakistan). Head shape differences due to age and size.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


darker spots is designated as II. I or II may be absent and this fact was noted.
Spot size was estimated by counting the total number of scales covered. Finally
the relative darkness of the spots in both rows was noted. A longitudinal series
of lighter (usually white) marks is associated with row II (Fig. 5, see remarks
below).
The dorsal body color pattern was placed into an additional one of two
categories: individuals with reduced melanin in the dorsal spots, the spots
usually with light centers (Fig. 6A) and those with dark, clearly marked
blotches (though these may vary in size) (Fig. 6C).

Ventral color pattern.-- In the populations studied this character ranges
from uniformly clear (no markings), to heavily spotted. The markings of each
specimen were subjectively placed in one of four categories; 0 = uniform
white, cream, etc., without traces of any spots; 1 = very dim spots; 2 = medium
intensity spots; 3 = deeply pigmented spots (Fig. 7). Additionally, the color of
the ventral markings varied from greyish brown to nearly black, and in size
from small to large dots. The last two variables were not tabulated.

Infralabial pattern.-- Many individuals have three dark blotches on the
infralabials. Others grade from this condition to uniform white or cream. We
use 0 for clear, 1 for dim, 2 for medium, and 3 for dense melanin (Fig. 8).

Lateral body color pattern.-- A series of white crescentic marks usually
occur on the lateral body surface in association with pattern cycle II (Fig. 6).
In some parts of Pakistan the crescent tips join to form a zig-zag line (= zig-
zag "yes"), and in other areas they fail to touch (= zig-zag "no") (Fig. 5).
Additionally, the number of rows of small lateral dark spots was tabulated for a
random sample of specimens (see above and Fig. 5).

Head color pattern.-- The top of the head and nape is always marked
with a distinct pattern. These markings vary from an arrow- to a cross-shaped
mark (Fig. 9), in which the figure may be broad or narrow. For analytic
purposes, we tabulated the trident mark (Fig. 9A) as 4, and the arrow mark
(Fig. 9B) as 3, the broad cross (Fig. 9C) as 2, the narrow cross as 1.
Intermediate conditions are indicated by a decimal (i.e., 3.5, 2.5).


SAMPLE AREAS


Figure 10 shows the locations and general size of sample areas chosen for
this study. The geographic limits of the samples were selected primarily on the
basis of sample size (museum material available), but in some cases partly on






AUFFENBERG & REHMAN: THE GENUS ECHIS


ZIG-ZAG
INTERBLOTCH
...: :- "...:::;: :.:

"' ~ DORSAL BLOTCH

... .... GROUND
"X N -CRESCENTS


* *0 0


Figure 5. Color pattern elements on dorsal and lateral body surfaces of Echis.


B C


Figure 6. Major variational types of dorsal body color patterns. (A) BMNH 86-9-2-23-42, Chilez
(= Chilas), Baluchistan; (B) UF 61232, Makli Hill, Sindh, showing an abnormal vertebral dark
zig-zag in an otherwise typical pattern; (C) SMF 57356, Astola Island, Baluchistan, holotype E. c.
astolae.


1 11







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


.-. Y'-











B


~--~ei~Fs ~-~l~s~ 4 4A


Figure 7. Ventral pattern categories used in this study. (A) uniform, or nearly so, no obvious
pattern = 0, MCZ 5405A, 100 mi. S Amballa, U.P., India; (B) markings dim, small, = 1, BNHS
2437, Gosht, Baluchistan, Iran; (C) medium intensity, larger markings, = 2, MCZ 1839,
Mednapur, West Bengal, India; (D) darker, larger markings, = 3, MCZ 2247, 100 mi. S Amballa,
U. P., India; (E) dark, smaller, = 4, SMF 57356, holotype, E. c. astolae, Astola Island,
Baluchistan.







AUFFENBERG & REHMAN: THE GENUS ECHIS


B


Figure 8. Infralabial pattern categories used in this study. (A) uniform, no pattern = 0, SMF
57356, holotype, Astola Island, Baluchistan; (B) dim markings = 1, SMF 63137, Nushki,
Baluchistan; (C) medium intensity = 2, SMF 63136, Hingol, Baluchistan; (D) intensely
pigmented = 3, SMF 55015, Turkmen SSR.


Figure 9. Head pattern categories used in this study. (A) broad cross, = 2, BNHS 2441, Mand,
Baluchistan, Pakistan; (B) arrow, = 2, SMF 63136, Killi Mangal (Hingol), Baluchistan; (C)
narrow cross, = 1, BMNH 86-9-21-124, Chilez, Baluchistan; (D) a rare trident pattern form (not
tabulated separately, but placed with "arrow" category), SMF 57256, holotype, E. c. astolae,
Astola Island, Baluchistan.


f-`







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 10. Location (and general size) of sample areas used in this study. Abbreviations used
are as follows: A, Afghanistan; CS, Central Sindh; GT, Gosht; Gu, Gujarat; I, Islamabad; K,
Karachi; Ki, Kirthar Range; L, Lahore; LS, Lal Suharna National Park; M, Multan; MC,
Mekkran Coast; NP, Nagar Parkar;; N, Nushki; NWFP, Northwest Frontier Province; NWR,
Northwest Rajasthan. P, Panjgur; Q, Quetta; Raj, Rajasthan; R, Rattankot; S, Shadadkot; SB,
Shah Bundar; TP, Thar Parkar; Th, Thatta; Z, Zabol.


276






AUFFENBERG & REHMAN: THE GENUS ECHIS


environmental differences between closely approximated geographic areas (i.e.,
elevation, major environment). Each of these sample areas served as the basis
for all calculations and evaluations, so that all specimens available from each
area were considered as constituting the same sample for computational
purposes.


VARIATION


Geographic Variation


Ventrals.-- Within Pakistan the number of ventrals vary from 145 to 189
and sample means from 155 to 189 (Islamabad and Nushki samples). The
general geographic pattern produced by the sample means is shown in Figure
11.
The geographic pattern is characterized by an area of high ventral counts
in both northwest Baluchistan (mean 181-183) and the northern Thal Desert
(mean 181) areas; and areas of low counts in both the northern mountains
(154-158) and the southern part of the Thal Desert (158). The means of
geographically intermediate sample areas are numerically intermediate. The
means of the two areas with high mean values are significantly different from
those with low means (t-test, at at least the 2 % level, see Table 1). Along the
Mekkran Coast the mean number of ventrals decreases steadily from west to
east and the trend is continued into adjacent coastal India (172 Gosht, Iran, to
162 in Gujarat, India). Separately, these coastal populations are significantly
different in mean ventral counts from inland populations in Baluchistan and
Sindh (Table 1), though the zone of high ventral counts in the coastal areas of
the latter is quite narrow (Fig. 11). The trend is reversed along the frontal
ranges of the Himalayas (154 NWFP to 162 Lahore), but without statistical
evidence for any significant differences in the means of mountain and plains
populations. There is no significant difference in mean ventrals of northern
Pakistan and northern India (Ganges Plain, 157) from that in southern
Pakistan and adjacent Gujarat. However, both coastal and mountain samples
from India southeast of Gujarat have mean ventral counts significantly lower
than those of southern Pakistan (Bombay 146, Poona 146) (Table 1). On the
Indian peninsula the trend to decrease ventrals southeastward is continued,
with the lowest values of all observed in the Goa sample (143; a significant
difference in means occurs between the adjacent Goa-Bombay samples, p <
0.05 % level, though not between any other population pair members in the
rest of the peninsula).







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 11. Mean number of ventral scales in Echis carinatus populations studied in Pakistan
(left) and in Indian peninsula (right), with estimated positions of isophenes of identical values.


181

181






AUFFENBERG & REHMAN: THE GENUS ECHIS


In the Chagai area (Nushki sample) the high mean value (183) cannot be
statistically separated from that of the Zabol, Iran (181), and Afghanistan
samples. However, the Nushki mean is very significantly different from the
adjacent Quetta one (165, p < 0.001). In the same way, the high mean value of
the northern NW Rajasthan sample is very significantly separated from all
surrounding samples (p's all < 0.05).
Several neonate samples were available for study (UF, SMF, ZSI). These
samples demonstrate something of the level of variation occurring in the
ventral and subcaudal scales among the siblings of single broods (Table 2).
These broods were available from mothers originating in Shadatkot, Karachi,
Thar Parkar, Quetta, Goa and Kerala. No significant difference between the
mean number of either ventrals or subcaudals of the siblings and the means of
these variables in the home area of the female was shown in t-test analyses.

Subcaudals.-- Analysis of the subcaudal counts results in a geographic
pattern similar to, but less pronounced than that for the ventrals (Fig. 12). The
highest mean values of subcaudals are found in the non-coastal areas of
Baluchistan, adjacent Iran and Afghanistan. The most significant difference in
means of these populations and adjacent ones occurs between the Nushki-
Quetta pair populations (means 34-29, t-test = 7.2, df 22, p < 0.001). The
seaboard populations demonstrate generally low, but not significantly different
values from those of the remainder of Baluchistan. Nor are any of the other
values significantly different for adjacent pair members through the remainder
of the country. The general trend is for values to be reduced into and through
peninsular India, but with the Goa and Kerala coastal populations again
showing a significantly different mean value. NW Rajasthan (and Lahore)
have high mean values (33-34), but they are not significantly different by t-test
analysis.

Dorsal Scale Rows.-- Figure 13 shows the geographic pattern of the mean
number of dorsal scale rows in Pakistan. The pattern is simple, with no
demonstrable significant differences between the means of any adjacent
samples. However, the general aspect is similar to that for the subcaudals.
Thus the Nushki sample is statistically indiscriminate from the Afghan sample.
The Indus River Valley means tend to be low, continuing into India, where
peninsular samples have lower means than northern ones. However, the
means of the west coast peninsular samples are not significantly different from
those of the remaining ones. The NWFP and Islamabad samples are low when
compared to the Indus Plains samples (but not significantly different). There
is no demonstrable difference between the mean of the NW Rajasthan sample
and those of surrounding areas.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 12. Mean numbers of subcaudal scales in Echis carinatus populations studied in Pakistan
(left) and in Indian peninsula (right), with estimated positions of isophenes of identical values.


30

33

34







AUFFENBERG & REHMAN: THE GENUS ECHIS


Figure 13. Mean numbers of dorsal scale rows in Echis carinatus populations studied in Pakistan
(left) and in Indian peninsula (right), with estimated positions of isophenes of identical values.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Oblique Scale Rows.-- The general pattern for this character (Fig. 14) is
similar to the previous one for dorsal scale rows. The two, are in fact,
correlated (R = 0.88). There is no demonstrable statistical difference in
means between any adjacent Pakistan population. The means of samples from
eastern Iran (7), western Baluchistan (7) and Afghanistan (8) are very clearly
higher than the means for any other part of the country (3-6), with the
exception of NW Rajasthan (7). The lowest value (3) occurs in Nagar Parkar.
In general, the Indus Valley has rather low values.

Gular Scale Rows.-- In spite of the fact that the range in gular scale row
variation is rather narrow, means of adjacent samples are often significantly
different. The geographic pattern is shown in Figure 15. Means range from
5.0 (northwestern Baluchistan and Karachi) to 3.0 (Shah Bundar). The entire
Thar Desert and Ganges areas are higher than surrounding samples, but the
means are not significantly different. However, the Nushki sample mean (5.0)
is significantly different from that of the Quetta sample mean (3.9; t = 9.4, df
19,p < 0.001). The Shah Bundar and adjacent Gujarat mean values (3.0, 3.3)
are significantly different from all the surrounding ones (p < 0.025--< 0.001).
While none of the peninsular Indian samples differ significantly from one
another, that from Gujarat is clearly separable from that from Bombay (t =
6.7, df 24,p < 0.001). South along the west coast of India there are additional
significantly different mean values in adjacent populations, especially that
separating the Goa sample from it's neighbors (t = 5.5, df 42,p < 0.001). The
Sri Lanka and Tamil Nadu mean values are also significantly different (t = 8.6,
df 11,p < 0.001).

Number of Suboculars.-- The lowest mean values for subocular scale
rows are found in southern peninsular India and Shri Lanka and the highest
ones in the northwestern parts of the range, such as Afghanistan and the
border area surrounding it. Typical examples of geographic variation in
subocular scalation is shown in Figure 3; the pattern of geographic variation is
shown in Figure 16.

Supraocular development.-- In some individuals, an enlarged supraocular
scale may be present on one side and not on the other (to 25% of those with
developed supraoculars). However, the majority are symmetrical. The
geographic variation in the presence of enlarged supraocular scales is shown in
Figure 17. Here, the pattern is one in which there are significant changes
between the southern Afghanistan population (identical to Transcaspian
material) and all border samples in adjacent Iran and Pakistan. The Cholistan-
Thar Desert and Nagar Parkar samples are also distinctive in all individuals
having a well-developed supraocular. However, the Shah Bundar sample,







AUFFENBERG & REHMAN: THE GENUS ECHIS


5 / 4
5 5

4 4
4


4 4


4LV






Figure 14. Mean numbers of oblique scale rows in Echis carinatus populations studied in
Pakistan (left) and in Indian peninsula (right), with estimated positions of isophenes of identical
values.


7 6







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 15. Mean numbers of gular scale rows in Echis carinatus populations studied in Pakistan
(left) and in Indian peninsula (right), with estimated positions of isophenes of identical values.







AUFFENBERG & REHMAN: THE GENUS ECHIS


C-
,3Q


II

Figure 16. Mean minimum numbers of subocular scale rows in Echis carinatus populations
studied in Pakistan (left) and in India (right), with estimated positions of isophenes of identical
values.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 17. Mean percent of sample areas with developed supraoculars in Echis carinatus
populations studied in Pakistan and India, with estimated positions of isophenes of identical
values.






AUFFENBERG & REHMAN: THE GENUS ECHIS


having a low proportion of individuals with well developed supraoculars, is
surrounded by samples in which significantly few individuals have supraoculars.

Dorsal body color pattern.-- Figure 6 shows the extremes of dorsal color
pattern found in Pakistan samples. The dorsal color pattern of Pakistan saw-
scaled vipers is characterized by having a series of short, light, transverse ovals
across the the paravertebral area (Figs. 5, 6) from just behind the head to at
least above the cloaca, often onto the tail. These ovals are often broken into
two paravertebral spots that are sometimes connected with a diagonal dark
zone that in some individuals produces a dark vertebral zig-zag, part or all the
way down the back (Fig. 6); in the Astola Island population (E. c. astolae)
these markings become distinct narrow transverse bars (Fig. 6). Specimens
from the adjacent Mekkran Coast population are sometimes similar, but less
distinctly barred. Table 4 summarizes the most common color pattern types in
each sample area. A single individual out of over 350 examined by the junior
author from Makli, Sindh, was provided with three vivid longitudinal stripes,
without any evidence of spot rows I and II.
In Pakistan, the mean number of dorsal body spots varies from 31
(Lahore and Rattankot) to 40-41 (Cholistan-Thar Desert area). Baluchistan
populations have more dorsal spots than those in the Indus Valley (Fig. 18). In
spite of the low level of variation over the entire area studied, there are many
statistically significant differences between adjacent population means,
suggesting that this is a sensitive character, useful in differentiating Echis
carinatus populations. These differences are shown in Table 3.
In general, the Pakistan pattern is part of the general trend within the
Indian Subcontinent, i.e., higher values northwestward and lower values
southeastward. The trend into peninsular India is an extension of the Pakistan
pattern, except that Goa and Kerala have a significantly lower mean number of
spots of all of the remaining Indian samples. It differs most from the Tamil
Nadu sample (p < 0.001). The Sri Lanka and Tamil Nadu samples are not
significantly different from one another.
The density and distribution of melanin in the dorsal blotches varies
geographically in Pakistan, though exhibits a fair degree of variation within
single populations. Character state A (Fig. 6A), in which the melanin
distribution may be dense, though found on only parts of the scales making up
the blotch (mean number of scales involved 13, only a few of which are
completely covered), is more or less restricted to arid environments (soil types
camborthids and torri- and ustipsamments, fide Atlas of Pakistan, Govt.
Pakistan: Islamabad, 1986). It is most common in individuals from extreme
western Punjab and the Irano-Pakistan borderlands.
The most common and widespread category in Pakistan is represented as
B in Figure 6. It is one in which the melanin is dense and tends to cover most
scales entirely. However, the spots tend to be rather small (total 18 scales in







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 18. Mean numbers of dorsal body spots in Echis carinatus populations studied in Pakistan
(left) and in Indian peninsula (right), with estimated positions of isophenes of identical values.






AUFFENBERG & REHMAN: THE GENUS ECHIS


mid-body blotches). It is found throughout most of the country, occurring on a
variety of soil types (ustorthents to ustipsamments).
The largest blotches are found in the population of Astola Island,
Baluchistan (Fig. 6C; blotches include a mean of 22 scales at mid-body). In
this population the center of the blotches are usually dark brown and the outer
scales black.
As expected, there is significant variation in ground color. In general, it is
lightest in populations living on light-colored soils, though not necessarily
darkest on the darkest soils. Within Pakistan the darkest individuals are
apparently found at Parachinar, NWFP, in which the ground color is grayish
brown. In most other localities it is a shade of grey, sometimes locally with
tinges of greyish-yellow or pinkish-yellow in life (locally).

Ventral Color Pattern.-- In general, the geographic variation in this
character is the same as that exhibited by other variables described above,
except that the values tend to be reversed (high means in the other patterns
tend to be low means in this one). The highest mean values (darkest spots) are
found in inland localities of eastern Iran (Zabol, Gosht, etc.), extreme northern
Pakistan (NWFP) and western coastal India (Goa, Kerala). The lowest means
(lightest ventral color) are found in central Sindh, NW Rajasthan, and the
Dekkan Plateau of peninsular India. Figure 19 shows the geographic pattern
of variation in this character. Table 4 gives the values and significance levels
for sample pair comparisons.
The density of the color of the ventral markings, or whether they even
occur at all, is often rather variable within a single sample. Thus, along the
Persian Gulf and Mekkran Coast (N = 38) individuals with class 0 belly
pattern (clear) occur in 8 percent, class 1 in 50 percent, class 2 in 21 percent,
and class 3 in 21 percent. The same level of variation can be demonstrated in
the Gujarat and Rajasthan samples. On the other hand, some populations are
very homogeneous in this regard, particularly those in desert areas (Nushki,
Cholistan-Thar Desert, etc.), where the spots are consistently light in intensity.
Of all the variables examined, the most significant interpopulational
differences occur in this character. Table 4 provides pertinent data for those
population pairs in which the means of the degree of ventral spotting are
significantly different. Figure 19 shows that the trends established through
Pakistan are continued into the adjacent parts of India. Prominent features are
the slightly darker venters of those populations from both the northern
Gangetic and Indus Plain in both countries, extending east to the delta of the
Ganges River in West Bengal. The lightest venters are found in a somewhat
diagonal strip running from the northwest to the southeast. The Goa area
stands out as having significantly darker bellies than surrounding populations
(Goa-Bombay, t-test, df 32,p < 0.001; Goa-Kerala, df 16,p < 0.005).







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 19. Mean intensities of belly spotting in Echis carinatus populations studied in Pakistan
(left) and in Indian peninsula (right), with estimated positions of isophenes of identical values.


1.5 3.0 3.

1 1.





1.5
1.3 2.0
1.5 1.0






AUFFENBERG & REHMAN: THE GENUS ECHIS


The size of the ventral spots also varies geographically. In general the
spots are largest in those Pakistan populations from both coastal and northern
mountain areas, and smallest in individuals from interior desertic
environments. The character was not easily divisible into clearly recognizable
categories and so was not treated statistically.

Infralabial and genial scale color.-- While the infralabials of some
Pakistan populations are more or less uniform white to light grey, others
possess up to three (usually two) dark spots on the infralabial scale series.
Figure 8 shows the extent of variation in the distribution and density of
melanophores on the infralabials and genial scales. In general, those
populations with the least melanin are found from northwest India west
through most of central Pakistan, thence into adjoining Iran and Afghanistan.
Regions in which dark infralabial spots are found are the west coast of India,
coastal Pakistan, and the front ranges of the Himalayas in both these countries.
Most Pakistan Echis have no melanin on the genial series on the throat;
some do. When present, a dusky mark is seen on each of the scales. This
character is statistically correlated with the presence of dark infralabial marks
(R = 0.72) and has the same general distributional pattern, but less easy to
categorize. We have not analyzed this character in detail. Darkness of the
infralabials and chin is positively correlated with darkness of ventral spotting
(R = 0.78).

Lateral Body Color Pattern.-- The Pakistan populations can be separated
into two major groups on the basis of lateral body patterns, those with light
lateral zig-zag markings and those with light-colored lateral crescents (Figs. 7,
20A, C). In the material we examined, the zig-zag condition occurs in all
individuals in Chagai and adjacent Afghanistan (Fig. 21). It remains common
through all of western Pakistan, being present in from 50 to 64 percent of the
population. Samples from the entire Indus Valley eastward, except Lahore and
Shah Bundar, lack the zig-zags entirely. Instead, the marks are represented by
light inverted crescents. In the Himalayan foothills from NWFP east and into
the Ganges Plain in India, the zig-zag lateral pattern is found in from 25 to 50
percent of each of the samples. The populations are thus intermediate
between the two extreme character states, if not in geographic location. The
coastal populations of both Iran and Pakistan are similar. Those from NWFP
south along the foothills of the Sulaiman Range are intermediate in both
condition and location. Along coastal Iran lateral light markings of any kind
are often entirely absent, as they nearly are in insular populations in western
Pakistan (i.e., Astola Island). In the eastern half of Pakistan the crescent type
pattern predominates (Fig. 21). Between the two extremes (zig-zag and
crescent) every gradation of intermediate condition, including bright, evident
crescents connected by faint zig-zags. In some populations both zig-zag,







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


A B


cS



C


.> r- n~ v,;~-~
*J~_~LT ~. --,-



D


Figure 20. Typical lateral body patterns. (A) dark ground color, bright narrow zig-zag, no
evidence of spot row I, faint traces of spots on II, no light crescents, BNHS 2448, Dera Ishmael
Khan, Punjab, Pakistan; (B) medium ground color, very dim, wide zig-zag, spot rows I and II
both evident, SMF 57356, holotype E. c. astolae, Astola Island, Baluchistan; (C) light ground
color, crescents present, spot row I absent, II evident, AMNH 81980, Las Bela, Baluchistan; (D)
light ground color, zig-zag present, no evidence of spot rows I and II, SMF 63136, Killi Mangal
(Hingol), Baluchistan.



crescent and intermediates are equally common; a few individuals have both
pattern cycles on their body, the zig-zag usually being found anteriorly and the
crescents posteriorly.
Lateral spot rows I and II also vary geographically. Over much of
Pakistan, row I is completely missing or very faint, while the spots of row II
may be very dark (Fig. 20D). In western Punjab row II may also be absent or
only faintly indicated (Fig. 20A). On Astola Island, both rows, with spots of
medium density, are present (Fig. 20B).





AUFFENBERG & REHMAN: THE GENUS ECHIS


0


I7
75 .'


Figure 21. Percent of sample populations with evident light lateral zig-zag.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Head and Nape Color Pattern.-- Figure 9 illustrates major pattern
categories seen on the head and neck of Pakistan Echis. The trident pattern
(Fig. 9D) has only been seen in the Astola Island population. It is undoubtedly
a modification of the wide arrow type. The most common types in Pakistan
are "arrow" (Fig. 9B) and"cross" (Fig. 9A, C). In general, the narrow cross
pattern is mainly found in Baluchistan and adjacent parts of Iran and
Afghanistan to as far north as Turkmen SSR. In Chagai District, Baluchistan,
it is found in almost all individuals. It occurs in a few individuals of all samples
as far east as Andra Pradesh, India. The arrow type is most common in Sri
Lanka and southern peninsular India. The wide cross type is most common in
the geographically intermediate areas. It is rarely dominant in any of the
Pakistan samples studied, the narrow cross type being the most prevalent.


Sexual Variation

An analysis of the entire sample demonstrates that there is no significant
difference in the number of males and females, the sex ratio being essentially
1:1. However, Rehman (MS) shows that this sample ratio varies seasonally,
depending primarily on the activity level of the males and females during
different months of the year. He shows that such differences as exist are
correlated primarily with different phases of the reproductive cycle in both the
males and the females.
A sample of 64 adult males and an equal number of adult females was
randomly drawn from the largest sample available (Thatta, N = 387). On
these specimens, the ventrals, subcaudals, dorsal scale rows and the number of
dorsal spots were counted, keeping individuals of the two sexes separated. In
addition, 66 neonates from several Pakistan and Indian localities (see Table 2
for sample sizes and localities) were similarly examined. No statistically
significant differences could be demonstrated between sex and the mean of any
of the variables analyzed, except the mean number of ventral scales. For this
variable, the mean number in females was significantly higher at a level of
about one percent (t = 2.84, df 62, p < 0.01), though the differences between
them are slight (males 164.9 6.6, mean deviation 4.6 ventrals; females 169.6
+ 7.7, mean deviation 8.2 ventrals. The females not only have a slightly higher
mean number of ventral scales, but they are more variable (CV = 4.5 percent,
as opposed to 4.0 in the males).






AUFFENBERG & REHMAN: THE GENUS ECHIS


SUMMARY AND DISCUSSION

Systematics

There is no evidence to suggest that more than one species of Echis
occurs within the boundaries of Pakistan. We base this on three major points.
First, all the characters studied showed complete blending between them.
All gradations of intermediacy were observed, although many of the
intermediate conditions were restricted geographically (see below). If two
species are represented in the Pakistan populations, one would expect evidence
of bimodality in at least some of the characters studied. There is no such
evidence in any of the characters. As an example, one of the most
geographically sensitive characters we analyzed is the mean number of ventral
scales in different populations. Figure 22 compares the frequency curves for
ventral scales in individuals taken from Pakistan, where two species of Echis
were believed to occur, and the same for central to southern Indian peninsula
populations, where all workers agree that only one species is found. Neither
curve shows any visual evidence of bimodality; nor is there any statistical
evidence of bimodality; (moment coefficients of skewness 4.06 for the Pakistan
populations and 3.92 for those from the Indian peninsula). However, both are
highly kurtotic (moment coefficient of kurtosis 21.52 for the Pakistan sample
and 20.90 for the extra-Pakistan samples). This flattening of the frequency
distribution is caused by the demonstrated geographic variation in ventral
scales in both geographic areas.
Second, at least two forms previously called species (Echis carinatus and
E. sochureki) have bred in captivity and produced young with completely
intermediate characters (preserved hybrid HUB 18565).
Third, there is no hiatus in the saw-scaled viper populations of the Indian
Subcontinent from at southern Afghanistan and eastern Iran through Pakistan
to the furthest reaches of Peninsular India and Sri Lanka. Nor is there any
morphological evidence to support the contention that two or more species
exist throughout this area. All of the characters we have studied blend
completely from west to east through a series of variously intermediate steps.
We see no evidence of sympatry in northern Pakistan and/or southern
Afghanistan populations of Echis c. multisquamatus and E. c. sochureki (as
presumed by Cherlin 1981).
The name available for this species is Echis carinatus. However, within
the Indo-Pakistan area we recognize four distinct geographic races. On the
basis of priority rule, the nominate form is Echis carinatus carinatus
(Schneider), found in southern peninsular India and Sri Lanka. It's northern
limit is not clear to us because the geographic variation in all the characters
examined in peninsular India represent broad morphoclines in which narrow
zones of intergradation are not easily defined. An exception is the coastal plain







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


20

10

0

CU
O 20
L -
10



VENTRALS

Figure 22. Frequency distribution of ventral scales. Above, Pakistan specimens only;, below,
peninsular India specimens only.


of Goa and Kerala (India). Here each of a suite of several important
characters can be shown to be distinctly different from all surrounding
populations. Since the taxonomy of Indian populations is beyond the scope of
this paper, we leave clarification of the nomenclatorial status of these
populations to others in the future.
Within Pakistan three races of Echis carinatus are easily distinguished.
These are to carry the names already given them by other workers in the past.
One of these is to be called Echis carinatus multisquamatus (Cherlin). In
Pakistan it occurs in relatively "pure" form in the Chagai area of northwestern
Baluchistan (Fig. 23). From here its range extends generally northwest to the
type locality in Turkistan, USSR. Southward in western Baluchistan it is
replaced by variously intermediate populations. It extends to the Mekkran
Coast. The second race is Echis carinatus sochureki Stemmler. It is found
primarily in the Indo-Gangetic Plain, including most of eastern Pakistan and a
large part of northern India (Fig. 23). Intermediates between it and E. c.
multisquamatus occur in sometimes apparently isolated populations in
mountainous areas of eastern Baluchistan and western Sindh. Unfortunately,
the type locality (Pishen, Baluchistan) borders on a zone of intergradation with
E. c. nultisquamatus. Relatively pure forms of both races occur closest to one
another in the Nushki-Quetta transect, between which there is a narrow band
of intergradation. Variously intermediate populations occur in a large part of
coastal Baluchistan. As pointed out above, intermediate populations between
E. c. sochureki and E. c. carinatus occur over a large part of central India. A






AUFFENBERG & REHMAN: THE GENUS ECHIS


MULTISQUAMATUS


ASTOLAE

ASTOLAEI


Figure 23. Taxonomic allocation of Pakistan Echis carinatus populations as applied in the current
study. The clear area represents the major zone of intergradation.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


third subspecies, Echis c. astolae Mertens occurs only on Astola Island,
Baluchistan. Presumed intermediates between it and Echis c. sochureki X E. c.
multisquamatus are known from several localities along the Mekkran Coast. In
many ways this is the most distinct Echis population in Pakistan, probably
because of its isolated, insular distribution.
Diagnoses of the races occurring in Pakistan are as follows:

Echis carinatus multisquamatus.-- A race characterized by generally more
scales in almost all parts of the body and head when compared to conspecifics;
i.e., more gulars (means of local populations usually > 4.5), ventrals (means of
local populations > 180), subcaudals (means 30-34), serrated oblique lateral
scales (means > 7), dorsal scale rows (means > 32), and suboculars (means
2.0-2.8). The supraoculars are almost always discernable. Mean number of
dorsal body spots 36-39, distinct white blotch interspaces, and a lateral color
pattern, while similar to that of the other races, is provided with a rather wide,
vivid light zig-zag stripe. The belly is marked with scattered dark dots; there
are usually three dark spots on the infralabials.

Echis carinatus astolae.-- A race characterized by fewer scales of the types
mentioned under Echis c. multisquamatus (population mean of ventral scales
170 subcaudals 31, dorsal scale rows 31 oblique scales 5.5 gulars 4.3;
supraoculars variably distinct or not). It has a body pattern that is considerably
darker than that of the other races, with larger dorsal blotches (mean number
38), almost no white blotch interspaces, and in which there are two distinct
rows of alternating, rather large dark spots laterally. The belly is marked with
scattered small but dark dots. Only about 25 percent of the individuals have a
light lateral zig-zag.

Echis carinatus sochureki.-- A much more variable race in regard to
almost all of the characters studied. In local populations the mean scalation is
as follows: ventrals 154-181, subcaudals 27-34, dorsal scale rows 28-32, oblique
serrated scale rows 3-7, gulars 3.2-5.0, subocular scale rows 2, discernable
supraoculars in from 33 to 100 percent of local populations. The mean number
of dorsal spots varies from 31 to 41; white blotched interspaces are always
present; belly spotting varies from none to very intense and lateral zig-zag
ranges from none to about half of the individuals in some local populations.

Within the range of Echis c. sochureki there are several smaller areas that
are important because they contain populations unlike those typical of the
subspecies. One of these areas is NWFP and northwestern Punjab. Here a
series of characters are statistically different from typical E. c. sochureki. With
additional material, some future workers may prefer to segregate them as a






AUFFENBERG & REHMAN: THE GENUS ECHIS


separate race. We feel that the nomenclature used here defines the biological
situation with sufficient accuracy.
A second area of interest lies in the Cholistan-Thar Desert, straddling the
Indo-Pakistan border. Here populations of E. c. sochureki are distinctly
different from surrounding populations in both color and scalation. We believe
this population represents an incipient geographic subspecies of Echis carinatus
that is adapted for life in sandy deserts. However, we do not think it warrants
nomenclatorial status.

Zoogeographical Implications

Pakistan (24-37 N lat., 61-770 E long.) is a rather large country (ca. 2000
km east-west and 2200 km north-south, or ca. 805,000 km2). It serves as a
northwest to southeast link between the highly distinct biotas of the steppe,
deciduous forests and deserts of temperate Central Asia and the deciduous
forests and deserts of Tropical South Asia on the one hand, and as an east to
west, arid to mesic link along the northern mountain ranges.
Climatic conditions of Pakistan are characterized by great areas of aridity
in almost all parts of the country, including some of the high northern
mountain ranges. Except for the Indus River and its tributaries, large rivers
and hydric or even mesic riverine forests are few and then only of small extent.
Mean annual precipitation varies from less than 100 mm/year in some regions
(Chagai and Cholistan-Thar Deserts) to nearly 2000 mm in others (parts of
Azad Kashmir); within the range of Echis carinatus, it ranges only to about
500 mm. For the most part, abundant rainfall is only experienced in the
southern ranges of the northern mountain areas. Within the distribution of the
saw-scaled viper, mean warm season temperatures vary from 25-380C. This
viper inhabits southern temperate, subtropical and tropical climatic regions
and the following major physiographic provinces: Northern Mountains, Iranian
Plateau (much of western Baluchistan), Indus Valley, the Central Mountain
Borderland (separating the last two provinces), and the Coastal Plain along the
Arabian Sea.
The only previous study of the zoogeography of the entire Pakistan reptile
fauna is by Khan (1980), which is more a compilation than an analysis and
provides little new information. Other than the current study on Echis
carinatus, there is no single-species analysis of reptiles for Pakistan that
provides details on the probable boundaries of geographically distinct
population gene pools. We would have liked to be able to analyze such
boundary effects in greater detail, especially from a statistical standpoint.
Unfortunately, there are no satisfactory statistical tests of ordination for
zoogeographical studies (Gauch 1982). As a result, our conclusions regarding
such factors rest on a series of Student t-tests between adjacent sample
populations. Concordance of statistically significant differences in the mean






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


values between the same adjacent populations suggests significant differences
between them in several characteristics. We take this level of concordance as a
primary factor in determining the location of zones of least gene flow. We
hope to analyze other species in the future and to examine the degree of
concordance in gene barriers between species to obtain a more accurate
picture of the factors important in the zoogeography of Pakistan reptiles.
Analysis of geographic variation in 13 morphological characters of Echis
carinatus makes obvious a small number of underlying geographic patterns in
this species within Pakistan. Most of the statistically significant differences in
mean character values can be shown to exist in a limited number of sample
pairs. These sample pairs and the number of characters that are significantly
different between them are shown in Figure 24. It illustrates that of all the
potentially interacting sample pairs, the greatest number of differences is
between the Nushki-Ouetta pair (7). It also shows that there is a north-south
line representing many differences between adjacent east-west sample pairs
from Shadatkot through Central Sindh to Rattankot, separating the Indus
Valley populations from those of central and western Baluchistan. There are
few significant differences between the various mainland Mekkran Coast
samples. Nor are there many significant differences between samples in the
remainder of central and northern Pakistan. However, NW Rajasthan is
significantly different in a number of characters from all surrounding samples.
Lastly, the Indus delta area is represented by a character complex in which
almost all samples are significantly different from neighboring ones in one or
more characteristics. A similar mozaic pattern of deltaic populations has been
described from the mouth of the Mississippi River (U. S. A.) (Wilson 1970).
In Pakistan the Indus Delta is the site of some species endemism among both
the lizards and the turtles (Minton 1966). We presume that this mozaic of
character state distributions in the Indus Delta was produced through repeated
isolation of populations caused by dramatic changes in deltaic water
distribution channels. The appearance, disappearance, and changing courses
of distributaries in the Indus Delta during the past several hundred years are
well documented (De Terra and Hutchinson 1936, see Flam 1986 for a recent
review) and much older changes during the Quaternary are known for several
sites in eastern Pakistan and adjacent India (Oldham 1886, Marshall 1931, De
Terra and Patterson 1939, Erikkson 1959). We believe the isolation and
reconnection of populations of Echis carinatus in this area by changing
distributary patterns are directly responsible for the mosaic pattern of
character states seen in the delta area. Thus the major part of the between-
sample differentiation in this area is believed to have a historic basis.
The north-south line of differentiated populations paralleling the
Sulaiman Range and its outliers also seems to have a largely historic basis, for
environmental conditions are largely identical on both sides of the dividing
zone (Fig. 25). We conclude that the populations on either side of the







AUFFENBERG & REHMAN: THE GENUS ECHIS


1\


-" 3


Figure 24. Numbers of statistically different character states between adjacent populations in
Pakistan.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Figure 25. General distribution of natural vegetation types in Pakistan (simplified, but based on
Map 52, The Atlas of Pakistan, 1985, Rawalpindi, Pakistan). (1) Desertic; (2), Tropical thorn
forests; (3) Tropical deciduous forests; (4) Temperate pine-juniper forests; (5) Subtropical dry
pine forests; (6) Temperate dry pine forests; (7) Temperate broad-leafed forests; (8)
Temperate pine/steppe (usually interspersed between mountain ranges in north, variously mixed
with juniper in south).






AUFFENBERG & REHMAN: THE GENUS ECHIS


mountains arrived there from different directions. Those west of the
mountains are clearly closely related to Transcaspian and Iranian Plateau
populations and must have arrived from the west and/or northwest. Those
east of the mountains have been associated with the Indus Plain for thousands
of years. The two populations are more or less genetically isolated by the high
Sulaiman Range, the mountains having few passes at low elevations. However,
a natural avenue leading from lowland arid lands in the Indus Plain to the arid
lands of the Iranian Plateau occurs from near Jacobabad (and the Shadadkot
sample) to Quetta across the Bolan Pass (1650 m). This pass is unquestionably
a major path for genetic interchange between the Trans-Sulaiman population
in northwestern Baluchistan and those of the central Indus Plains. Through
appropriate similar habitat (Fig. 25) and climate, the Quetta population is also
genetically closely related to those of the NWFP. Thus the fact that the Quetta
sample is more closely related to those from Shadadkot (Indus Plain) and from
NWFP than to the geographically much closer Nushki sample (Iranian
Plateau) seems to be largely a matter of more rapid gene flow through similar
habitats than across the borders of dissimilar ones. Apparently a more
southern lowland gene pool and/or an eastern subtropical one of Echis
(sochureki) was brought into close juxtaposition with a temperate plateau pool
(multisquamatus). There is no evidence that the two populations are
genetically incompatible, or that they are sympatric, i.e., forming the northern
ends of a Rassenkreis around the Sulaiman Mountains and through the Bolan
Pass. Rather, the two forms seem to blend evenly and completely into one
another over a narrow zone of intergradation.
The fact that the E. c. multisquamatus character complex is approached
by populations in similar desert environments along the eastern borderland of
Pakistan (Thar-Cholistan Desert) suggests substantial similarity in certain
selective pressures, in spite of the fact that the northwestern deserts are
temperate, and the eastern ones are tropical. While populations from this area
are distinct from surrounding ones in several characters, the number and kind
of differences suggest they are most closely related to those of the Indo-
Gangetic Plain. The differentiation witnessed is probably based on factors
associated with the Pleistocene local desertification of the area (Karpov and
Nebolsine 1964, and others).
The character differences exhibited between the Nushki-Quetta and NW
Rajasthan-Multan populations reflect points along two ecoclines (sensu
Auffenberg 1955), rather than geoclines, for environmental conditions on
either side of the intermediate zones are significantly different (desert versus
non-desert). Other ecoclines may include the change in characters between
the population of northern and southern India, and from Nushki to the
Mekkran Coast. Probably most significant are the series of lines between the
Himalayan foothills (more mesic) to the central Indus Plain (more arid). The
differences in the latter are along a north to south, cooler to warmer






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


environmental gradient. Among others, Edgren (1961), Smith (1956), and
Christman (1980) have demonstrated the common existence of important
north-south lines in the mean scale values of snakes, so that this particular
series of lines in Echis is perhaps expected. However, other important lines
in Pakistan Echis carinatus are clearly east-west. The several character lines
along the Mekkran Coast Coast are examples. While the Himalaya-Indus
Plain changes are clearly ecoclines of some sort, those of the Mekkran Coast
are not, for environmental conditions from the Iran border to at least the Hab
River are very similar, yet changes in mean character states occur in a regular
pattern. Fox et al. (1961) have shown a positive correlation between
developmental temperature and meristic counts in snakes. Thus the
developmental temperatures of embryonic snakes may be the mechanism
maintaining the dines witnessed, but if so, the adaptive significance remains to
be discovered.

Possible Selective Forces

In Echis carinatus some of the most important characters showing
significant geographic variation are scales. Some of the dines demonstrated in
Echis meristic characters are clearly ecoclines (sensu Auffenberg 1955),
maintained by adaptation to environmental conditions that also vary clinally. A
true geocline occurs in northern peninsular India. Within Pakistan, the
patterns of variation in such characters as ventral scales are ecoclines on the
basis of the fact that they are repeated at the ecotones between desert and non-
desert environments at opposite ends of the country.
Cherlin (1983b) suggested there is an important positive relationship
between the mean number of body scales (longitudinal ventral rows and
transverse dorsal rows) of Echis species and climate (mean relative humidity X
average annual temperature/average annual daily range of air temperature).
However, his formula has never been tested by analysis of meristic characters
within a single species.
Scrutiny of the isophenic maps of geographic variation in the scalation of
Echis in Pakistan suggest that these patterns may indeed be correlated with
climatic condition. Thus, higher scale counts and lighter colors are often
associated with arid environments (particularly the Cholistan-Thar Desert
[=NW Rajasthan sample, in part] and Chagai Desert [=Nushki sample, in
part]; see Fig. 25 for major vegetation zones in Pakistan; Table 5 relates our
sample areas to these vegetative zones). We thus turn our attention to an
analysis of these characters in reference to climatic factors.
Spearman Rank Correlation values (= Rs) and corrected z values were
calculated for the relationship between a number of the more distinctive
geographic patterns of meristic characters and certain climatic factors. The
meristic characters selected were the mean number of dorsal spots, ventrals,






AUFFENBERG & REHMAN: THE GENUS ECHIS


and gulars; the climatic factors were mean annual temperature, and the mean
daily maximum and mean daily minimum temperatures for April, May, June
and July (separately). These are the months when saw-scaled vipers are most
active in Pakistan (Rehman, MS). We also obtained correlation for these
important meristic characters and the Cherlin climatic "equitability" formula.
Our results show there is no significant correlation between the mean
number of ventral scales and Cherlins climatic value for the same geographic
points (Rs 0.10, z 0.36). However, we found a significant negative correlation
(p 0.02) between mean ventrals and mean annual rainfall at each sample site
(Rs 0.35, z 1.22, p 0.02). Samples with low rainfall areas are not only arid, but
hot in summer, and we believe that temperature is, in this instance, more
important than precipitation (many desert areas in Pakistan fail to get rain for
several successive years). The correlation between ventrals and mean annual
daily maximum temperature is not significant (Rs 0.06, z 0.21, p < 5%). Nor is
there a significant correlation between mean daily maximum temperatures and
mean ventral number in April (0.22, z 0.45 respectively). However, the same
correlations for June (0.33, 1.15) and July (0.32, 1.12) are both barely
significant at the five percent level. For May, the correlation shows a higher
significant value (0.53, 1.84, p < 4%). In general, the mean daily minimum
temperatures shows a similar range of from poor to fair correlations with mean
ventral number (May Rs 0.43, z 1.46, June Rs 0.18, z 0.64). July daily minimum
temperatures shows the best correlation of all (Rs 0.63, z 2.18, p < 0.05).
These minimum temperatures occur during the night, when most individuals
forage (Rehman, MS).
The number of dorsal blotches also show a geographic pattern
significantly correlated with the mean number of ventral scales (Rs 0.79, z 3.96,
p < 0.02), but a consistently low correlation with mean daily maxima or
minima. Nor is there a good correlation between any climatic factor and the
mean number of gulars. Thus of all the significant geographic patterns in
meristic characters, ventral scales appear to show the best correlation with
climatic factors.
Figure 26 provides data on the distributional pattern of mean summer
high temperatures. It shows that within the Pakistan area the highest mean
summer highs are achieved in western Baluchistan and NW Rajasthan--exactly
those areas in which most of the mean scale counts are highest. Within the
range of Echis carinatus in Pakistan, the mean daily minimum summer
temperatures occur in areas of deciduous and mixed, rather than thorn brush
habitats (Table 5, Fig. 26). These areas also tend to have proportionately
lower scale counts. The same pattern can be demonstrated in dorsal and
ventral color density, for the colors are least intense in desert areas and most
intense in forested areas (a typical, world-wide pattern). In the Indian
peninsula, Echis carinatus from the relatively cooler (Fig. 26), more moist and






306 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


/
FJ,-


45


/1


I
Figure 26. Mean high summer temperatures in the Indian Subcontinent (modified from Atlas of
Pakistan 1986 and Koteswarum 1974).






AUFFENBERG & REHMAN: THE GENUS ECHIS


more heavily forested Goa-Kerala area are significantly darker and with fewer
scales than those populations in the remainder of the peninsula.
In his study of the effects of temperature on the scalation of Echis species,
Cherlin (1983b) developed what he termed a "dorsal scale index." It is
determined by multiplying the number of ventrals by the number of middorsal
scale rows. This is a very useful statistic, for it provides a comparative
statement of the relative number of scales covering the back. From a
physiological standpoint it is useful in providing an index of the amount of skin
between the scales, for this is where water loss occurs, and thus evaporation
takes place. Snakes with more scales have proportionately more skin exposed
per body surface. The more evaporation the greater potential for lowering
high body temperatures during near-lethal limits at the high end. The
physiological basis of the importance of Cherlin's index has not yet been
unquestionably demonstrated. However, when we plot the values
geographically we find that the geographic pattern of mean values follows the
common one shown previously for ventral and dorsal scales; i.e., highest values
in northwestern Baluchistan (Chagai Desert) and northwestern Rajasthan and
adjacent Pakistan (Cholistan-Thar Desert), the lowest in NWFP and along the
foothills of the Himalayas. Statistically significant differences in the means of
adjacent populations occur between Nushki-Quetta, Karachi-Kirthar, Multan-
NW Rajasthan, Lahore-NW Rajasthan, and Rajasthan-NW Rajasthan samples.
There is no statistical correlation between Cherlin's scale index and mean daily
maximum summer temperatures (R2 0.01), or the scale index and mean
minimum summer temperatures (R2 0.03).
While critical thermal maximum value has not yet been determined for
Echis carinatus, we presume it is close to 400C, as such values are for most
snakes (Lillywhite 1987), regardless of their environmental preferences
(Spellerberg 1972). Thus, during the hottest seasons of the year it is only at
night when temperatures are sufficiently reduced to allow active foraging on
the surface (30-370C Rehman, MS). It follows that the daytime highs are
probably the most critical during the hot seasons, and though the snakes are
then usually in abandoned rodent burrows (Rehman MS), daytime burrow
temperatures sometime exceed 350C in arid parts of Pakistan (Auffenberg
MS).
Brown (1957) defines centrifugal speciation as that process by which
populations central to the point of speciation for that subspecies evolve much
faster than peripheral populations. Thus the most distant populations receive
adaptive genes more slowly, determined by the rate of gene flow in that
direction. The more distant a population, the more time it will take for an
evolutionary change to reach it. Populations completely isolated (such as Echis
carinatus astolae, on an island) may never acquire newer adaptations, and thus
remain "primitive" with respect to those adaptations. We believe this







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


hypothesis best explains most of the kind and degree of geographic variation
witnessed in this study.
Our studies on Echis populations in the Indo-Pakistani area suggest there
are five recognizable major adaptive centers. These are Transcaspia, Iranian
(out of our main area of study), Astola Island (minor), Indo-Gangetic Plain,
Himalayan foothills, and Cholistan-Thar Desert. We believe that the gene flow
from these four adaptive areas account for most of the geographic patterns
seen in the characters we studied. The only other factor considered important
is the pattern mosaic in the Indus Delta, which is believed due to distributor
shifts during the Pleistocene and possibly Post-Pleistocene.




LITERATURE CITED

Auffenberg, Walter. MS. Behavioral Ecology of the Bengal Monitor.
Beg, A. R. 1983. Wildlife habitats of Pakistan. Pakistan Forest Inst., Peshawar. 56 pp.
Brown, W. L. 1957. Centrifugal speciation. Quart. Rev. Biol. 32:247-77.
Cherlin, V. A. 1981. [The new saw-scaled viper Echis multisquamatus sp. nov. from southwestern
and middle Asia]. Proc. Zool. Inst. Acad. Sci. USSR 101:92-5. (in Russian).
1983a. [New data on taxonomy of snakes of the Echis genus]. Vesnik Zool. 1983:42-6.
(in Russian).
S1983b. [Dependence of scale pattern in snakes of the genus Echis from climatic factors].
Zoologicheskii Zh. 62:252-8. (in Russian).
Christman, S. P. 1980. Patterns of geographic variation in Florida snakes. Bull. Florida State
Mus., Biol. Sci. 25(3):157-256.
Constable, J. D. 1949. Reptiles from the Indian peninsula in the Museum of Comparative
Zoology. Bull. Mus. Comp. Zool. 103:59-160.
De Terra, H., and G. E. Hutchinson. 1936. Data on Post-glacial climatic changes in
northwestern India. Curr. Sci. 5:5-10.
and T. T. Patterson. 1939. Studies on the Ice Age in India and associated human cultures.
Carnegie Inst. Publ (493):1-115.
Edgren, R. A. 1961. A simplified method for analysis of dines; geographic variation in the
hognose snake, Heterodon platyrhinos Latreille. Copeia 1961 (2):125-32.
Erikkson, K. G. 1959. Studies of the dry bed of the River Ghaggar at Rang Mahal, Rajasthan,
India. Acta Archeol. Lundensia, ser. 4 (3):1-63.
Flam, L. 1986. The Indus River and the Arab Period in Sindh. Sindhological Stud. Inst.
Sindhology, Univ. Sindh, Jamshoro Summer 1986:5-13.
Geoffroy, Saint Hillaire, E. 1827. Description de l'Egypte. Histoire naturelle, Reptilia. Paris. 344
p.
Khan, M. S. 1980. Affinities and zoogeography of herptiles of Pakistan. Biologia 26(1/2):113-
171.
Koteswarum, P. Research in the humid tropical Asia. 12th National Congress UNESCO. pp. 1-
30.
Lillywhite, H. B. 1987. Temperature, energetic, and physiological ecology. Pp. 422-477 in R. A.
Seigel, J. T. Collins, and S. S. Novak (eds.). Snakes: Ecology and Evolutionary Biology.
MacMillan Co., New York. 529 p.
Marshall, J. 1931. Mohenjo-daro and the Indus Valley Civilization. MacMillan Co., London.
244 p.
Mertens, R. 1969. Die Amphibien und Reptilien West-Pakistans. Stuttgarter Beitr. z.
Naturkunde (197):1-96.
Oldham, R. D. 1886. On probable changes in the geography of the Punjab and its rivers. J.
Asiat. Soc. Bengal (55):322-43.







AUFFENBERG & REHMAN: THE GENUS ECHIS


Schneider, J. G. 1801. In B. G. E. Lacepede, Naturgeschichte der Amphibien. Vol. 4,:156-256. J.
M. Bechstein, Weimar.
Smith, P. W. 1956. A blotch-count gradient in snakes. Herpetologica 12(3):156-60.
Spellerberg, I. F. 1972. Temperature tolerances of southeast Australian reptiles examined in
relation to reptile thermoregulatory behavior and distribution. Oecologia 9:23-46.
Stemmler, 0. 1969. Die Sandrasselotter aus Pakistan: Echis carinatus sochureki subsp. nov.
Aquaterra 6:118-24.
Wilson, L. D. 1970. The racer Coluber constrictor (Serpentes: Colubridae) in Louisiana and
eastern Texas. Texas J. Sci. 22(1):67-85.






TABLES


Table 1. Geographic variation in number of mean ventral scales in those adjacent populations
(Pakistan and border sites) showing significant differences in their means.


Sample Pairs Ventrals df* p*


Afghanistan-Quetta 181-165 22 < 0.001
Quetta-Nushki 165-183 22 < 0.001
Thar Parkar-Gujarat 158-162 27 < 0.01
Thar Parkar-Nagar Parkar 158-168 25 < 0.01
Multan-NW Rajasthan 165-181 10 < 0.03
Lahore-NW Rajasthan 162-181 8 < 0.05
NW Rajasthan-Rajasthan 181-158 14 < 0.01


* df, degrees of freedom; p, probability on basis of Student's t-test.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Table 2. Variation in ventral and subcaudal scales among siblings in single broods of Pakistan
and Indian Echis.


Ventrals Subcaudals
Locality

(N) OR* Mean SD* OR* Mean SD*


Goa, India (4) 140-143 142.2 1.7 24-26 25.3 1.0
Kerala, India (3) 149-152 150.3 1.3 29-30 29.7 0.6
Thar Parkar, Pakistan (7) 154-160 157.0 1.9 25-27 26.3 1.0
Thar Parkar, Pakistan (12) 154-162 158.6 2.6 23-30 26.4 2.5
Karachi, Pakistan (12) 159-173 164.8 4.7 27-35 29.1 2.1
Shadadkot, Pakistan (20) 162-177 169.6 5.0 26-36 29.4 2.4
Quetta, Pakistan (8)) 162-171 165.7 3.1 24-34 28.4 3.7



* OR, Overall range; SD, standard deviation.


Table 3. Geographic variation in number of dorsal spots in those adjacent populations (Pakistan
and border sites) showing significant differences in their means (p < 0.5) (abbreviations as in
Table 1).


Sample Means df p


Nushki-Quetta 39-35 22 < 0.001
Quetta-Shadadkot 35-31 19 < 0.1
Karachi-Ratan kot 36-31 31 < 0.001
Karachi-Mekkran Coast 36-38 29 < 0.25
Shah Bundar-Thatta 37-33 302 < 0.01
Shah Bundar-Thar Parkar 37-40 33 < 0.1
Shah Bundar-Gujarat 37-33 27 < 0.1
Thar Parkar-Thatta 40-33 310 < 0.1
Thar Parkar-Gujarat 40-32 27 < 0.01
Thar Parkar-Central Sindh 40-32 22 < 0.1
Thar Parkar-Rajasthan 40-33 15 < 0.5
NW Rajasthan-Lahore 41-31 8 < 0.1
NW Rajasthan-Multan 41-34 11 < 0.1
NW Rajasthan-Central Sindh 41-32 12 < 0.1







AUFFENBERG & REHMAN: THE GENUS ECHIS


Table 4. Geographic variation in degree of darkness of ventral spots in those adjacent
populations (Pakistan and border sites) showing significant differences in their means (p < 0.5)
(abbreviations as in Table 1).


Sample Means df p


Nushki-Quetta 1.4-0.9 22 < 0.001
Karachi-Mekkran Coast 2.6-1.5 28 < 0.001
Shadatkot-Cent. Sindh 0.7-2.0 15 < 0.010
Lahore-NW Rajasthan 2.0-0.3 8 < 0.050
NW Rajasthan-Cent. Sindh 0.3-2.0 11 < 0.050
Rajasthan-Thar Parkar 1.0-1.9 27 < 0.025
Thar Parkar-Nagar Parkar 1.9-1.0 23 < 0.025
Gujarat-Nagar Parkar 1.8-1.0 22 < 0.010
Karachi-Kirthar 2.6-1.3 32 < 0.010








Table 5. Echis sample areas and the general vegetative zones in which each is located.


Sample Area Zone Sample Area Zone


Multan Thorn Brush Islamabad Subtrop. Pine
Cent. Sindh Thorn Brush Lahore Trop. Decid.
Thar Parkar Thorn Brush NWFP Trop./Subtr.
Pine
Thatta Thorn Brush Quetta Temp.
Jun./Pine
Karachi Thorn Brush Nagar Pakar Trop. Decid.
Ghost Thorn Brush Zabol Temp. Desert
Mekkran Coast Thorn Brush* Afghanistan Temp. Desert
Panjgur Thorn Brush Nushki Temp. Desert
Rattankot Thorn Brush NW Rajasthan Trop. Desert
Shah Bundar Thorn Brush Shadadkot Trop. Desert
Rajasthan Thorn Brush Gujarat Thorn Brush
Parachinar** Temp. Pine


* Mekkran Coast thorn brush is rather different from that of the remaining Indus Valley types
(see Beg 1983 for review).
** Included in NWFP throughout this publication, but separated here because Echis carinatus
from Parachinar are very dark.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


APPENDIX I


Locality data and present museum location for specimens examined during this study.


Locality


Museum


nr. Abadan, Iran
Ajmere, Rajasthan, India
Allahabad, Gujarat, Iran
Amballa (100 mi. S), India
Aram, Fars, Iran
Aschabad, Turkmen SSR
Astola Isl., Baluchistan, Pakistan
Badin, Sindh, Pakistan
Balchany, Transcaspia SSR
Bampur, Iran
Bamrud, Iran
Bangalore, Karnataka, India
Bhavnagar, Gujarat, India
Bhit Poochari, (Punjab ?), Pakistan
Bombay, Maharashtra, India
Bushire, Iran
nr. Cancona, Goa, India
Cape Monze, Sindh, Pakistan
Chabar, Persian Gulf, Iran
Chumbum, Tamil Nadu, India
Colombo, Sri Lanka
Deesa, Gujarat. India
Dera Ismail Khan, NWFP, Pakistan
Dhabiji, Sindh, Pakistan
Dir, NWFP, Pakistan
Dureji, Baluchistan, Pakistan
Durun, Transcaspia SSR
Duschak, Transcaspia SSR
Fars, Iran
Fatehjang, Punjab, Pakistan
Ft. Sandeman, Zhob, NWFP, Pakistan
Goa, India
Gondal. Gujarat, India
Gosht, 42 mi. N Dizak, Seistan, Iran
Gumtur Dist., A. P., India
Gurshir, Iran
Hab Chouki, Baluchistan, Pakistan
Henjam Isl., Persian Gulf, Iran
Hinadan crossing, Baluchistan, Pakistan
Hingol Natl. Park, Baluchistan, Pakistan
Hingolgadh, Gujarat, India
Hungo, Kohat, NWFP, Pakistan
Hyderabad, Sindh, Pakistan
Islamabad
Jabalpur, Madha Pradesh, India


SMF
ZSI
ZSI
ZFMK, MCZ
FMNH
SMF
SMF
ZSD
ZIN
SMF
ZIN?
ZSI
BNHS
RSM
BNHS
ZSI
ZSI
RSM
BNHS
BNHS
SMF
BNHS
BNHS
ZSD
PMNH
RSM
ZIN?
SMF
CAS
PMNH
BNHS
UF
BNHS
ZSI, BNHS
ZSI
ZIN?
AMNH, RSM
BNHS
AMNH
UF
BNHS
UF, ZSD
AMNH
UMMZ, PMNH
ZSI







AUFFENBERG & REHMAN: THE GENUS ECHIS


Locality


Jaisalmere, Rajasthan, India
Jamsanda (nr. Bombay) Maharashtra, India
Jask, Persian Gulf. Iran
Jati, Sindh, Pakistan
Jiwani, Baluchistan, Pakistan
Jhang, Punjab, Pakistan
Jungshai, Sindh, Pakistan
Kacha, Baluchistan, Pakistan
Kalagan, Baluchistan, Pakistan
Kalicut, Kerala, India
Kanpur, U. P., India
Karachi Dist., Sindh, Pakistan
between Karman amd Shiraz, Iran
Kaur, Baluchistan, Pakistan
Kaweit, Iran
Kerman, Iran
Kerula, Punjab, India
Khandahar, Afghanistan
100 mi SW Khandahar, Afghanistan
Khandala, Afghanistan
Khar Centre, Sindh, Pakistan
Khokhrapur, Baluchistan, Pakistan
Khudsil Khan (nr. Quetta), Baluchistan, Pakistan
Khuzistan, Iran
Killi Mangal, Baluchistan, Pakistan
Killi Jamaldini, Baluchistan, Pakistan
Kirthar Natl. Park, Sindh, Pakistan
Kojdar, Mekkran Coast, Iran
Krasnowodki, Transcaspia SSR
Lal Suharna Natl. Park, Punjab, Pakistan
Larkana, Sindh, Pakistan
Madras, Tamil Nadu, India
Makli, Sindh, Pakistan
Mand, Baluchistan, Pakistan
nr. Manjhand, Sindh, Pakistan
Mango Pir, Sindh, Pakistan
Matiari (nr. Hyderabad), Sindh, Pakistan
Mednapur, West Bengal, India
Megas, E. Iran
Mekkran Dist., Baluchistan, Pakistan
Miranshah, NWFP, Pakistan
Mithi, Sindh, Pakistan
Mohen-jo-daro, Sindh, Pakistan
Multan, Punjab, Pakistan
Munbhum Dist., Bihar, India
Nabisar Nota, Sindh, Pakistan
Nagaur Dist., Rajasthan, Pakistan
Nasrie, Iran
Nazirabad, Iran
Nellore Dist., Andra Pradesh, India
Nushki, Baluchistan, Pakistan
Panjgur, Baluchistan, Pakistan
Patho Pass (Pab Hills), Baluchistan, Pakistan


ZSI
USNM
BNHS
RSM
SMF
UMMZ
RSM
BNHS
ZSI
BNHS
FMNH, USNM
SMF, ZSD, NMW, UF
ZSI
BNHS, BMNH
BNHS
ZIN?
FMNH, MCZ
ZFMK
CAS
FMNH
UF
SMF
ZSD
NMBA
SMF
SMF
ZSD
BNHS
USNM
UF
BNHS, UF
BNHS, MCZ
ZSD, UF
BNHS
AMNH
ZSD
ZSD
FMNH, MCZ
ZIN
BNHS
BNHS
SMF
SMF
ZSD, UF
ZSI
RSM
ZSI
ZIN
BNHS
BNHS
SMF, RSM
SMF, ZSM
RSM


Museum







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 35(5)


Museum


Persepolis, Iran
Peshawar, NWFP, Pakistan
Point Calimere, Tamil Nadu, India
Pol-Absinch, Fars, Iran
Poona, Maharashtra, India
Pugal, Rajasthan, India
Puttur (Chittor) Andra Pradesh, India
Quetta, Baluchistan, Pakistan
Quilon, Kerala, India
Ratan Kot, Sindh, Pakistan
Saman (Dasht Dist.), Baluchistan, Pakistan
Sehirabad, (nr. Buchara, Afghanistan ?)
Schechradj, Baluchistan, Pakistan
Shah Bundar. Sindh, Pakistan
Shadadkot, Sindh, Pakistan
Soman Dist., Baluchistan, Pakistan
Spinkaras, Baluchistan, Pakistan
Sirohi, Rajasthan, India
nr. Stalinabad, Uzbekistan SSR
Thar Parkar, Sindh, Pakistan
Thatta, Sindh, Pakistan
Trivandrum, Kerala, India
Turkmen SSR
nr. Urak, Baluchistan, Pakistan
Valpoi, Goa, India
nr. Zabol, Iran
Zhor, Sindh, Pakistan
nr. Zirkuch, E. Iran


FMNH
UF, ZSD
BNHS
FMNH
ZSI
ZSI
USNM
SMF, ZSD, RSM
AMNH, UF
SMF
ZSI
ZIN
ZMFK
CAS
SMF
ZSI
SMF
ZSI
NMBA
SMF
UF, ZSD, AMNH, SMF, RSM
UF, UMMZ
SMF, ZMFK
SMF
ZSI
SMF, FMNH
ZSD
ZIN?


Locality








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