D o l . J. Linn. SOC.,57; 1-57. With 3 plates and 3 3 figures
August 1975
The taxonomy, distribution and faunal succession
of B u l h i n u s (Pdmonata: Enidae) in Israel
JOSEPH HELLER
Department of Zoology, The Hebrew University, Jenisalem
Accepted for publication September I974
The taxonomy, intraspecific variation and distribution of each of the eight Israeli species of the
rock-dwelling landsnail Buliminus is described, with a new subspecies labrosus jiftliki. The
general biology of th e genus is briefly described, and a key to all species and subspecies is given.
Intraspecific and interspecific variation is partly associated with different environmental
conditions. In the more h o t regions of their distribution areas, the snails have a narrow shell, a
smaller aperture, a more smooth shell surface and thick lips. The possible adaptive significance
of these characteristics in h o t regions is discussed.
The distribution limits of many species closely coincide with both isohyets and types of
substratum, suggesting that rainfall and substratum are the most important physical factors
determining dismbution. Vegetation in the Mediterranean region could perhaps influence such
aspects of th e distribution as the choice of northern or southern slopes.
It is suggested th at in Israel, in each climatic zone and within the limits determined by these
physical fa cto n , a faunal succession is today in operation. In the mediterranean, steppe, steppe
periphery a nd arid regions, species which evolved o r arrived recently are displacing more ancient
elements. As compared with the ancient fauna, the new species have less rigid ecological
requirements and are more capable of adjusting themselves to severe environmental conditions
by altering their shell proportions. The last relics of the ancient fauna survive in the Judean
Hills and along the isohyet 7 0 mm, on the brim of the extreme desert.
In 1. labrosus the presence of a reverse cline, in both shell and genitalia, near the borderline
with 1. spirectinus, and, in addition, the presence along t h e borderline of a few intermediate
individuals, suggests that 1. labrosus is “rolling back” 1. spirectinus b y keeping u p a narrow belt
of hybridization.
CONTENTS
. . . . . . . . . . . . . . . . . .
Introduction
General biology of Buliminus
. . . . . . . . . . . .
Materials a n d methods
. . . . . . . . . . . . . .
Key t o the Israeli species and subspecies of Buliminus . . . . . .
Taxonomic descriptions . . . . . . . . . . . . . .
. . . . . . . . . . . .
Genus Buliminus Beck
labrosus Olivier
. . . . . . . . . . . .
1. labrosus Olivier
. . . . . . . . . . .
1. jiftlikii ssp. nov.
. . . . . . . . . . .
1. spirectinus Bourguignat
. . . . . . . . .
diminutus Mousson
. . . . . . . . . . .
. . . . . . . . . .
therinus Bourguignat
alepensis Pfeiffer . . . . . . . . . . . .
a. marsabensis Westerlund
. . . . . . . . .
1
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29
J. HELLER
2
lamprostatus Bourguignat
sinaiensis Heller
. .
glabratus Mousson
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negevensis Heller . .
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Discussion
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Taxonomic remarks
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lntraspecific and interspecific clines
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Ecological factors correlated to distribution
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Climate
Substratum
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Vegetation
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Succession of Buliminus in Israel
. . . .
Steppe region . . . . . . . .
Steppe periphery . . . . . . .
DeadSea region
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Mediterranean region
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Acknowledgements
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References
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30
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54
INTRODUCTION
Landsnails are particularly suitable for zoogeographic research. They are easy
to collect, and, moreover, the empty shells sometimes suffice for the evidence
required. Accurate distribution-maps can therefore be compiled, which are an
important aid in interpreting the environmental factors which determine each
distribution.
On the other hand, Israel is particularly well suited for researches in
ecological and historical zoogeography. It is situated on the dispersal route of
animals moving between Asia and Africa. Further, it is a meeting point between
Mediterranean, steppic and desert conditions (terms as defined by Atlas of
Israel, 1956). As a result, it has been subject to many faunal invasions (Haas.
1952; Steinitz, 1954; Tchernov, 1965).
The landsnail Buliminus Beck (Enidae) is a very common Israeli pulmonate
which successfully inhabits a very wide range of habitats, from Hermon (over
1500 mm precipitation) to the Negev (25 mm rain). Its species form a natural
group of morphologically closely related species, differing from one another
mainly in shell proportions and sculpture, and in details of the reproductive
system. Its wide distribution and large number of species (Pallary, 1929;
Forcart, 1940) make Buliminus well suited for studies in zoogeography.
Outside Israel, the Buliminus species of Syria and Lebanon have been described
by Germain (1922) and Pallary (1929, 1939), those of Turkey by Forcart
(1940) and those of the Iranian Plateau by Biggs (1937, 1959).
This paper will be concerned with the taxonomy, distribution and
intraspecific variability of the Israeli species of Buliminus, all of which are
illustrated in Plate 1 . It will be shown that, whilst some Israeli species are found
also in Syria and Lebanon, others, as far as we know, do not occur outside
Israel. In the discussion an attempt will be made to determine the physical
factors influencing the distribution of these species and then, partly upon this
basis, to follow the faunal succession of Buliminus and some other genera in
Israel.
GENERAL BIOLOGY OF BWLIMNWS
Buliminus is a rock-dwelling genus which inhabits rock crevices and the
undersides of stones. It is active in winter and the early spring, the snails
TAXONOMY OF BULIMINUS IN ISRAEL
3
emerging from aestivation with the onset of the first rains in OctoberNovember. The snails are nocturnal, though sudden showers occasionally bring
them out also during the day.
By sampling the species at fortnight intervals over a period of two years
(Heller, in prep.) it was found that during summer, when the snails are in
aestivation, the follicles of the hermaphrodite gland are packed with sperm and
ripe ova; within a fortnight of the first rain they are almost empty, and they do
not fill again till late in the following summer. This suggests that mating occurs
only once in a year, immediately after the first rain. Eggs are laid on the
undersides of rocks and within crevices, as the snails usually do not bury into
the soil in the manner that slugs and many Helicidae do. Juveniles appear
within about a fortnight. They complete their shell growth within two years,
after which sexual maturity is reached.
I inspected the stomach contents of all Israeli species. In mediterranean,
steppe and desert species alike it consisted of the remains of dead leaves and
stems, spores of saprophytic mushrooms, pollen and awns of cereals, and
occasional bits of soil. Apparently the snails feed upon decaying vegetable
matter which they collect from the upper layers of the soil.
At the end of spring (March-April in arid zones April-May in Mediterranean
ones) the snails aggregate into certain rock crevices and aestivate. Ifi one such
crevice, in the Lower Galilee, I found 63 aestivating snails and 76 empty shells.
The many empty, undamaged shells found in this and in other crevices suggest
that dessication is a major cause of mortality, and that many snails do not
survive the hot, dry Israeli summer.
The most important predators are nocturnal rock-dwelling rodents,
especially the spiny mouse Acomys caliirinus, which depends upon the water
content of snails t o balance its water regime (Shkolnick & Borut. 1969). To
extract the snail the rodent holds the shells in its front paws, bites through the
apex and then works its way downwards towards the aperture. In front of one
rodent’s burrow on Mt. Carmel I found 120 such damaged Buliminus shells.
Acomys is found throughout the distribution area of Buliminus in both
mediterranean, steppe and desert areas. Thrushes appear not to be as important
predators as rodents, perhaps because they hunt during the day, when the snails
are concealed beneath stones. I never found empty Buliminus shells near thrush
anvils in Israel.
MATERIALS AND METHODS
This taxonomic study is based upon the inspection of 4000 shells and the
reproductive system of 200 animals. Measurements of the shell are illustrated in
Fig. 1, nomenclature of the genitalia in Fig. 2. All measurements are in
millimetres, accurate to 0.1 mm.
The distribution maps are based partly upon material collected by Professor
G. Haas over the past 40 years and partly upon a field survey which was carried
out by me from 1968 to 1971. Most of the material from Sinai was collected
by A. Haim during a rodent survey. The area covered by this survey is bordered
by the 1967 cease-fire line, as presented in the distribution maps by a dashed
line. Collecting sites were usually 4-5 km apart but in the Negev and Sinai
distances were greater, up to 1 5 km. General maps or remarks concerning the
distribution outside Israel are based upon records in literature and upon the
J. HELLER
4
c--
M i n D-
Figure 1. Shell measurements used in this work. LW, Lip width; Max. D., maximum shell
diameter; MD, mouth diameter; MH, mouth height; Min. D., minimum shell diameter; SH, shell
height.
5 mm
B
01
/
Figure 2. Nomenclature of t h e reproductive organs (specimen of 1. Zabrosus from Elon, Upper
Galilee). A, Penial appendix; B, bursa; C , penial flagellum; Db, ductus bursae; Di, diverticle of
the receptaculum seminis; E, epiphallus; Fe. epiphallian flagellum; Ped, pediculus; Pen, penis;
Ra, appendicular retractor; Rp, penial retractor; U, uterus; V,vagina; Vd, vas deferens.
TAXONOMY OF BWLIMINUS IN ISRAEL
.'*errno"
i
PO
Figure 3. Localities mentioned in this text. 8, Atlit; 17, Bet Zait; 40,Dimona; 34, Ein Boqeq;
31, Ein Gedi; 1 6 , Ein Hemed; 1 9 , Ein Kerem; 1 9 , Ein Turaba; 1 , Elon; 4 3 , Hamakhtesh
Hagadol; 4 1 , Hamakhtesh Haqatan; 5 0 , Har 'Arif; 4 2 , Har Avnon; 4 7 , Har Gerafon; 36, Har
Hemar; 49, Har 'Oded; 2 0 , Har Ora; 37, Har Qomot; 2 3 , Hebron; 2 5 , Jericho; 5 2 , Jerash; 1 8 ,
Jerusalem; 24, Jiftlik; 5 , Kafr Manda; 2 7 , Khirbet Za'arur; 1 4 , Kiryat Anavim; 5 3 , Kitte; 1 2 ,
Latrun; 4 5 , Ma'ale Aqrabim; 48, Makhtesh Ramon; 2 6 , Marjaris; 28, Marsaba; 15, Martyrs
Forest; 6, Merkaz Hacarmel; 4 , Meron; 33, Mezada; 10, Nablus; 46, Nahal 'Avdat; 21, Nahalin;
44, Nahal Hatira; 7 , Nahal Oren; 2 , Nahal Namer; 5 1 , Nahal Timna; 32, Nahal Zeelim; 35, Nahal
Zohar; 38, Saruhen; 3, Sasa; 1 3 , Sha'ar Hagai; 2 2 , Si'ir; 9 , Tul Karm; 1 1 , Urn Safa; 30, Wadi
Mashash.
5
J. HELLER
6
collection of the Hebrew University, not on localities inspected by myself.
Localities mentioned in the text are figured either in the relevant distribution
maps (Figs 4 , 2 0 , 22 and 26), or in Fig. 3 .
Data on the climate, substratum and vegetation in the distribution areas are
from the Atlas of Israel (1956), the Israel Meteorological Service (1961, 1967),
Ravikovitch (1969), Dan & Raz (1970), Zohary (1955), Gruenberg-Fertig
(1966) and Danin (1970).
The Buliminus collection is kept at the Zoology Department of the Hebrew
University, Jerusalem. Also deposited with the collection are tables containing
the observed range, mean and standard deviation, of each measurement. for
104 of my samples.
KEY TO ISRAELI SPECIES AND SUBSPECIES OF BULIMINUS
(Adult specimens)
Shell height is almost always more than 24 mm
. . . 2
Shell height is 24 mm or less
. . . . . . .
. 5
2. (1) The shell surface is finely granulated (best observed under x80
magnification, see Plate 1B)
. . . . . . . . . 3
The shell surface is not granulated and has a smooth striated
. . . . . . . . . . . . . .
4
surface
3. (2) Mouth height is large as compared with shell height (see Figs 1 3
and 14)
. . . . . . . . . . .labrosus Eabrosus
Mouth height is small as compared with the shell height (see
. . . . . . . labrosus spirectinus
Figs 1 3 and 14)
4.(2) Peristome more than 1.8 mm thick
. . . labrosus jiftliki
Peristome less than 1.8 mm thick
. . . . .diminutus
5. (1) Shell is glossless and its surface granulated (best observed under
x80 magnification, see Plate 1J)
. . . . . . therinus
The shell is glossy and its surface smooth or striated
. . 6
6. ( 5 ) The index SH/MH is almost always more than 2.50
. . 7
The index SH/MH is 2.50 or less . . . . . . . . 8
7. (6) The minimal shell diameter is small, as compared with shell
height (Fig. 31). The ultimate whorl usually has a palatal
lamella, seen from the outside as a white stripe
. glabratus
The minimal shell diameter is large, as compared with shell
height (Fig. 3 1). N o palatal lamella
. . .negevensis
8. (6) Peristome less than 1 .O mm thick
. . . . . .
9
Peristome more than 1.0 mm thick
. . '. . lamprostatus
9. (8) Mouth diameter almost always less than 6.6 mm
. . . 10
Mouth diameter almost always is more than 6.6 mm
. . 11
10. ( 9 ) Mouth diameter is narrow, as compared with mouth height (see
Fig. 27)
. . . . . . . . . alepensis marsabensis
Mouth diameter is large, as compared with mouth height (see
Fig. 27)
. . . . . . . . . . . . . sinaiensis
11. ( 9 ) Mouth height is almost always more than 9.5 mm
.diminutus
Mouth height is almost always less than 9.5 mm
. sinaiensis
1.
TAXONOMY OF BULIMINUS IN ISRAEL
7
TAXONOMIC DESCRIPTIONS
Genus Biiliminiis Beck
Buliminus Beck, 1837.
Petraeus Albers, 1850: 1 5 3.
Petraeus sensu stricto: Hesse, 1933: 192.
Type species. Bzllimus labrosus Olivier.
The shell is dextral, horny brown or purple, composed of 5'/2-91/2whorls. The
diameter of the protoconch is much larger than its height. The whorls of the
protoconch are continuous with the other apical whorls. The aperture has no
teeth. The reproductive system has a well developed epiphallian flagellum.
Buliminus differs from Pene Pallary in the proportions of its protoconch
(Heller, 1974), and from Ena Turton in its strongly developed epiphallian
flagellum (Hesse 193 3).
Buliminus labrosus Olivier
(Plates 1A-G and 2)
Bulimus labrosus Olivier, 1804 (2): 30, pl. 3 1 , fig. 10 A, B. type locality:
Lebanon.
Bulimus thaumastus Bourguignat, 1876: 2. From Nahr-el-Kelb and Beirut.
Bulimus exochus Bourguignat, 1876: 3. From Antilebanon, North of Jordan
River.
Buliminus (Petraeus) granulatus Westerlund, 1892: 42. From Samaria.
Buliminus (Buliminus) labrosus Zilch, 1959: 190; fig. 660.
The shell varies from cylindrical-conic t o low cylincrical and from light to
dark horny brown, sometimes purple. I t is composed of 6-7 whorls which, in
the two mediterranean subspecies 1. labrosus and 1. spirectinus, are glossless
delicately striated and have a very delicate granulation (best observed under
x80 magnification; see Plate 1B). In the more arid-dwelling subspecies 1.
jiftliki, the whorls are not granulated. The height and diameter of the
penultimate whorl are larger than those of the preceding ones The lips are
connected by a well-developed callus. The peristome is reflected outwards,
sometimes also backwards.
Measurements (mm): Shell-height
Max. diam.
Min. diam.
Mouth-height
Mou th-diam .
Shell-height/Max. diam.
Shell-height/Mouth-height
23.8 -38.9
12.0 -18.0
9.0 -14.4
12.0 -20.0
8.3 -14.1
1.75- 2.47
1.72- 2.31
The reproductive system of labrosus was described by Schmidt (1855) who
examined specimens from Lebanon and Jerusalem. Hesse (193 3) and Forcart
(1940) described the reproductive system of egregius Nagele, which Forcart
considers a northern subspecies of labrosus. In the present study over 50
specimens were examined. The following description refers to a specimen from
Elon (Fig. 2).
The albumen gland is aeamish and bent at its distal end. The pediculus is
longer than the diverticle. The combined length of the bursa and its duct is
8
J . HELLER
slightly less than the length of the diverticle. The vas deferens widens slightly
upon ascending towards the epiphallus. The epiphallian flagellum is longer than
the epiphallus. The epiphallus inserts on the penis laterally, leaving a penial
flagellum (=“caecum” of Hesse, 1933). The penis is thinner than the proximal
part of the appendix, which has a very long distal part. The penial and
appendicular retractors, of equal length, unite into a common muscle which
inserts at the diaphragma.
Synonyms
B. exochus Bourguignat (see Pallary, 1929: pl. 1, fig. 15) is a slightly more
ventricose shell of lubrosus which appears in many samples from Upper Galilee
and Hermon. Plate 2A illustrates the continuous transition within one sample
from lubrosus to exochus, because of which the latter is considered a synonym.
B. thaumastus Bourguignat (see Pallary, 1929: pl. 1, fig. 20) is a lubrosus with a
very dilated aperture. The shape of the aperture in lubrosus is highly variable
and many transitions to a “thaumastus” aperture were found in my collection.
B. grunulutus Westerlund (type specimen lost, Dr H. Walden, pers. comm.) is
characterised by its finely granulated shell. As nearly all Israeli labrosus are
finely granulated, this too is considered a synonym.
Distribution (Figs 4 to 6)
The species is distributed on the lower slopes of Mount Hermon and in the
hilly regions of Upper Galilee, Lower Galilee (west of the watershed) and
Mount Carmel; further south, it is very common in Samaria and the Judean
Hills, with an isolated population on Mt. Gilboa and a few populations along
the coast. The southernmost point is Si’ir, north of Hebron.
Ninety per cent of the localities are within the limits of the 500 mm isohyet.
Only in northeastern Samaria do populations of 1. lubrosus extend into more
arid regions, down to 200 mm, and, as will be described later, this is
accompanied by alterations in the size, proportions and sculpture of the shell.
Throughout the entire distribution area the rainy season is at least six months
long (calculated by combining maps 4/3a and 4/3f in Atlas of Israel, 1956).
B. lubrosus inhabits limestone and dolomite almost exclusively. It has only
twice been found upon kurkar (a hardened, calcareous dune sand), and never
upon chalkstone or basalt, or in alluvial soils. This is illustrated in Fig. 6 where,
as terra-rossa is the soil derived from limestone and dolomite, the distribution
map of lubrosus is mounted on the map of this soil. Taking Mount Carmel and
Lower Galilee (where the soil map is most complex and any correlation to
substratum would therefore be more emphasized), all localities but one are
upon terra-rossa. The one exceptional locality is on kurkar, and is maritime. A
similar situation exists in other parts of the distribution area.
The species inhabits the Mediterranean phytogeographic region, as defined
by Zohary, 1955. It is more common in open landscapes than in oak maquis,
and correspondingly more common on south than on north-facing slopes. On
Mount Carmel, for example, the two slopes of Nahal Oren were each searched
for one hour and the following material was found:
South-facing slope: 136 empty shells, 18 live snails.
North-facing slope: 30 empty shells, 6 live snails.
TAXONOMY OF BULIMINUS IN ISRAEL
Figure 4. Distribution of labrosus and diminurus in Israel. Most localities of 1. labrosus and of 1.
spirectinus are within the limits of the 500 rnrn isohyet. 0 , 1. labrosus; 0 , 1. spirectinus; a,
localities at which intermediates between 1. labrosus and 1. spirectinus were found: A, 1. jiftiiki;
*. diminutus.
9
J . HELLER
10
Figure 5 . General distribution of labrosus
subspecies of labrosus.
( 0 ) ; 0,
egregius, which Forcart ( 1 9 4 ) considers a
i
Figure 6. Distribution of Iabrosus ( 0 ) in Lower Galilee and o n Mt. Carrnel, as compared with
the distribution of terra rossa soil (stippled areas).
TAXONOMY O F BULIMINUS IN ISRAEL
11
Thus the hot, dry habitat has 3-4 times more snails than the cool. damp one.
The difference in proportions between the empty shells (1 : 4) and the live
snails (1 : 3) could be because of the somewhat different collecting method
involved in each case. For the first half hour only empty shells were gathered,
from those scattered on the ground and from those in front of rodents’
burrows. For the next half hour only live snails were collected, by overturning
rocks.
Figure 5 presents the distribution of labrosus as recorded in the literature
(Blankenhorn, 1889; Germain, 1921; Pallary, 1929, 1939; Forcart, 1940; Haas,
1952; Najim, 1959) and in the Hebrew University’s collection. Outside Israel
the species is found in Lebanon, Syria and Jordan. From Southern Turkey t o
the Caspian sea it is represented by egregius, which is considered by Forcart
( 1 940) to be a subspecies of labrosus.
hi traspecific variability
Three subspecies are found in Israel (Fig. 4): 1. labrosus, 1. jiftliki ssp. nov.
and I. spirectinus.
B. labrosus labrosus Olivier
(Plates 1A-E and 2 )
The shell varies from spindle-shaped t o ventricose. The aperture is large in
comparison to shell-height, the ratio shell-height/mouth-height being usually
less than 2.00 (see Figs 1 3 and 14). In the reproductive system the penis,
appendix and pediculus are elongate, and a penial flagellum is nearly always
present (Figs 2 and 11B-C).
Measurements (mrn): Shell-height
Max. diam.
Min. diam.
Mouth-heigh t
Mouth-diam.
Shell-height/Max. diam.
Shell-heigh t/Mouth-height
24.7
12.0
9.7
12.5
-36.2
-18.1
-13.6
-20.0
9.0 -13.2
1.75- 2.43
1.72- 2.31
This subspecies is found in Hermon, Galilee, Camel, Gilboa and Samaria
(Fig. 4). In Upper Galilee the shells are large, massive, and the peristome is very
thick and curled around itself (Plates 1C and 2B), resembling Olivier’s holotype
of labrosus from Beirut. Another form found in Upper Galilee resembles a
small labrosus var. jordani Charpentier: the shell is cylindric, the sutures are
shallow, the peristome is thin and not curled around itself (Plate 1D). The
transition between these two forms is usually quite continuous within a sample
(Plate 2C), but may occasionally be abrupt, between samples even only 5 km
apart. Table 1 shows that samples from Sasa and Har Meron ( 5 km apart and
with similar ecological conditions) have n o overlap whatsoever in five
shell-measurements.
In Lower Galilee the shells are usually roundish, a ventricose form which
resembles labrosus var. kervillei Germain from Syria (Plate 1E). The intergradation between this and the cylindric form of Upper Galilee is quite gradual
and continuous (Plate 2D). In comparison with the samples of Upper Galilee,
12
3
3
3
3
N
3
3
3
3
3
m
m
m
OI
TAXONOMY OF BCJLIMINUS IN ISRAEL
13
Figure 7. Mean shell-height of 1. labrosus in Upper and Lower Galilee.
those of Lower Galilee are much more homogeneous (Fig. 7), perhaps because
the latter has a much more uniform topography and climate than the former.
In Mount Carmel, the samples are again extremely variahle (see Fig. 8 and
Table 1). Ventricose, cylindric and narrow shells, with many intermediates,
were found together within a sample (Plate 2E), in several samples. Actually
some of these narrow, small-mouthed shells have a shell-height/mouth-height
ratio higher than 2.00 and thus do not fit the description of the subspecies as
given above if one relies upon shell characters alone. Unfortunately the
morphology of the genitalia of such slim-shelled snails is unknown t o me, as no
live specimens were found.
In northern Samaria the samples are more uniform. The shells are, generally,
cylindric. Shell-height rises clinally south-eastwards, from Tul Karm towards
Nablus (Fig. 9 ) . Other measurements change only slightly. From Tul Karm
southwards the shells become wider and the shell’s mouth becomes relatively
larger (Fig. lo), as the cylindric form of northern Samaria gives way t o a very
ventricose form (Plate 1 E) with rapidly expanding whorls. In the reproductive
system the pediculus and epiphallian flagellum become exceptionally long and
a well-developed penial flagellum is present (Table 2, and Fig. 11C).
B. labrosus jiftliki ssp. nov.
(Plate 1F)
Diagnosis. B. 1. jiftliki differs from the nominate subspecies in its thick lip and
smooth, ungranulated shell.
J . HELLER
14
n
180
2.00
220
240
'y;,,~
Figure 8. Frequency of the ratio shell-heighthnaximum diameter in I. labrows from Kafr
Manda, Lower Galilee (dots) and from Merkaz Hacarmel, Mt. Carmel (line); and in 1. spirecrinus
from Nahalin, Judean Hills (stripes).
Description. The shell is cylindric to ventricose, thick, light horny brown,
glossy and not granulated. The mouth reaches more than half the shell's height.
The peristome is more than 1.8 mm thick.
Measurements (mm):
Shell-height
Max. diam.
Min. diam.
Mouth-height
Mou th-diam
Lip-width
Shell-height/max. diam.
Shell-height/max.-height
Holotype
(No. 4139)
24.0
12.2
9.3
12.9
9.3
2.4
2.01
1.93
40 paratypes
Range
23.8 -28.7
12.0 -14.4
9.2 -11.7
12.7 -15.6
9.3 -11.0
1.8 - 2.5
1.87- 2.32
1.78- 2.01
Mean
26.03 f 1.32
13.01 k 0.65
10.07 ? 0.48
13.78 f 0.90
9.85 f 0.50
2.00 k 0.16
2.04 f 0.07
1.90 k 0.06
The reproductive system (two specimens from Jiftlik were inspected, of
which one is shown in Fig. 11A), is similar to that of 1. lubrosus. A
well-developed penial flagellum is present. Holotype: Buliminus No. 41 39
(and 40 paratypes, No. 641-654 and 2271-2305) in moll.-coll., Hebrew
University of Jerusalem, Zoology Dept.
Type-locality (Fig. 1 3 ) . Samaria, Nablus-Damiya-BetShe'an crossroad,
1%km E. of Jiftlik;-200, 25 km S.E. Nablus (35" 44' 23" E, 32" 12' 44" N ,
or 197171 Israel grid).
Jiftlik, the name of which the subspecies bears, receives only 200mm
rainfall. The substratum consists of large, well-spaced boulders and projecting
TAXONOMY O F BULIMINUS IN ISRAEL
160
w
15
longitude
170
*E
Figure 9 . Changes from north-west to south-east in the shell-height ( 0 ) and in the ratio
shell-height/mouth-height (a) of I . Iabrosus from the Tulkarm-Latrun road, Samaria.
rocks, with brown xeric rendzina soil. To the west a narrow strip of chalkstone
isolates these boulders from the rocky outcrops and terra-rossa soil inhabited
by 2. labrosus. Phytogeographically Jiftlik is within the Irano-Turanian Batha
steppe (Zohary, 1955).
From Nablus eastwards, as populations of 1. Zabrosus extend beyond the
limits of the 500 mm isohyet, the shells gradually become shorter and the
peristome, in relation to mouth height, gradually thickens. B. 1. jiftfiki
represents, in this respect, the climax of clines for shell-height and lip thickness
10
10
4Q
t.0
K "3
Figure 10. Changes in the ratios shell-heightlmax. diameter ( m ) and shell-heightlmouth-height
of L labrosus from north to south on the Tulkarm-Latrun road, Samaria. and o represent
I spirectinus from the Judean Hills.
(0)
16
J . HELLER
Figure 11. Reproductive system of labrosus. A, 1. jiftliki, Jiftlik; B, 1. labrosus, Sabastia
(north-west of Nablus), northern Samaria; C, 1. labrosus, Um Safa, southern Samaria; D, 2.
spirectinus, Nahalin. Judean Hills; E, F, 1. spirectinus. Har Ora, Judean Hills. The reproductive
system of F resembles that of 1. labrosus but its shell is of a typical 1. spirectinus. This specimen
is probably a hybrid.
TAXONOMY OF BULIMINUS IN ISRAEL
17
0
0
0
0
2b
I
I
I80
West
0
0
185
I90
195
long~tud.
I1
L
East
Figure 12. B. labrosus, east Samaria. Changes in shell-height ( 0 ) and in ratio mouth-height/
lip-width (0)from west eastwards. Small symbols, 1. labrosus; large symbols, 1. jiftliki.
which originate in central Samaria (Fig. 12). The lack of granulation however is
not clinal, as samples of 1. labrosus neighbouring 1. jiftliki are distinctly
granulated.
B. labrosus spirectinus Bourguignat
(Plate 1G)
Bulimus spirectinus Bourguignat, 1876: 1.
The shell is elongate to cylindrical-conic. The aperture is small in comparison
t o shell-height, the ratio shell-height/mouth-height usually being more than
1.95 (see Figs 1 3 and 14).
Measurements ( m m l : Shell-height
24.5 -34.0
Max. diam .
11.6 -15.7
Min. diam.
9.0 -12.2
Mouth-height
12.0 -15.9
Mouth-diam.
8.3 -11.3
Shell-height/Max. diam.
1.89- 2.47
Shell-height/Mouth-height
1.88- 2.25
The reproductive system was inspected in specimens from Nahalin
.(Fig. l l D ) , Martyrs Forest, Zur Hadassa Har Ora (Fig. 11E) Jerusalem and
Sha’ar Hagai. Compared with that of the neighbouring 1. labrosus from
Southern Samaria, the genital system of Lspirectinus has a very short penis,
appendix and pediculus, and no penial flagellum (compare Fig. 11C with
Fig. 11D-E, and see also Table 2).
J. HELLER
18
Table 2. Measurements of the reproductive system of 1. labrosus
from Umm Safa (southern Samaria) and of 1. spirectinus
from Nahalin (Judean Hills)
Pediculus
Epiphallian flagellum
Penis
Penial flagellum
Appendix
Ductus bursae
1. labrosus, Umm Safa
1. spirectinus, Nahalin
23
14
4.5
1.5
5.0
Three times as long as bursa
8
6
1.5
0
2.5
Twice as long as bursa
The type locality described by Bourguignat (1876) is "Dans les gorges des
montagnes, i l'Orient du lac Bahr el Houle". I searched the wadis east of Hula
basin very carefully on several occasions, but no Buliminus whatsoever was
found. I found 1. spirectinus in the Judean Hills (Fig. 4),from Si'ir up to Sha'ar
Hagai, Qalandia and Shillo. This area receives up to 780 mm rain. The
distribution limit in the south, east and west coincides with the 500mm
isohyet. Northwards, in southern Samaria, 1. spirectinus meets the southernmost populations of 1. Zubrosus. I could find no edaphic, climatic or
phytogeographic significance in this borderline. Along it the differences
between the two subspecies are greatest since it is here, where confronting
+
++
c+
...
.
+
.
+
+
+
f
+
t
t
+
+
+
+
+r
+
&+
+
++
+++
++
++
+
24
I
13
I
I
15
14
* O U T *
I
16
1
I7
I
18
I
1s M Y
"ClC*T
Figure 13. Relation between shell-height and mouth-height in 1. labrosus, from Samaria (+) and
1. spirectinus, from Judean Hills
(0).
TAXONOMY OF BULIMINUS IN ISRAEL
19
narrow, small-mouthed 1. spirectinus, that 1. labrosus has populations containing its most ventricose, big-mouthed shells (Fig. 10). Only further north
does it have cylindric and narrow shells (these are responsible for most of the
overlap in shell measurements seen in Figs 1 3 and 14). On Mount Carmel all
shell forms intermediate between the two subspecies are found.
Though the labrosus-spirectinus borderline is usually well-defined, there is
evidence that a certain extent of interbreeding takes place. Some spirectinus
samples (Bet Zait, 'Ein Hemed, 'Ein Kerem, Kiryat 'Anavim) have a few shells
of labrosus and a few intermediates. A specimen from Har Ora, which has a
typical spirectinus shell, has in its reproductive system a minute penial
flagellum, a 4.5 mm long penis and a 9 mm long appendix (Fig. I l F ) , as in
labrosus.
6C
.O
,.._.
,......
20
,.......
J ' ~ll .
.
.... ..
Figure 14. Frequency of ratio shell-heighthouth-height in 1. Zubrosus from Samaria (......) and 1.
spirectinus from Judean Hills (---).
Bulirninus diminutus Mousson
(Plate 1H and 3A)
Bulimus labrosus var. dirninutus Mousson, 1861: 37. Type locality "environs de
la sainte citt".
Petraeus sikesi Preston, 1907: 94. From near Jericho.
Buliminus (Petraeus) labrosus var. dirninutus: Germain 1921 : 276.
The shell is cylindrical-conic t o low-cylindrical, light brown, glossy and
composed of 6-6s whorls. The aperture varies from roundish t o elongate. The
peristome is thin and reflected outwards.
20
J. HELLER
Measurements (mrn): Shell-height
Max. diam.
Min. diam.
Mouth-height
Mouth-diam.
Shell-height/Max. diam.
Shell-height/Mouth-height
Mouth-height/Mouth-diam.
18.1 -27.5
9.2 -13.8
7.3 -10.1
9.3 -13.6
6.6 - 9.6
1.69- 2.20
1.76- 2.17
1.28- 1.48
The reproductive system was examined in specimens from Marsaba, Khirbet
Za'arur (Fig. 16A) and Wadi Qilt. N o significant differences could be found
between the reproductive system of diminutus and that of 1. labrosus.
Figure 15. Intraspecific distribution of Levantina in Israel. The south-western limit of L. s.
caesareana is very similar to that of 1. Iabrosus, and its south-eastern limit is very similar to that
of diminurns (compare with Fig. 4).
21
Figure 16. Reproductive system of A , diminutus (Marsaba); B, fherinus (Saruhen); C , D,
alepensis (Ein Fashkha); E, lamprostatus (Wadi Mashash); F. lamprostams (Ein Gedi); G,
sinaiensis (Har 'Arif).
J. HELLER
22
B. diminutus resembles 1. labrosus, differing from it mainly by its small size
and absence of granulation. In East Samaria, where the two species come
within about 20 km of each other, diminutus differs in that its minimal
diameter, mouth-height and mouth-diameter are smaller and its peristome is
much thinner (Figs 17 and 18). Outside Israel, the two species come close
together in Jordan. Our data from here are scanty but two 1. labrosus samples
.......
9
10
I1
12
.....
I3
........
14
IS
16
l7mm
Figure 17. Frequency of mouth-diameter (A) and mouth-height (B) in 1. labrosus from east
Sarnaria (......), as compared dith that of diminurus (-).
in the Hebrew University collection, from Jerash and Kitte, have a thin
peristome, as in diminutus. Conchologically, they are closer to diminutus than
any other labrosus sample, differing from it mainly in their finely granulated
shell. More material from Jordan might bring more evidence as to the
taxonomic relationship between 1. labrosus and diminutus. B. diminutus may
well be only a subspecies of labrosus.
Synonyms
Preston (1907) describes sikesi from the neighbourhood of Jericho as having
a cylindric shell with a relatively small aperture. His description well fits the
TAXONOMY OF BULIMINUS IN ISRAEL
23
Fr
30
1:
21
A
1
1
......I
07
.... ..
.
I
I
I
0.0
0.9
1.0
II
1.2
13 mrn
Fr
20~
B
16-
I?.
8.
4-
Figure 18. Frequency of peristome width ( A ) and minimal diameter (B) in I. lubrosus from east
Samaria, as compared with that of dirninurus. Legend as in 15.
samples of dirninutus from this area. Plate 3A shows the gradual, continuous
transition from dirninutus t o sikesi, because of which the latter is considered a
synonym. (In the holotype, measurements of the aperture are 10.1 x 7.4, not
9 x 4.7 as in Preston’s original description.)
Distribution (Fig. 4)
The species is confined t o the Judean Desert and, outside Israel, t o the slopes
east of the Jordan River. Germain (1921) mentions it from Jaffa but this is
most probably an error, as diminutus has not been found by any other
collector in that well-searched locality.
The Judean Desert receives 100-400 mm rain annually, most of it falling in
the high, western parts. Where diminzitus is found the substratum consists of
J. HELLER
24
soft chalkstone which is unfavourable for diminutus because it lacks sheltering
crevices. The snails were found amongst boulders of flint which are scattered
amongst the chalk, in the crevices of small cliffs of hardened chalk, or in
canyons that cut through the chalk to the underlying limestone. Vegetation
consists of semi-steppe Batha communities (Zohary, 1955).
SI
1.1.0
aa
A
e
0
100
A
19<
A
I.¶<
A
100
170
210
I90
longitude
7
205
/
A
/tY.
B
A
A
A
I80
West
I90
0
210
I O n g 1 1u d c
East
Figure 19. B. diminutus. Clinal changes in the ratios shell-heighthaxima1 diameter (A) and
shell-heighthouth-height (B) from west eastwards.
TAXONOMY O F BUIJMINUS IN ISRAEL
25
Intraspecific variability
The shell proportions alter clinally. In the western, rainy parts of the Judean
Desert the shells are ventricose and have a large aperture. Eastwards, towards
the arid surroundings of the Jordan River, they rapidly become slimmer and
their aperture is reduced in size (Fig. 19). The Jordan River is the climax of this
cline. Beyond it, in the high rainier areas of Gilead, the shells are once more
ventricose and have wide apertures.
Buliminus therinus Bourguignat
(Plates 11, J and 3B, C)
Bulimus therinus Bourguignat, 1876: 5. Type locality “Vallee du Liban, Syrie”.
Bulimus halepensis: Mousson, 1861: 6. From Marsaba.
Buliminus (Petraeus) carneus var. reconditus Germain, 1921. From the environs
of Jerusalem.
Petraeus therinus: Pallary, 1929: 1 5 ; pl. 1, fig. 22.
The shell is cylindrical-conic to low-cylindrical, thick, dark to light horny
brown and glossless. It is composed of 5%-6?4 whorls which are striated and
have a very strong granulation, best observed under x80 magnification
(Plate 1J). The penultimate whorl is of subequal diameter to the prepenultimate one. The height of the aperture is less than half the shell’s height.
The peristome is reflected outwards, sometimes backwards.
Measurements (mm): Shell-height
Max. diam.
Min. diam.
Mouth-height
Mouth-diam.
Shell-height/Max. diam.
Shell-height/Mou th-height
Mouth-heigh t/Mouth-diam.
13.7 -21.0
5.9 - 9.5
5.3 - 7.3
5.8 - 9.2
4.3 - 7.0
1.88- 2.68
2.00- 2.53
1.23- 1.63
The reproductive system was examined in specimens from Saruhen
(Fig. 16B), Wadi Qilt, Jericho and Dimona. The ductus bursae is thicker, in
relation to the bursa, than in other species.
The coarse granulation of the shell (Plate 1J) enables therinus t o be easily
distinguished from other species.
Synonyms
Mousson (1861) mentions halepensis from Marsaba. The description,
measurements and locality fit therinus very closely. Today, no specimens from
this locality labelled “halepensis” can be found in Mousson’s collection (Dr H.
Jungen, pers. comm.). His description most probably refers to a shell from
Marsaba labelled B. uriae. I have seen this specimen, which is a typical,
granulated therinus. Bourguignat’s description of thexinus appeared only 15
years after Mousson’s article.
G e r m i n (1921) describes carneus var. reconditus from the vicinity of
Jerusalem. The glossless shell, the small size, the proportions and the locality all
J. HELLER
26
suggest that this variety is a slightly more ventricose specimen of therinus (see
Plate 3B). I t has no resemblance whatsoever to carneus.
Distribution (Fig. 2 0 )
In Israel the species inhabits the Judean Desert and the northern and central
Negev. The distribution area lies between the 100 mm and 250 mm isohyets
and coincides with the Irano-Turanian phytogeographic region in Israel. B.
therinus occupies small piles of flint, limestone rock or chalky limestone. I t was
not found in alluvium or sand.
1
/
I
Figure 20. Distribution of therinus in Israel. Most localities are between the 100 rnrn and
250 rnrn isohyets.
TAXONOMY OF BCJLZMINUS IN ISRAEL
27
Data on the distribution outside Israel are scanty. The type locality is in
Lebanon. Specimens from Suleimanik (Iraq) and the Diarbekir-Maltia road
(Turkey) are in the Geneva Museum.
IIZtraspecific variability
Shell measurements and ratios alter clinally from south-west to north-east. In
the western Negev the shells are large, very ventricose (shell-height/max.
diam. = 1.98) and have large apertures. Towards the Judean Desert mouthdiameter decreases by 25% (Fig. 21), mouth-height and maximal diameter by
18% (shell-height/max. diam. = 2.33), shell-height by 12% and the aperture
changes from roundish to elongate. Plate 3C illustrates these clines.
A 4.90
A600
,b.17
Figure 21. B. rhen‘nus. Changes in mouth-diameter from north-east to south-west.
Buliminus alepensis Pfei ffer
Helix (Cochlogena) Alepi De Firussac, 1821: 55. Type locality “Alep, cote de
Syrie, au dit la Coupe, P une demie lieue de ville; Corn m. Olivier”
Buliminus alepensis Pfeiffer, 1841: 45.
Bulirninus (Petraeus) halepensis: Germain, 1921: 278-279.
Petraeus halepensis; Pallary , 1929: 15.
Petraeus halepensis: Forcart, 1940: 165.
N
W
Table 3. Measurements and ratios of a. alepensis (from Jordan and Lebanon) and a. rnarsabensis (from Israel)
N
Shell-height
Max. diarn.
fin. diam.
Mouth-height
Mouth-diarn.
SH/Max. D.
SHIMH
MHIMD
40
16.3-21.1
18.42
1.32
7.2-9.5
8.37
0.60
6.0-7.6
6.91
0.43
7.0-8.6
7.89
0.5 3
5.0-6.5
5.61
0.38
2.01-2.58
2.208
0.11
2.18-2.51
2.331
0.08
1.27-1.47
1.403
0.04
'Ein Fashkha (3 krn S of) 15
M.
S.D.
16.7-20.0
18.79
0.87
8.0-9.1
8.39
0.32
6.5-7.4
6.95
0.27
7 23-8.7
8.22
0.30
5.6-6.2
5.95
0.23
2.1 1-2.41
2.267
0.07
2.09-2 .SO
2.284
0.10
1.33-1.42
1.383
0.02
'Ein et-Turaba
5
17.6-20.0
18.72
1.04
8.4-9.0
8.83
0.29
6.9-7.2
7.08
0.12
7.5-9.0
8.28
0.57
5.7-6.5
6.08
0.30
2.02-2.18
2.088
0.06
2.22-2.3 5
2.264
0.05
1.32-1.39
1.362
0.02
18
17.2-21.0
18.87
1.10
8.0-9.1
8.63
0.38
6.5-7.6
7.04
0.27
7,5-9.0
8.22
0.47
5.6-6.6
5.98
0.89
1.99-2.38
2.190
0.10
2.17-2.49
2.297
0.08
1.33-1.43
1.377
0.03
4
14.9-1 6.9
15.68
0.86
8.0-8.9
8.33
0.40
-
7.5-8.5
7.83
0.47
5.8-6.3
6.03
0.25
1.80-1.93
1.885
0.005
1.99-2.05
2.005
0.003
1.29-1.35
1.313
0.03
Lebanon: Founoul
M.
S.D.
10
19.7-22.0
20.75
0.66
9.0-9.9
9.56
0.1 3
7.3-8.2
7.81
0.1 5
8.3-9.6
8.93
0.13
6.2-7.1
6.63
0.08
2.05-2.32
2.171
0.06
2.17-2.43
2.326
0.04
1.21-1.25
1.343
0.18
Lebanon: Tourboul
21
17.5-22.1
19.85
1.09
8.8-10.1
9.46
0.36
7.4-8.8
7.77
0.38
8.2-9.2
8.70
0.36
6.2-7.2
6.66
0.29
1.96-2.22
2.092
0.08
2.17-2.43
2.278
0.06
1.23-1.37
1.306
0.04
Locality
a marsabensis
'Ein Fashkha
M.
S. D.
M.
S.D.
'Ein el Ghuweir
M.
S.D.
a alepensis
Jordan: Shuni
M.
S.M.
M.
S.D.
TAXONOMY OF BULIMINUS IN ISRAEL
29
B. alepensis marsaberisis Westerlund
(Plates 1 K and 3E-G)
Bztliminus marsabensis Westerlund, 1887(3): 57. Type locality “Marsaba.”
The shell is cylindrical-conic to low-cylindrical, dark to light horny brown
and glossy. I t is composed of 6-7 whorls which are delicately striated. The first
3-4 whorls increase rapidly in height and diameter, and form the apex of the
shell. The last 2-3 whorls hardly increase in diameter, and the penultimate
whorl is usually not wider than the prepenultimate one. The aperture is
elongate, oval and less than half the shell’s height. The peristome is less than
one millimeter thick, and reflected slightly outwards.
Measurements (mm): Shell-height
Max. diam.
Min. diam.
Mouth-height
Mouth-diam.
Shell-height/Max. diam.
Shell-heigh t/Mouth-height
Mouth-height/Mouth-diam.
15.2 -21.1
7.1 - 9.7
6.1 - 7.7
6.6 - 9.0
4.9 - 6.6
1.95- 2.41
2.09- 2.50
1.27- 1.47
The reproductive system was examined in five specimens from ’Ein Fashkha,
of which two are shown (in Fig. 16C-D). The bursa is well-defined and has a
thin duct (contrast to therinus). The penial flagellum, when present, is
roundish. In some specimens it is absent (Fig. 16D). The two retractors may
unite into a common muscle or, in other specimens (Fig. 16D) may insert
separately at the diaphragma.
B. a. marsabensis differs from a. alepensis from Lebanon in that its
maximum and minimum diameter are narrower in relation to mouth diameter
(Table 3). Compared with a. alepensis from Jordan the Israeli shells are higher
and their aperture is narrower (Table 3).
Plate 3E presents two marsabensis shells from ’Ein Fashkha which are very
similar to damascensis var. minor Pallary (compare with Pallary, 1929: ~ 1 . 1 ,
fig. 28). B. a. marsabensis and demascensis are probably closely related or even
synonyms. More material from Syria and Jordan would enable a better
Table 4. Measurements of the reproductive system of a. marsabensis from
’Ein Fashkha (five specimens), as compared with those of a. alepensis
from north Syria (from Hesse 1933: 198)
a. alepensis
a. marsabensis, ’Ein Fashkha
Ackbes
(from Hesse 1 9 3 3 )
Pediculus
Ductus bursae
Diverticle
Penis
Epiphallus
Epiphallian flagellum
Appendix
Appendicular flag
7
3.2
7
3.7
5
7
5.5
19.0
5.5
2
5
3
2.5
2.5
4
13.5
4
2.5
6
2
2.5
2.5
3.5
10.0
4.5
3.5
7
2
2
2.5
7
14.0
5
2.5
5.5
3.5
2.5
4
4
12.0
4
2.5
5
3
3.5
2.5
3
11.5
30
J. HELLER
understanding of the relationship between the two. Forcart (1940) states that
in alepensis the columella terminates in the upper half of the columellar lip. In
the Israeli populations the columella appears to terminate in the upper, middle
or lower half, but this is a difficult character. Plate 3F-G present the extent of
variability in shell-height and maximum diameter, within one sample.
Hesse (1933) examined the reproductive system of three alepensis specimens
from Akbes, North Syria. Table 4 presents the measurements of the Israeli
specimens in comparison to those of Hesse. The Israeli specimens differ in that:
1, the pediculus is shorter, both absolutely and as compared with the diverticle;
2, the epiphallus usually reaches only half the length of the North-Syrian ones;
3, the epiphallian flagellum is less than half as long as in North Syria.
Distribution (Figs 22 and 23)
B. a. marsabensis inhabits the cliffs and rocky slopes of the northwestern
shores of the Dead Sea, from ’Ein Turaba to Qumran. Two samples from Wadi
Qilt (leg. G. Haas) extend the distribution a little northwards. The area receives
only 90 mm rain (for comparison, areas in Lebanon, Iraq or Turkey, where a.
alepensis is found, receive over 900 mm). The mean temperature is 31” C in
August and 15”C in January, and the daily relative humidity ranges from 36%
(July) to 46%(January). The area immediately west of the cliffs is occupied by
therinus. Southwards, near ’Ein Turaba, a. marsabensis meets the northernmost
populations of lamprostatus. Here, a single sample shows peristome width
which is intermediate between the two species (Plate 3D), indicating that some
interbreeding may occur.
Figure 23 presents the distribution of alepensis outside Israel, as recorded in
literature (Pfeiffer, 1841; Boettger, 1898; Germain, 1921; Biggs, 1937, 1959;
Pallary, 1939; Forcart, 1940; Haas, 1952; Gittenberger, 1967) and in the
Hebrew University’s collection. The species is found in a broad arc from Iran
via Kurdistan, South-East Turkey, Syria and Lebanon to Jordan and Israel.
Buliminus lamprostatus Bourguignat
(Plates 1L and 3H)
Bulimus lamprostatus Bourguignat, 1876: 3. Type locality “Les vallkes du
Liban, sur les rochers”.
Bulimus carneus: Tristram, 1865: 537. From Mezada.
Buliminus labrosus var. asphaltinus: Westerlund, 1887( 3): 57; from Dead Sea.
Petraeus fourtaui Pallary, 1923: 210-212; pl. 12, fig. 4, 7. From Gebel Hellal.
Petraeus lamprostatus: Pallary, 1929: 1 5 ;pl. 1, fig. 2 1 .
The shell is cylindrical-conic t o low cylindrical, very thick, light horny
brown and very glossy. It is composed of 6-634 weakly striated whorls. The
penultimate whorl is usually wider than the preceding ones. The aperture is
elongate and less than half the shell’s height. The outer and inner lips are
connected by a very strong callus. The peristome is more than one millimeter
thick and reflected slightly backwards.
TAXONOMY OF BULIMINUS IN ISRAEL
a
Hebron
Figure 22. Distribution alepensis marsabensis (o), lamprostatus (A), glabratus (m) and negevensis
in Israel. The upper limit of glabratus closely coincides with the 1 5 0 mm isohyet (stippled
area represents the sand and alluvium areas of Mishor Rotem and Nahal Ef'e).
(0)
31
32
J . HELLER
Figure 23. General distribution of alepensis.
Measurements fmml: Shell-height
Max. diam.
Min. diam.
Mouth-height
Mouth-diam.
Shell-height/Max. diam.
Shell-height/Mouth-height
Mouth-heigh t/Mouth-diam.
16.5 -24.0
8.0 -11.0
7.1 - 9.1
8.0 -10.5
6.0 - 8.3
1.83- 2.36
1.94- 2.37
1.25- 1 . 4 4
The reproductive system was examined in ten specimens, from ’Ein Gedi
(Fig. 16F), Nahal Zeelim, Wadi Mashash (Fig. 16E) and ’Ein Boqeq. Usually the
genitalia are larger than those of a. marsabensis. and differ from it in that the
penial flagellum is always represented by a small protuberance. Those from
Wadi Mashash (Fig. 16E) which is very close to the meeting-point with a.
marsubensis, have a smaller reproductive system and may lack the penial
flagellum as in marsubensis. This is further evidence that these two species may
interbreed where they meet (see above, and Plate 3D).
Live snails of lamprostatus differ from all other Israeli species in that their
skin is black, not yellowish. Shells of lamprostutus differ from alepensis
marsabensis in their thicker peristome (Fig. 24) and larger aperture in relation
to shell-height (Fig. 25). They differ from diminutus in their smaller aperture
and from lubrosus jiftliki by their smaller shell-height.
Synonyms
Tristram (1865) mentions curneus from Mezada. His description, in
particular concerning “the solidity of the peristome and the callosity largely
TAXONOMY O F BULIMINUS IN ISRAEL
33
F,
14
............
. .
I1
.
.:
...
.....
..
10
8
6
...
....
....
...
.
_......
...
.
.....
...
....
..
...
... .,..,.
. .
.. ...
i......
4
2
.
.:
....
.....
.
........
r
Figure 24. Frequency of peristome width in alepensis marsabensis (......) and [amprostatus
(---).
extended over the last whorl at once distinguish every specimen” (p. 5 37) does
not fit,carneus with which he was probably unacquainted (“I have not been
able to compare my specimens with the type”). It does, however, very well suit
lamprostatus, which is the only species found today near Mezada. Bourguignat
described lamprostatus 11 years after Tristram’s article.
The holotype of B. labrosus var asphaltinus Westerlund is lost (Dr H. Waldtn,
pers. comm). The lectotype, which Dr Waldkn kindly figured for me, proved
upon examination to be a typical lamprostatus.
Pallary (1923) describes B. fourtaui from Sinai, His description, especially
concerning “Ouverture ovalaire, a bord externe et inferieur bien cambrts”
(p. 210) fits lamprostatus well. Plate 3H presents four shells of lamprostatus
from ’Ein Gedi which are very similar to the types of fourtaui as figured by
Pallary (1923: pl. 12, figs 4-7). I therefore consider the latter a synonym.
Distribution (Figs 2 2 and 26)
B. lamprostatus is abundant mainly on the cliffs bordering the south-western
shores of the Dead Sea. This area is extremely arid (50-85 mm rain) and hot:
mean temperature of the hottest month (August) is 33.8”C, of the coldest
(January) 15.8”C, and daily relative humidity ranges from 33% (July) to 56%
(January). The substratum consists of hard limestone. The Lisan marl areas
below the cliffs are uninhabitable and the stony areas above it are occupied by
another species, glabratus. B. lamprostatus is confined mainly to the barren,
Saharic habitats of its area, and is absent from the damp areas of lush
vegetation surrounding the few springs (such as ’Ein Boqeq or ’Ein Gedi) that
run down the cliffs. Further north lamprostatus is replaced in these habitats by
a. marsabensis. N o climatic, edaphic or phytogeographic significance could be
attributed to the borderline between the two.
The species is also found in a few localities in north Sinai (Fig. 26) in areas
which receive only 20-60 mm rain. The type locality in Lebanon could be an
3
34
J. HELLER
( 0 )
24
I
t
e
2:
'
r
22
-
21
*
2c
-EE
s
0
+
.-
x
s
I
19
0)
d
18
17
..
-
-
...
. ;.. :"
..
.'
.
.
. .
4
* +
... . .
*
t.+
.
a
i
t
..
16
15
7
8
9
Mwth-height (mm)
10
II
error, as there are no further records from there. There are no habitats in
Lebanon which could be considered Saharic (Atlas of Israel, 1956).
In trasp ecific variability
The shells of Sinai differ from those of the Dead Sea area in that they are
broader (shell-height/max. diam. = 1.97, as compared with 2.12 in the 'En Gedi
sample), and the surface of the shell is slightly less glossy and more striated.
Along the Dead Sea itself no constant pattern of variation could be found.
Buliminus sinaiensis Heller
(Plate 1M)
Buliminus (Buliminus) sinaiensis Heller, 1970: 279-280, figs 1, 4, 5. Type
locality "Negev, Har 'Arif".
The shell is cylindric, light horny brown glossy, semi-transparent and is
composed of 5%-6 whorls. The apex consists of *3 slightly convex whorls, the
other whorls are nearly flat. The suture is very shallow, except on the last
whorl, where it is slightly deeper. The aperture is roundish and less than half
35
TAXONOMY O F BULIMINUS IN ISRAEL
24
23
22
-E
-i
21
20
P
c
-
6a
19
18
"
17
..
. .
..
16
15
5
6
7
8
9
Mouth - diam.(mm)
Figure 25. Relation of shell-height to mouth-height ( A ) and to mouth-diameter ( B ) in alepensis
marsabensis ( 0 ) and lamprostatus (+).
the shell's height. The peristome is less than one millimeter thick and reflexed
slightly backwards.
Measurements (mm): Shell-h eight
Max. diam.
Min. diam.
Mouth-heigh t
Mouth-diam.
Shell-height/Max. diam.
Shell-heigh t/Mou th-height
Mouth-height/Mouth-diam.
17.6 -21.6
8.5 -10.0
6.7 - 7.9
7.6 - 9.5
5.9 - 7.4
1.84- 2.19
2.09- 2.37
1.17- 1.36
The reproductive system of sinaiensis was described previously (Heller,
1970), and no live specimens have been found since. The genitalia (Fig. 16G)
are similar to those of a. marsabensis from Ein Fashkha. B. sinaiensis differs
from alepensis marsubensis in that its mouth-diameter is much bigger in relation
to mouth-height (Fig. 27). Its thin peristome and smaller shell-height distinguish it from lamprostatus. In Makhtesh Ramon, shells of sinaiensis were
found together with those of therinus in front of a rodent's burrow. B.
J . HELLER
36
50 km
I1
Figure 26. Distribution of lamprostatus
(A) and
sinaiensis
(0)
in Sinai.
sinaiensis could here be easily distinguished from the latter in its more
cylindrical shell, roundish mouth and lack of granulation.
Distribution(Fig. 26)
In the Negev the species was found at Har 'Arif, Makhtesh Ramon, Har
'Oded and Nahal Timna'. In Sinai it was found at Gebel Tarbush and Gebel
Sirbal. If quantity of shells found per unit time expresses population size, then
the biggest population is at Har 'Arif, which towers 200-350 m above its
TAXONOMY O F BULIMINUS IN ISRAEL
37
A
I
50
I
I
54
I
1
58
1
I
1
62
I
66
I
I
7.0
I
I
7.4
Mouth diom (mm)
Figure 27. Relation of rnouth-height to mouth-diameter in sinaiensis (a) and alepensis
rnarsabensis (a).
immediate surroundings and receives up to 70 mm rain. Har 'Arif is within the
Saharic phytogeographic region, just beyond the most south-western point of
the Irano-Turanian region (which is populated by therinus). The shells were
found on a stony slope beneath a cliff, on the northern side. The Saharic areas
south of Har 'Arif are unpopulated because of their extremely hot, dry climate.
In southern Sinai the shells were found on magmatic rocks in an Irano-Turanian
enclave which is completely surrounded by Saharic regions. Mean annual
rainfall in the Irano-Turanian enclave at St. Catharine, near Gebel Musa, is
61.8 mm (Aloni, in Haim, 1969).
Intraspecific variability
I t is remarkable that the samples of Har 'Arif and of south Sinai are so
similar (Table 5 ) , even though they are 275 km apart, and though Har 'Arif
consists of limestone whereas southern Sinai consists of granite.
Buliminus glabratus Mousson
(Plate 1N-0)
Buliminus carneus var. glabratus Mousson, 1861: 37, Type locality
"Es-Zenore".
NOT Petraeus cameus glabratus: Forcart, 1940: 169-170; pl. 2, fig. 32.
The shell is cylindrical-conic to tower-like, light horny brown and glossy. I t
is composed of 7 - 9 s finely striated whorls. The first 3-4 whorls are convex and
increase rapidly in height and diameter, forming the apex of the shell. The
remaining 4-5 whorls are flatter and hardly increase in size. The aperture is less
w
m
Table 5 . Measurements and ratios of sinaiensis
~
~~
Locality
N
Shell-height
Max. diam.
Min. diam.
Mouth-height
Mouth-diam.
SH/Max. D.
SHIMH
MHIMD
Har 'Arif
M.
S.D.
44
17.7-20.5
19.43
1.36
8.5-10.0
9.39
0.43
6.3-7.9
7.52
0.33
7.7-9.5
8.74
0.41
6.2-7.4
6.84
0.32
1.84-2.19
2.038
0.08
2.09-2.3 7
2.192
0.06
1.17-1.36
1.276
0.04
Makhtesh Ramon
M.
S.D.
6
18.3-21.2
19.93
1.oo
8.8-9.6
9.38
0.33
7.3-7.8
7.52
0.17
8.3-9.0
8.60
0.32
6.3-7.3
6.78
0.35
2.04-2.31
2.12s
0.10
2.18-2.49
2.320
0.1 3
1.22-1.35
1.270
0.04
Sinai: G . Tarbush
13
17.7-19.3
7.7-9.3
6.7-7.7
7.2-8.5
5.6-6.8
1.98-2.30
2.23-2.54
1.19-1.29
18.28
0.70
8.69
0.48
7.20
0.30
7.78
0.42
6.24
0.30
2.117
0.69
2.361
0.10
1.249
0.03
and G . Sirbal
M.
S. D.
TAXONOMY OF BULIMINUS IN ISRAEL
39
than half’ the shell’s height. The outer lip usually protrudes slightly from the
penultimate whorl like a small shelf. Outer and inner lips are connected by a
strong callus which is usually more pronounced than in other Israeli species.
The peristome is thin and reflected slightly outwards. On the inside of the
outer wall of the body whorl there is usually (see below) a palatal lamella, seen
from the outside as a white stripe which begins near the insertion of the outer
lip onto the penultimate whorl, curves along the body whorl and ends at the
middle or lower half of the outer lip.
Measurements ( min): Shell-heigh t
Max. diam.
Min. diam.
Mouth-height
Mouth-diam.
Shell-height/Max. diam.
Shell-height/Mouth-height
Mouth-height/Mouth diam.
12.0 -20.5
4 . 8 - 7.1
4.2 - 5.7
4.4 - 6.2
3.2 - 4.5
2.26- 3.16
2.51- 3.54
1.24- 1.54
The reproductive system was examined in specimens from Wadi Mashash
(Fig. 28A), Hamakhtesh Haqatan and Dimona. The entire female system, from
albumin gland to vagina, is ensheathed in black connective tissue, unlike any
other species in Israel.
Figure 28. Reproductive system of (A) glabrants (Wadi Mashash); (B) negevensis (Har Avnon).
B. glubrutus differs considerably from other species, in its narrow shell and
small aperture (note the high values of shell-height/max. diam. and shellheight/mouth-height) and in the palatal lamella, which was found in all samples
though not in every specimen. I t is probably not as closely related to the other
Israeli species as the latter are related amongst themselves.
Synonyms
Mousson (1861) provisionally placed “var” glubrutus within the species
carneus Pfeiffer. Forcart (1940), accepting this classification, considers
glubrutus as a senior synonym of curneus minor. He therefore describes
“curneusglubrutus” from South Anatolia. I consider this classification incorrect
J. HELLER
40
for two reasons: (a) The shell of glubrutus has a very marked callus (Plate
1 N - 0 ) . Mousson (1861: 38) stresses that “La partie la plus particulikre est
toutefois l’ouverture, dont le bord n’est form6 que d’un arrondi blanc, qui
continue en courbe rtguliPre de la columelle sur la callositt de l’avantdernier
tour jusqu’i l’insertion droite; tout le contour se trouve en un meme plan, ce
qui donne ii l’ouverture quelque chose de dtlicat et d’tltgant.” This description
fits both the syntype and the 200 specimens I collected. In contrast, curneus
has no callus whatsoever (Forcart, 1940: 168 “Mundungswand ohne Callus”
and his fig. 32-33; also specimens in Hebrew University’s collection). (b)
Mouth-diameter of glubrutus is much narrower than in carneus.
Distribution (Fig. 22)
The species is found from Wadi Mashash in the Judean Desert t o Har
Gerafon in central Negev. It encircles the sands and alluvial soils of Mishor
Yamin, Rotem and Nahal E f e westwards, towards Dimona, and eastwards
towards Hamakhtesh Haqatan. The type locality, Es-Zenore, is today identified
with Nahal Zohar (Forcart, pers. comm.). The distribution area of the species
in Israel receives 70-150 mm rain (the 150 mm isohyet actually defines the
species distribution border to the west) and has a Saharic vegetation which
consists largely of the Zygophyllum dumosum associations. The snail inhabits
bare rocky outcrops of limestone, dolomite, flint or hardened chalkstone but
A
r-
B
Figure 29. B. glabrufus. A. Changes in shell-height from north to south and south-west. B.
Changes in shell proportions (on left of triangle-shell-heightlmaximaxmal diameter; on rightshell-heightlmaximouth-height).
TAXONONY OF BULIMINUS IN ISRAEL
41
not sand or alluvium. Mousson’s remark about identical shells from Cyprus is as
yet the only record of glubratus from outside Israel.
In traspecific variability
The shells in the west are smaller and stouter than the slender, elongate
forms of the east and north. From Dimona to Wadi Mashash, over 65 km,
shell-height rises by 38%(Fig. 29), maximal diameter by 22%and mouth-height
by 17%. Correspondingly, the western shells have 7 4 % whorls, the southeastern have 7%-8%and the large northern ones have 8%-9%.N o other Israeli
species has such high variability in the number of whorls.
Though in all samples at least some of the shells have a palatal lamella, in the
southeastern range of the species the distinctness of the palatal lamella within
one sample may vary considerably. I t may be strongly developed, completely
absent or in an intermediate position (Plate 10). In the northern and western
range of the species all shells have a distinct lamella.
Buliminus negevensis Heller
(Plate 1P)
Buliminus (Buliminus) negevensis Heller, 1970: 280-285, figs 2, 3 , 6. Type
locality “Negev, above Mezad Zohar”.
The shell is elongate, light brown, glossy and semitransparent. I t is composed
of 7-9 whorls, which are striated. The first k 4 whorls increase rapidly in height
and diameter, and form the apex of the shell. The last 3-4 whorls increase
slowly in height, but hardly in diameter. All whorls except the penultimate are
convex. The aperture is elongate, less than half the shell’s height. The peristome
is thin and reflexed slightly backwards.
Measurements ( m m l : Shell-height
Max. diam.
Min. diam.
Mouth-heigh t
Mouth-diam.
Shell-height/Max. diam.
Shell-height/Mouth-height
Mouth-height/Mouth-diam.
14.0 -23.2
5.9 - 9.0
5 . 1 - 7.3
5 . 1 - 7.8
3.9 - 5.6
2.21- 2.97
2.43- 3.18
1.24- 1 . 5 0
The reproductive system of negevensis was described in a previous paper
(Heller, 1970), and no live snails have been found since. The genitalia
(Fig. 28B) resemble those of glabratus in their elongate, slim structure, but
differ in that the female system is not ensheathed in black connective tissue.
The shell of negevensis differs from that of glubratus in its complete absence of
a palatal lamella, and in the relation of shell-height to maximal diameter,
minimal diameter and mouth-diameter (Figs 30 and 31). These shell
characteristics put negevensis in an intermediate position between glabratus and
the other Israeli species.
Distribution (Fig. 22)
The species was found at Har Hemar, Har Avnon, HaMakhtesh Hagadol,
Nahal Hatira and Ma’ale Aqrabim. These localities are on the easternmost
42
++ +
+
+
++
A
+
+
+ +
+
+
+
+
t +
+
+
t
t
+ t
+
+
t
+
t
. . . .;
+.
+
+
+ +
+
0
0
0
0.
0.
0
.
0.
0
0
00
.
~...~.
..
.
. . ....!...
. ..
p ++ +
+
0
H
0.
0.m
Do
0:
+,*+
t
+
0
0
0
0
0 0
+
I++ ,+
++t+
0
8-
8.
9
st
.
0
0
:
0..
.
0
0
0
0
m
+
+
+
t
a
++
+
+
0.
0
TAXONOMY OF BULIMINUS IN ISRAEL
43
23
+
21
+
.
0
.
.
.
0
I
"'
0 0 %
+
+
.
:
-
+
I
15
.
13.
.
40
44
*
4.8
5.2
5.6
Min D
6.0
6.4
6.8
7.2
(mm)
Figure 3 1 . Relation of shell-height to minimal diameter in negevensis (+) and glabratus
(0).
slopes of the central Negev, on the verge of the steep descent towards the Rift
Valley and Zin Desert, where extremely arid conditions prevail. The distribution area receives 70-100 mm rain. Northwards and eastwards, where better
conditions prevail, the area is occupied by therinus and glabratus. The
vegetation type in the distribution area is Saharic.
In traspecific variability
Shells in the north-east are 30% taller than those of the south-west (20.04 in
Har Hemar as compared to 15.23 in Har Avnon], mainly because of an addition
of whorls, hardly accompanied by changes in shell-diameter or mouth-height.
This pattern of change is very similar to that of glabratzts, which neighbours
negevensis to the north.
44
J. HELLER
DISCUSSION
Taxonomic remarks
B. diminutus, which in this work is considered a separate species, may well
be only a subspecies of labrosus. On the other hand marsabensis, which is here
regarded as a subspecies of alepensis, may perhaps be a fully separate species. I t
is difficult t o evaluate the precise position of these taxa in the Linnean
hierarchy, mainly because of the very scarce material available from Jordan and
Syria. On evidence of the shell and genitalia it appears that diminutus is very
closely related to labrosus, and that sinaiensis may be related to a. marsabensis.
B. glabratus and negevensis are probably closely related to each other and both
of them stand slightly apart from the other Israeli species. Except for
diminutus which, as shall be discussed elsewhere, has almost certainly evolved
from labrosus, it is practically impossible at our present state of knowledge to
suggest whether or not any of the Israeli species has evolved from any other of
the recent ones.
Int raspecific and int erspecific clines
The shell proportions of diminutus alter clinically (Fig. 19) from a ventricose
and large-mouthed shell in the cool, rainy regions of the Judean Desert (near
the Judean Hills) t o a slim, small-mouthed shell in the hot, arid regions (near
the Jordan River). Similarly, also the shell proportions of both therinus
(Fig. 21, and Plate 3C), glabratus (Fig. 29B) and negevensis alter clinally from
ventricose, large-mouthed shells in the relatively cool Negev, t o slim,
small-mouthed shells in the relatively hotter Judean Desert; and shells of
lamprostatus from Sinai are more ventricose than those from the Dead Sea
region, which is very hot. There is thus a general trend within most Buliminus
species to develop narrow shells with small mouths in the hotter regions of
their distribution area. A small mouth could reduce the loss of water by
evaporation via the aperture, whilst a snail with a narrow shell would be able to
enter narrower rock crevices to avoid the heat of the summer.
B. labrosus and therinus, the species which in Israel inhabit the more cool,
rainy regions within the distribution area of Buliminus, are both granulated
(Plate lB, J). All other Israeli species, which dwell in more arid conditions than
these two, have non-granulated, relatively smooth shells; and lamprostatus,
which inhabits the most hot, arid regions within the range of Bulirninus, has an
exceptionally smooth and glossy shell. Further, where labrosus encounters the
relatively arid conditions of eastern Samaria, its shell loses its granulation. A
somewhat similar situation occurs also in the Israeli species of Sphincterochila, where the Mediterranean-dwelling cariosa is coarsely granulated whereas
the steppe-dwelling boissieri and the arid-dwelling arabica have smooth, glossy
shell surfaces. As a glossy surface reflects radiation better than a granulated one,
smooth, ungranulated shells could, in hot arid conditions, add to the efficiency
of thermal isolation, and thus be of some selective advantage.
In lamprostatus, which inhabits the hottest and dryest habitats within the
range of Buliminus, the shell has an extremely thick lip (Plate 1L; Fig. 24).
Further, in Mediterranean-dwelling labrosus the lip thickens clinally, as the
TAXONOMY O F BCJIJMINUS IN ISRAEL
45
species encounters the relatively arid conditions prevailing in eastern Samaria
(Fig. 12). It is noteworthy that also amongst the Enidae of Cyrenaica,
Cyrenaeus kaltenbachi, which inhabits the most arid habitats of the genus (see
Heller, 197 1 for generic classification), has extremely thick lips; and that
amongst the Enidae of Saudi Arabia, arid-dwelling Mastus labiosus has an
unusually thick lip. A thick lip may perhaps enable a more firm adherence t o
the substratum, or maybe enable a firm, tight attachment of the epiphragma t o
the shell’s aperture. In both cases this would further reduce the loss of water
via the aperture of the shell.
Ecological factors correlated to distribution
Climate
The borderline of Zabrosus spirectinus coincides well with the 500 isohyet
(Fig. 4); therinus is delimited between the isohyets 250 mm and 100 (Fig. 20);
glabratus is found only in areas receiving 70-150 mm rain (Fig. 22); lamprostatus in areas of 70-50 mm. Of special interest is the 70 mm isohyet along
which pockets of three taxa of restricted distribution (sinaiensis, negevensis,
alepensis marsabensis) are found. The close correlation of these distribution
patterns to the isohyet map leaves little doubt that rain is a major factor in
these species distribution.
Within this context the species fall into two categories:
(1) Allopatric species, which do not share borders with any other species of
the genus. The dependence of such a species range on rain, directly or
indirectly, seems unequivocal. This includes Mediterranean labrosus spirectinus:
between it and steppe-dwelling therinus there is a gap of 40 km and of 250 mm
rain (Fig. 4).
(2) Parapatric species pairs, as therinus-glabratus or glabratus-Zamprostatus.
The location of the frontier between these species may well be influenced by
interspecific competition, and the rain may only be influencing the point of
equilibrium between the species. Other Israeli Enidae which show a correlation
between rainfall and distribution are Paramastus episomus Bgt and Pene
sidoniensis Ferussac. Both are the southern-most species of their genera; the
former does not range beyond the 500 mm isohyet (Heller, 1971), the latter
not beyond the 400 mm (Heller, 1974).
Similar correlations were found also in the Israeli species of Jaminia
(Rochanaburanahda, 1968).
Subs tratum
Buliminus being a rock-dwelling genus, the distribution of its species, within
the boundaries of the relevant isohyets, is further limited by lithic requirements. As the nature of the substratum is reflected to a great extent in the type
of soil it produces, snail distribution also shows a considerable correlation with
soil distribution.
The connection between distribution and substratum is most conspicuous in
Zabrosus, because the lithologic map in its distribution area is most complex.
Over 96% of the localities of this species consist of limestones or dolomite
(Fig. 6), the remaining 4% are on kurkar (a hardened, calcareous dune sand).
N o shell was found on chalk, basalt or on alluvial plains,
46
J. HELLER
This unequivocal correlation to substratum probably stems from the fact
that limestone and dolomite have well-developed systems of crevices, which
provide the only available shelter from the severe heat and drought prevailing
throughout the Israeli summer. A snail which fails to reach the shelter of a deep
enough crevice cannot survive. Chalkstone does not have crevices, and the rocks
of hardened chalk (“Nari” rocks) which are found in some regions amongst the
chalk do not have fissures of sufficient depth. Therefore, presumably lubrosus
cannot inhabit chalk. Similarly, i t is absent from plains and valleys devoid of
rock because it is incapable of burying itself deep into the soil to avoid
drought, in the manner that slugs and many Helicidae do. Basalt has suitable
crevices, and absence here is presumably the result of lack of calcium (Dan &
Raz, 1970). Oldham (1929) has found that shells of Ariunta can develop only
very thin shells under experimental conditions of calcium deficiency. Boycott
(1934), Burch (1955), Voelker (1955), Lozek (1962) and Murray (1966) all
consider calcium a major factor in snail distribution.
Because lubrosus is dependent upon hard limestone, its distribution closely
follows that of terra rossa, the soil which in mediterranean climates is derived
from limestone. The question therefore arises whether terra rossa itself might
be of any importance in the snail’s distribution. The following two observations
suggest that this is not the case: (1) B. labrosus is not found in terra rossa
where it contains no rocks. If the snail depended upon the soil, one would
expect to find it also here. (2) At one single locality on the coast, as
mentioned, lubrosus inhabits kurkar, where the soil is not terra rossa, but
Hamra. Kurkar is both calcareous and has fissures in which to aestivate.
The two main steppe-dwelling species, therinus and glubrutus, inhabit a very
wide spectrum of substrata. B. therinus seems capable of inhabiting practically
any small heap of stones within its reach. Though usually frequent amongst
limestone rocks, it was found also upon chalk, flint or any other rocky outcrop
available. It is, from this aspect, the most euryoecous species of the genus. Its
distribution coincides with that of the brown lithosol soil (as defined and
mapped by Dan & Raz, 1970). Surprisingly, the species is not found in the
heaps of stones that are on terra rossa or rendzina soils, even when these are
not occupied by other species. Such is the case in the Judean Hills, where
therinus reaches the last hill bordering to terra rossa, but does not venture
further. As a steppic species, therinus may not be adapted to the draining
regime of mediterranean soils. The efficiency in which water is drained from
crevices and from the undersides of stones could be crucial for the survival of
the eggs. and juveniles of rock-dwelling snails, and apparently labrosus is
adapted to the conditions of terra rossa, whereas therinus is not. A somewhat
similar situation occurs in Britain where Cepaea nemorulis, which buries its
eggs, occurs in light soils whereas Ar-iantu ubustorurn, which lays its eggs on the
surface beneath leaves, occurs mainly in wet soils (Prof. A. J. Cain, pers.
comm.).
B. glubratus is found in the steppe periphery, just beyond the area of brown
lithosol. It too is capable of exploiting a wide range of substrata and inhabiting
very small piles of stones, though to a lesser extent than the former species.
The significance of stones and rocks as a shelter from desiccation is very
pronounced in the desert. All desert species of Buliminus have a disjunct
distribution; sinaiensis is found in central Negev, and separately, in south Sinai;
TAXONOMY 0 1 : BU1,IMINCJS I N ISRAEL
47
lamprostatus inhabits the Dead Sea area, and also central Sinai; alepensis is
found near the Dead Sea, cut off from the major part of the range in Jordan,
Syria, Lebanon, Turkey and Iran; negevensis is known, as yet, only from the
central Negev, where it is split into two isolated areas. Such a distribution
pattern in snails is characteristic of relics. Cliffs and crevices, where they occur
in the desert, provide a refuge to those species of high rainfall habitats which,
whether by eremiation or by interspecific competition, have become extinct in
most of their original areas. They now cling to those peripheral localities where,
owing to substratum which supplies a suitable microclimate, conditions for
survival still prevail. Hard limestone rocks hardly absorb water, and have much
surface run-off after even slight quantities of rain (Danin, 1970). The water
drains into the cracks and crevices, whereby a rock-dwelling snail can survive
amidst the very dry conditions which prevail in the immediate surroundings.
The ecological significance of desert rocks and cliffs as a refuge for relics is
demonstrated by diminutus, which in the higher, rainy region (300-400 mm) of
the Judean Desert populates continuous areas, but in the low, dry region
(100 mm) is limited to the cliffs and canyons.
Vegetation
The distribution of labrosus corresponds to that of the Mediterranean oak
maquis; diminutus is concentrated mainly in areas of the Batha communities;
therinus corresponds t o the Irano-Turanian shrub-steppes of Artemisia herbaalba associations, and glabratus to the associations of Zygophyllum dumosum.
Apparently both snail and plant distribution are influenced by similar factors,
namely rainfall and substratum. The possibility that vegetation is directly
influencing snail distribution, in that different Buliminus species have specific
food preferences for different plants, could not be confirmed by my
observations. All species feed upon decaying vegetable matter and there is no
evidence to suggest that the food requirements of one species differ from that
of another.
However, vegetation may play an important role in microgeographic
distribution, especially in preventing snails from inhabiting certain habitats.
Evidence of this is offered by the fact that labrosus, the mediterranean species
which is seldom found outside the area delimited by the 500 mm isohyet, is
found within this area upon the hot, dry southern slopes, which have extreme
temperatures. It is very uncommon on the cool, damp northern slopes which
are covered with dense oak maquis. The south-facing slope is preferred also by
many other mediterranean landsnails (Levantina caesareana, Rima cariosa,
Helicella bargesiana, Jaminia septemdentata, J. saulcyi, Paramastus episomus).
Such a preference for the south slopes could perhaps be due to the annual
vegetation prevailing on it. Annual plants do not develop xeromorphic
characters and are therefore easily eaten and digested. The oak-maquis of the
north-facing slope may be cool, damp and offer more shelter, but it consists
mainly of perennial vegetation. The development of xeromorphic characters
necessary for survival in the dry season (such as the thick cuticle of the oak
leaves, Zohary, 1955) could perhaps be an obstacle for snail feeding and
digestion. The few specimens found on the north-facing slope were, indeed, all
from more open areas, where annuals, not maquis, predominate. Furthermore,
in regions where the vegetational distinction between north and south is not
J . HELLER
48
obvious, the distinction of the snails between the slopes also became obscure.
In the steppe, where there is almost no thick vegetation, there is no
difference in the mollusc fauna between north and south slopes. Microgeographic distribution is influenced by the distribution of rocky habitats over
the slopes rather than by distribution of annuals, which are common and
plentiful on both sides.
Plant cover in the steppe is higher than in desert regions (40% as compared
with 25%, Orshan & Diskin, 1968), which means that food is presumablymore
abundant. In this respect therinus, which inhabits the area of the Artemisia
herba-alba associations, has more favourable conditions of food abundance
than glabratus, which dwells in the area of Zygophyllum dumosum.
Succession of Buliminus in Israel
The previous section discussed the influence of physical factors upon
Buliminus distribution. I t was shown that rainfall, substratum and, perhaps,
vegetation, could account for many of the distribution borderlines within the
genus.
The present section will refer to certain borderlines which could not be
explained by such physical factors. They may offer evidence that a faunal
succession is today taking place within the genus, involving forces of
competition and mutual geographic exclusion between species or subspecies
inhabiting the same climatic region, whether in the Mediterranean, steppe,
steppe-periphery or in the Dead Sea area. Our discussion would benefit greatly
from fossil evidence, but unfortunately fossil material is very scarce (Table 6).
Our argument is therefore based only upon the present distribution, upon the
factors influencing it as discussed previously, and upon our knowledge of the
general developments in the Middle East as derived from palaeontology (Haas,
1966, 1968; Tchernov, 1968), palaeobotany (Horowitz, 1968, 1970),
palaeoclimatology (Ferrand, 1971) and geobotany (Zohary, 195 5).
Table 6. Buliminus species described from Pleistocene sites in Israel and Lebanon
~~
~~
Lower
Micoquian Mousterian
Aurignac
Pleistocene c. 110,000- c. 30,000- c. 28,000
120,000 B.C. 70,000 B.C.
B.C.
B labrosus
a alepensis
Iamprostatus
damascensis
Buliminus sp.
Buliminus sp.
J.’
J.’
J.’
&;C
J.’
Natufian
c. 8000
B.C.
Post
Natufian
C.
C.
J.’
Y.’
L.’
c.3
J .’
Key: Y., Yarrnouk Valley; L., Lebanon (Ksar ’Akil); C., Carrnel (Camel Caves, Atlith, Geula); J., Judean
Desert (Wadi Dabber, Jericho). 1, Probably misidentified and actually another species, alepensis; 2, today
not found in this area; 3, species unknown.
Sources: Bate (1937), Shalem (1950), Neuville (1951), Biggs (1960), Altena (1962), Haas (1967).
Steppe region
The most widespread species in the steppe region is therinus (Fig. 20). It
extends also to the Jordan Rift Valley, where its survival, in spite of the very
TAXONOMY O F BULIMINUS IN ISRAEL
49
severe conditions prevailing in this valley, is most probably due to its ability to
adapt itself to hot environments by altering the proportions of its shell
(reduction of the diameter of the shell allows the snails to enter narrower
crevices, while reduction of the relative size of the aperture reduces the amount
of water loss through evaporation). In addition, because of its ability to inhabit
a wide spectrum of substrata, therinus has spread over most of the central
Negev. It is not, however, found in the steppe regions of southern Sinai.
This expansion of therinus over the whole steppe region of the central Negev
may be at the expense of another species, sinaiensis, which does not exhibit
such conchological and ecological flexibility. Today sinaiensis is found in two
separate areas (Fig. 26): Har’Arif (Negev) and the steppe region of southern
Sinai. These two populations, 220 km apart, were apparently once connected,
but later separated by the gradual development of the extremely arid desert in
between. I t is possible that when therinus invaded the steppe of the central
Negev, the Far Negev and Central Sinai had already undergone complete
eremiation, so its route t o southern Sinai was blocked. B. therinus has therefore
established itself in the steppe of the central Negev only, where it has excluded
sinaiensis from nearly all steppe localities. In the Negev sinaiensis today remains
only at Har’Arif, the southernmost steppic outpost on the very brim of the
extreme desert. This area is gradually becoming drier, and more extreme desert,
with its well-adapted fauna, is penetrating Har’Arif from the south. At present
the snail species associated with sinaiensis are Albea arabica Pallary, and
Helicella erkeli Kobelt, two typical representatives of the extremely arid fauna.
I t is thus suggested that colonization of the steppe was carried out in two
migratory waves. The ancient species is found in southern Sinai and relics of it
are still found in the central Negev. The new, dominant species is very common
in the central Negev and Judean Desert, but is not found in south Sinai. We
have some evidence that this line of succession is not restricted only to
Buliminus. Another rock-dwelling snail, Jaminia, shows a strikingly similar
distribution pattern: J. “desertorurn” (Rochanaburananda, 1968) is found, like
B. sinaiensis, only in some parts of the central Negev and separately in southern
Sinai (Hebrew University collections) while J. heptodon Martens (like B.
therinus) populates most of the central Negev and Judean Desert. An
alternative hypothesis, that sinaiensis speciated from the periphery of therinus,
is improbable: the two species differ so much in shell proportions, in sculpture
and in their discordant variation, that the probability of one originating from
the other is negligible.
Steppe periphery
A similar sequence of events may have occurred also in the steppe periphery,
in the region receiving 70-120 mm rain per annum. Here two different species,
glabratus and negevensis, are involved. Near the Dead Sea glabratus, a successful
species which (like therinus) has considerable shell plasticity and which is
capable of populating even the smallest piles of stones, covers large areas. I t
encroaches upon Har Hemar, an isolated locality where negevensis is found
(Fig. 22). B. negevensis survives on one of the last slopes before the cliffs falling
t o the Dead Sea Valley, an extremely arid desert with less than 40 mm rain.
Two empty shells of glabratus found at Har Hemar suggest that negevensis may
be ousted also from this refuge. Similarly, also near Hamakhtesh Hagadol, B.
4
50
J . HELLEK
glabratus surrounds negevensis from the north arid east with therinus in the
west. The species is limited to the cliffs of Hamakhtesh Hagadol and its outlet,
on the border of barren Zin Desert.
Dead Sea region
A faunal succession appears to be occurring also along the shores of the Dead
Sea, though here it could be in part because of climatic changes.
B. alepensis is a typical representative of steppe landscapes, ranging from
Jordan to Iran. In past times the species spread over vast areas of the Judean
Desert (Shalem, 1950; see also Table 6). Today, therinus has overtaken most of
these regions and alepensis has been reduced to the narrow strip of cliffs
(20 x 1 km) bordering the northwest of the Dead Sea. I t cannot spread
westwards, into damper and cooler regions, because these are occupied by
therinus. I t is hindered from spreading northwards by the uninhabitable Lisan
marl fo the Rift Valley. To the east lies the Dead Sea, and another species,
lamprostatus, at present inhabits the area t o the south (Fig. 22). Better adapted
t o arid conditions, lamprostatus has invaded the southern Dead Sea region, and
it now seems to be taking over the narrow biotope from alepensis. In referring
t o lamprostatus, Tristram (1865: 537) mentioned that he “never found it north
of Engedi”. Today it is found 1 5 km north of En Gedi. Though it is doubtful
that Tristram’s observations were accurate, had they been so, an advance of
150 m per year would have taken place. Morphological adaptations of
lamprostatus include a thick peristome (allowing better adherence t o the
substratum), a very thick shell (enabling better isolation) and a very glossy
surface (for reflecting radiation). We may also mention the black skin of the
snail, as a possible defence against excessive ultraviolet radiation. To the extent
that success of a species is reflected by its population density, it is noteworthy
that lamprostatus is three times more abundant than alepensis (measured as
number of live snails found per hour), even though alepensis inhabits the
northern cliffs where, as there is more rainfall, more food is abundant.
Even lamprostatus cannot survive the extreme aridity which prevails today in
the far Negev and central Sinai. Where the isohyets drop below 50 mm, snails
must cope with the irregularity of rain as well as with its scarcity. In central
Sinai precipitation is known t o fluctuate from 36 to 0 mm per year. This may
explain why lamprostatus is found, outside the Dead Sea area, only in a few
disconnected localities in central Sinai.
The faunal succession of Buliminus in the steppe and arid areas of south
Israel is schematically presented in Fig. 32.
Mediterranean region
In the Mediterranean region B. labrosus is represented by two subspecies:
The slim 1. spirectinus in the Judean Hills and the robust Z. labrosus from
Samaria northwards (Fig. 4). The border between the two, which includes a
narrow area of hybridization, cuts through a wide range of ecological areas but
remains totally unaffected by them (in contrast with the species’ range as a
whole, which is clearly delimited by physical borders). Strangely, this
inter-subspecific border coincides with that of another rock-dwelling snail,
Levuntina Kobelt, which also has a southern subspecies (s. spiriplana Olivier,
synonym: L. hierosolyma Boissier ; taxonomy follows Pfeiffer, 1949), in the
TAXONOMY O F BULIMINUS IN ISRAEL
'amprostotus
51
I
30 k M
3.5
,'
Figure 32. Evolution and succession of Bulirninus in southern Israel. Arrow-invasion;
fold-retreat .
Judean Hills and a northern one (L. s. caesareana Parreyss) from Samaria
northwards (compare Figs 4 and 15). The distribution patterns of the two
genera are so similar that I was able in the field to predict successfully which
subspecies of B. Zabrosus would turn up a t any given locality by means of the
Levantina present. As they occupy similar biotopes and as their distribution
patterns are so similar, we may assume that a similar past history has shaped
the distribution of the two.
It is suggested that in these genera, colonization of the Israeli Mediterranean
region took place in two migratory waves, the southern subspecies representing
the earlier element. In Levantina, the first wave brought spiriplana, which in
the beginning spread over the entire Mediterranean biotope. At a later stage a
new subspecies, caesareana, invaded from the North (or, possibly, evolved on
the spot: limestone rocks form a discontinuous biotope which may afford
sufficient isolation for speciation even in the centre of the distribution area). I t
52
J . HELLER
pushed out spiriplana which survives in the Judean Hills and a few other
isolated populations (North of Jiftlik, Fig. 15). Similarly, in B. labrosus, 1.
spirectinus represents the ancient element, which the later invader, 1. labrosus,
then met. B. 1. Zabrosus seems to have succeeded in “rolling back” (Mayr
1970: 3 18) 1. spirectinus, while keeping up a narrow belt of hybridization. I t is
interesting that the 1. labrosus populations closest to the confrontation line
with 1. spirectinus have extremely ventricose, stout, large-mouthed shells, and
in their reproductive system the pediculus, epiphallian flagellum and penis are
very elongate and the penial flagellum well-developed. These characters
strongly contrast with those of 1. spirectinus which has a slim, small-mouthed,
tapering shell, a short, stout pediculus, epiphallian flagellum and penis, and no
penial flagellum at all. A similar reverse cline has been recorded also in the
tropical landsnail Partula (Bailey, 1956). Some interbreeding occurs along the
labrosus-spirectinus borderline, as shown by a few intermediate specimens
found on both sides of it. Though perhaps the borderline (LatrunRammallah-Shillo) might have had ecological significance in the past, as
already stated it at present appears to bear none. The extremely variable
samples from Mt. Carmel, which have shell characteristics of both subspecies
(Fig. 8) may represent a local hybrid population, formed before more complete
genetic isolation was achieved.
The above suggestion of faunal succession is supported by two other
rock-dwelling pulmonates with similar distribution patterns (Hebrew University
collections). Cristataria petriboki Pallary is found on Mt. Gilboa and in
Samaria, whereas C. haasi Nordsieck occurs in the Judean Hills and south
Samaria; Jaminia saulcyi Bourguignat is found in the Galilee, Carmel and
Samaria up to the neighbourhood of Nablus, whereas J. chondriformes.
Mousson is found in the Judean Hills, south Samaria and as a relic on Mt.
Gilboa. Altogether then, four pairs of parapatric or allopatric rock-dwelling
snail taxa within the Mediterranean region have borderlines that cut across
Samaria.
An alternative hypothesis, that the northern taxa invaded the Israeli
Mediterranean first, and that at a later stage the northern species evolved from
their periphery, is improbable: Leuantina spiriplana inhabits, aside from the
Judean Mountains, also Karpathos in the Dodecanese Archipelago (Pfeiffer,
1949). Our case is thus not one of peripheral speciation but of isolates
remaining at the margins of the distribution area.
B. 1. spirectinus and 1. labrosus differ in their reaction to environmental
pressures. While spirectinus seems incapable of inhabiting the Mediterranean
batha regions beyond the 500 mm isohyet, labrosus has remained in the drying
regions of the Judean Desert, where it budded off two new taxa: B. diminutus
and B. 1. jiftliki. The Judean Desert is today an arid barrier separating the 1.
labrosus populations of Israel from those of Jordan. In pluvial periods,
however, a more rainy climate prevailed, which caused a temporary spread of
Mediterranean oak forest into the Judean Desert (Horowitz, 1970), together
with a typical Mediterranean fauna such as the woodpecker Dendrocopus
syriacus and the jay Garrulus glandarius, (Tchernov, 1962). We can safely
assume that under pluvial conditions the Judean Desert was inhabited also by 1.
Zabrosus (see also Table 6), and that there was a connection between the Israeli
and Jordanian populations. The extreme eremiation of the Judean Desert was
TAXONOMY O F BULIMINUS I N ISRAEL
53
accompanied, as in therinus, by selection for smaller forms which are able to
penetrate into narrower crevices during the hot, dry summer. Evidence of the
shell and genitalia (see pp. 20-22) leave little doubt that diminutus is but a
dwarfed 1. labrosus, its shell closely resembling the Jordanian populations of
the latter. Furthermore, its southern limit overlaps the borderline between
Levantina s. caesareana and L. s. spiripluna, as though it were, in this aspect, a
former continuation of 1. labroszts. B. diminutus could thus well be the result of
speciation in a local population, under the pressure of severe eremiation.
B. 1. jiftliki, on the other hand, inhabits the northern corner of the Judean
Desert. Climate here is mild, the Jiftlik region being only a border-district of
35
fobrosus
[ 1 spirecfmus
]iff/iki
J
f
--
31
I
/
i
’
30km
I
-
Figure 3 3 . Evolution and succession of B. Iabrosus and B. diminurus in Israel. Arrow-invasion;
fold-retreat.
54
J . HELLER
the Mediterranean region t o the east. This gentle gradient of climate allows a
continuous connection of 1. jiftliki with its mother-taxon, 1. labrosus.
The evolution of B. labrosus and B. dirninutus is schematically presented in
Fig. 3 3 .
We are today witnessing a faunistic succession within the genus Buliminus in
Israel. In the Mediterranean region 1. labrosus is overcoming 1. spirectinus; in
the steppe therinus has nearly completely replaced sinaiensis; in the steppe
periphery glabratus is overcoming negevensis and, near the Dead Sea
lamprostatus is ousting alepensis. In the Judean Mountains, and along the
isohyet 70, on the brim of the extreme desert, the last relics of the ancient
fauna survive. As yet we have no clue as t o from when the ancient fauna dates,
or when the modern fauna first arrived.
ACKNOWLEDGEMENTS
I am greatly indebted to Professor G. Haas, under whose supervision this
work was carried out, for his devoted guidance, interest and encouragement.
My thanks are also due to Dr Y . L. Werner, Dr U. Ritte and E. Tchernov, of
the Hebrew University, for critically reading parts of the manuscript and for
many valuable discussions. I am especially grateful to Professor A. J. Cain of
the University of Liverpool, for kindly reading through the script and for his
many useful comments.
Throughout this study I was greatly assisted by many museums. I thank Dr
H. Jungen (Zoologisches Museum, Zurich), Dr E. Binder (Museum National
d’Histoire Naturelle, Geneve), Dr H. Walden (Naturhistoriska Museet,
Goteborg), Dr A. Anderson (Naturhistoriska Museet, Stokholm), Dr B.
Metivier (Museum National d’Histoire Naturelle, Paris) and Mr J. Peake (British
Museum of Natural History, London).
I am grateful to Mrs M. Rosenthal and Y . Siman for technical assistance, to
Mr A. Niv for the photographs, and to E. Mizrahi who accompanied me on the
field trips. Miss H. Saendler (Oranim) and Mr Z. Bar (Bet-Ha’emeq) kindly put
their collections at my disposal.
Finally, I wish to express my deep thanks to Dr L. Forcart (Basel) for his
generous advice and continued interest in the many difficult taxonomic
problems concerning this work.
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EXPLANATION O F PLATES
PLATE 1
A.
B.
C.
D.
E.
F.
G.
H.
I.
J.
K.
L.
B. labrosus labrosus. Nahal Namer, Upper Galilee.
Detail of A. Notice fine granulation, typical of labrosus.
Detail of A. Strongly reflexed peristome, typical of many shells from Upper Galilee.
B. 1. labrosus var. jordani, Har Meron, Upper Galilee.
B. 1. labrosus var. kervillei, Umrn Safa, southern Samaria.
B. hbrosusjiffliki ssp. nov. Holocype. Jiftlik, eastern Samana.
B. labrosus spirectinus. Nahalin, Judean Hills.
B. diminutus. Marsaba, Judean Desert.
B. therinus. Wadi Qilt, Judean Desert.
Detail of 1. Notice pronounced granulation (same scale as B).
B. alepensis marsabensis. 'Ein Fashkha, Dead Sea area.
B. lamprostatus. 'En Gedi, Dead Sea area.
Zool. J . Linn. Soc., 5 7 (1975)
J . HELLEK
Plate 1
2001.J. Linn. SOC.,
57 (1975)
J . HELL.ER
Plate 2
Zool. J. Linn. Soc., 5 7 (lY75)
J. HELLEK
TAXONOMY O F B U / , / , W I N U S IN ISRAEL
M. B. sinaiensis. Holotype. Har 'Arif, Central Negev.
N. B. glabratus. Wadi Mashash, Judean Desert.
0. B. glabratus. Har Qomot, Central Negev. Notice variation in distinctness of palatal lamella.
Compare also with N.
P. B. negewensis. Holotype. Mezad Zohar.
PLATE 2
A. B. 1. labrosus. Nahal Namer, Upper Galilee. Gradual transition from labrosus t o exochus
within one sample.
B. B. 1. labrosus. Sasa, Upper Galilee. Spindle-shaped shell dominant in central Upper Galilee.
C. B. 1. labrosus. Tarshiha, Upper Galilee. Variation in the extent to which the lip curls around
itself, within o n e sample.
D. B. 1. labrosus. Kafr Manda, Lower Galilee. Gradual transition from t h e cylindric, jordani
form to the more ventricose kerwillei form.
E. B. 1. labrosus. Merkaz Hacarmel, Mt. Carmel. Gradual transition between very ventricose
and very slim shells within o n e sample. This high variation in shell diameter is typical of
many samples from Mt. Carmel.
PLATE 3
A. B. diminufus. Mar-Jaris, Judean Desert. Gradual transition from diminutus to sikesi within
o n e sample.
B. B. therinus. W. Qilt. Shell closely resembling recondirus Germain.
C. B. therinus. Gradients from north-east t o south-west in shell height, maximal diameter,
mouth height and mouth diameter. Localities, from left: Jiftlik, Beer Sheva-Halutza rd.,
Nahal Avdat.
D. Sample from Wadi Mashash, Judean Desert, which is near t h e borderline between alepensis
marsabensis and lamprostarus. T h e left shells are typical marsabensis, the right ones typical
lamprostarus, t h e middle ones are intermediates. This could be evidence that t h e t w o
species interbreed where they meet.
E. B. alepensis marsabensis. Ein Fashkha. Two shells closely resembling B. damaxensis Pallary,
var. minor.
F. B. alepensis marsabensis, Ein Fashkha. Variation in shell diameter.
G. B. alepensis marsabensis, Ein Fashkha. Variation in shell height.
H. B. lamprostarus, 'En Gedi. Four shells similar to B. fourraui (compare with Pallary, 1923:
pl. 1 2 , figs 4-7).
57