ecology and biology of scorpions in palmyra, syria

A. H. SHEHAB, Z. S. AMR, J. A. LINDSELL

333

Turk J Zool

2011; 35(3) 333-341

© TÜBİTAK

doi:10.3906/zoo-0904-19

Ecology and biology of scorpions in Palmyra, Syria

Adwan Hussien SHEHAB1, Zuhair Sami AMR

2,*, Jeremy Arthur LINDSELL

3

1General Commission for Scientifi c Agricultural Research (GCSAR), Douma, P. O. Box 113, Damascus - SYRIA

2Department of Biology, Jordan University of Science & Technology. P. O. Box 3030, Irbid - JORDAN

3Th e Royal Society for the Protection of Birds, Th e Lodge, Sandy, Beds, SG19 2DL - UK

Received: 21.04.2009

Abstract: Near Palmyra, in the Syrian Desert, 5 species of scorpions belonging to 2 families (Buthidae and Scorpionidae)

were observed; Buthacus tadmorensis, Androctonus crassicauda, Leiurus quinquestriatus, Orthochirus scrobiculosus,

and Scorpio maurus palmatus. B. tadmorensis accounted for 80.6% of the total number of recovered or observed

scorpions, while O. scrobiculosus was the least common (1.4%). Pitfall traps proved to be more effi cient at sampling

(304 individuals) than checking under stones (57 individuals). Pitfall trapping results showed that scorpion abundance

diff ered signifi cantly between the 3 survey areas, while their abundance showed no signifi cant diff erence among the 3

areas when employing the under-stone method. Notes on predation of scorpions (interspecifi c and intraspecifi c) and

predators of scorpions are also included. Seasonal abundance and emergence of scorpions is described briefl y. Biometric

data on collected scorpion species indicating their weight are given. Population structure of B. tadmorensis during the

study period is analyzed.

Key words: Scorpion, Buthacus tadmorensis, pitfall trapping, predation, population structure, Syria

Research Article

* E-mail: [email protected]

Introduction

Limited knowledge is available concerning the

scorpions of Syria. Only 2 recent papers addressed

the systematics of the scorpions of Syria (Kabakibi

et al., 1999; Stathi and Lourenço, 2003). Kovařík and

Whitman (2004) listed 5 species of scorpions from

Talilah National Park, within the vicinity of Palmyra,

Syria. On the other hand, there is a complete lack of

species richness and abundance data for scorpions

inhabiting the Syrian Desert.

In the Middle East, Warburg et al. (1980)

studied the scorpion distribution and diversity in

arid regions of northern Palestine. Warburg (2000)

studied several aspects of scorpion cohabitation. Th e

burrowing behavior of Scorpio maurus palmatus was

investigated by Rutin (1996). Fragmentary reports

dealt with predation of other animals on scorpions

in the Middle East (Tigar and Osborne, 2000;

Zlotkin et al., 2003) including reptiles and birds. No

previous study addressed the diversity and ecology of

scorpions in Syria.

Th e systematics of the genus Buthacus was revised

by Kovařík (2005). He placed in synonymy with

B. macrocentrus at least 3 other species, Buthacus

tadmorensis Simon, Buthus pietschmanni Penther, and

Buthacus yotvatensis Levy et al. However, Lourenço


334

(2006) argued on the validity of the new assignment of B. tadmorensis, and concluded that B. tadmorensis is a valid species, not related to B. macrocentrus as asserted by Kovařík (2005). Furthermore, analysis of the morphological characteristics of the species of Buthacus distributed in Sinai and the Middle East, including Syrian specimens, indicate only that Buthacus leptochelys nitzani Levy, Amitai & Shulov, 1973, could be a junior synonym of B. tadmorensis (Lourenço, 2006). For the purpose of uniformity in the literature, we will consider the name B. tadmorensis for the Palmyra scorpion.

While undertaking ecological fi eldwork near

Palmyra, Syria, we collected a large number of

scorpions and made a number of fi eld observations.

Th e aims of the present study were to investigate

scorpion diversity and abundance, compare 2

sampling methods, comment on the population

structure of Buthacus tadmorensis, and document

some fi eld observations of these scorpions.

Materials and methods

Study sites

Palmyra is located in the heart of the Syrian

Desert. Th e annual rain fall is about 127 mm (average

for the last 50 years), with the maximum annual

mean temperature reaching 25 °C and a minimum of

12 °C. January and February are the coldest months

in Palmyra, where in previous years the average

monthly minimum temperature reached -4.4 °C.

June, July and August are the hottest months, and the

monthly temperature can reach 42 °C in July. Near

Palmyra, within the Northern Bald Ibis Reserve,

3 sites were selected as study areas (Figure 1). Th e

following are descriptions for each site:

1. Site 1: about 596 m asl. Located 18 km

northeast of Palmyra. Th is is a fl at arid area,

with small wadi beds formed by fl ash fl oods.

Scattered stones are abundant over the hard

soil. Sand sheets caused by wind action

are formed in some parts of this area and

dominant vegetation consists of Peganum

harmala with less than 1% cover. Other plants

include Achillea fragantissima, Poa sinaica,

and Haloxylon articulata.

2. Site 2: about 732 m asl. Located 18 km northwest of site 1. Th is is an arid area with hard soil and few rocks. It is a fl at area with wadi systems. Vegetation cover is about 10%, with Anabasis syriaca being the dominant species. Other less common plants include H. articulata and Poa sinaica.

3. Site 3: altitude 924 m asl. Located 19 km west of site 2. A fl at arid area with loose soil and scarce stones. Vegetation cover is less than 1%

Figure 1. Landscape of the 3 study sites.


335

and consists of Anabasis syriaca, H. articulata

and Poa sinaica.

Within the study area, we identifi ed numerous

species of arthropods that were collected in pitfall

traps or were observed under stones. Hymenoptera

(ants) and coleopterans were the most frequently

encountered insects and represented several families

(Tenebrionidae: Trachiderma hispida, Erodius sp.,

Mesostena sp., Akis sp., Adesmia sp., Pimelia sp.,

Zophosis sp.; Carabidae: Calosoma chlorosticum;

Buprestidae: Julodis distincta; and Scarabaeidae:

Scarabaeus sacer). Solphugidae, Empusidae

(Berpharopsis sp.), and Acrididae were less abundant.

Scorpion collecting

Pitfall transects:

In each study site, 6 transects were selected. For

each transect, 5 plastic containers (13 × 9 cm) were

leveled in the ground, 50 m apart, and were without

bait. Scorpions were inspected, measured, identifi ed,

and then released every 2-7 days beginning from

April 2008 to the end of July 2008. Other arthropods

that were trapped in the plastic containers were also

identifi ed where possible, counted, and then released.

Under-stone collecting:

At each site, 20 stones (15-30 cm in diameter)

were lift ed to check for the presence of scorpions.

Th is method was employed every 15 days beginning

from April until the end of July.

Data analysis

Th e association of scorpion abundance with

locality and methods of survey were analyzed using

Genstat 7th edition soft ware for general analysis of

variance.

Predation activity

During the fi eld work, various activities of

collected scorpions were recorded, including

scorpion-scorpion predation and predation of other

animals on scorpions. Th e predation of these animals

was documented with digital images.

Biometry

Total body length (from the anterior edge of

the prosoma to the end of the telson) for collected

scorpions was measured by means of a digital calliper

(measuring range: 0-150 mm), and scorpions were

weighed using a digital balance (0.01 g increments).

Results

Scorpion diversity

A total of 361 scorpions representing 5 species (Figure 2) belonging to 2 families (Buthidae and Scorpionidae) were collected from the study area: Buthacus tadmorensis (Simon, 1892), Androctonus crassicauda (Olivier, 1807), Leiurus quinquestriatus Hemprich & Ehrenberg, 1829, Orthochirus scrobiculosus (Grube, 1873), and Scorpio maurus palmatus (Hemprich & Ehrenberg, 1829).

Table 1 shows the number of recovered species from the 3 study sites using the pitfall traps and under-stone collecting method. By far, B. tadmorensis was the most commonly collected species for both methods, accounting for 63.6%-86.6% of the total number of collected scorpions from all 3 sites. Orthochirus scrobiculosus was the least common species (2.7%-3.9% of the total number of collected scorpions) and was recovered from 2 areas. In site 2, B. tadmorensis was found in pitfall traps along with A. crassicauda, O. scrobiculosus, and L. quinquestriatus.

Comparison between pitfall and under-stone sampling methods

Pitfall trapping proved to be more effi cient than the under-stone method; while the fi rst method yielded 304 scorpions, the second yielded only 57 scorpions.

Both pitfall traps and searching under stones recorded 5 scorpion species during the entire period of the study. Orthochirus scrobiculosus and S. m. palmatus were recovered from under stones during April only.

Pitfall trapping method

Pitfall results showed that scorpion abundance diff ered signifi cantly between the 3 areas; Site 1 (M = 1.4 scorpions), Site 2 (M = 2.8 scorpions), and Site 3 (M = 3.4 scorpions), with the lowest in Site 1 and highest in Site 3 (F = 3.83, P = 0.028). Th e diff erence was signifi cant only between Site 3 and Site 1, while no signifi cant diff erences were observed in scorpion abundance between Site 2 and Site 3 or Site 2 and Site 1 (LSD = 1.487).

A signifi cant diff erence was found between the 5 scorpion species trapped in pitfalls (F = 41.78, P < 0.001), where B. tadmorensis was the most abundant


336

(M = 10.33 scorpions), while no signifi cant diff erence was found between the other 4 species themselves (LSD = 1.919), A. crassicauda (M = 0.63 scorpions), L. quinquestriatus (M = 1.21 scorpions), O. scrobiculosus (M = 0.13 scorpions), and S. m. palmatus (M = 0.38 scorpions).

Under-stone method

Scorpion abundance employing the under-stone method showed no signifi cant diff erence between the 3 areas: Site 1 (M = 0.25 scorpions), Site 2 (M = 0.50 scorpions), and Site 3 (M = 0.67 scorpions) (F = 2.74, P = 0.073).

A signifi cant diff erence was found among the 5 scorpion species recovered from under stones (F = 19.82, P < 0.001), where B. tadmorensis was the most abundant scorpion species (M = 1.79 scorpions), while no signifi cant diff erence was found among the other 4 species (LSD = 0.471), A. crassicauda (M = 0.12 scorpions), L. quinquestriatus (M = 0.29 scorpions), O. scrobiculosus (M = 0.08 scorpions), and S. m. palmatus (M = 0.08 scorpions).

Seasonal abundance of scorpions

Table 2 shows the number of collected scorpions over the study period. Numbers of scorpions collected

A B

C D

E

Figure 2. Scorpions collected from the study area. A) Scorpio maurus palmatus,

B) Androctonus crassicauda, C) Orthochirus scrobiculosus, D) Leiurus

quinquestriatus, and E) Buthacus tadmorensis.


337

in pitfalls increased signifi cantly over the season (F =

5.02, P < 0.001), with the highest levels in July (LSD

= 2.428); the number of scorpions observed under

stones decreased signifi cantly over the season (F =

2.28, P = 0.041), with the highest levels in the second

half of April (LSD = 0.596). For instance, both B.

tadmorensis and L. quinquestriatus increased about

six-fold.

Buthacus tadmorensis was the most common

species throughout the study period (Table 2),

reaching the highest level in July, with A. crassicauda

ranked second. Numbers of O. scrobiculosus

remained constant over the study period.

Predation by scorpions of other arthropods

We observed several occasions where scorpions

predated on other arthropods as well as other

scorpions; B. tadmorensis was observed feeding on a

mantid of the genus Eremiaphila sp. during day time

in a pitfall trap, while B. tadmorensis was observed

feeding on a 1-cm spider.

Interspecifi c predation among scorpions was

observed; an adult L. quinquestriatus fed on all

body parts except the metasoma of B. tadmorensis.

Intraspecifi c predation was also observed, with

an adult B. tadmorensis attacking a 3-cm young

individual from its chelicerae. In captivity, a gravid

Table 1. Recovered scorpions employing under-stone method and pitfall transects in the study sites.

Scorpion speciesUnder stones Pitfalls transect

TotalSite 1 Site 2 Site 3 Site 1 Site 2 Site 3

N % N % N % N % N % N % N %

A. crassicauda 1 9.1 2 10 0 0 3 5.4 8 7.1 4 2.9 18 5

B. tadmorensis 7 63.6 14 70 22 84.6 40 71.4 97 86.6 111 81.6 291 80.6

L. quinquestriatus 3 27.3 1 5 3 11.5 6 10.7 2 1.8 21 15.4 36 10

O. scrobiculosus 0 0 1 5 1 3.9 0 0 3 2.7 0 0 5 1.4

S. m. palmatus 0 0 2 10 0 0 7 12.5 2 1.8 0 0 11 3

Total 11 100 20 100 26 100 56 100 112 100 136 100 361 100

Table 2. Th e number of recovered scorpions using opportunistic collecting and pitfall traps during the

study period from the 3 sites.

Scorpion speciesMethod of

collecting

MonthTotal

April May June July

A. crassicaudaUnder stone 1 0 2 0 3

Pitfall traps 2 6 1 6 15

B. tadmorensisUnder stone 12 10 12 9 43


L. quinquestriatusUnder stone 1 0 4 2 7


O. scrobiculosusUnder stone 2 0 0 0 2


S. m. palmatusUnder stone 2 0 0 0 2


Total 40 61 120 140 361


338

female of B. tadmorensis gave birth to 21 individuals,

all of which later died; the mother ate all of them

within a day.

Predators of scorpions

During our study we observed several animals,

including beetles, ants, lizards, and birds, feeding on

scorpions. On one occasion, a lizard of the species

Acanthodactylus boskianus viciously attacked a 5-cm

B. tadmorensis (Figure 3A). Northern Bald Ibis,

Geronticus eremita, was photographed by Lubomir

Peske during June 2007 at Site 3, while feeding on B.

tadmorensis (Figure 3B). Droppings of the Bald Ibis

collected under the Ibis nesting sites yielded a few

remains of B. tadmorensis. Within the pitfall traps,

remains of B. tadmorensis were found in association

with the presence of ants, Trachyderma hispida,

Erodius sp., and Pimelia sp. Pellets collected near a

nest of a Little Owl, Athene noctua, revealed many

remains of B. tadmorensis, S. m. palmatus, and A.

crassicauda.

Biometric data on scorpions

A total of 14 B. tadmorensis representing diff erent

sizes were weighed. Scorpions measuring 4, 5, and

6 cm in length had an average weight of 0.39 g (n

= 6, 0.29-0.57 g, SD ± 0.124), 0.466 g (n = 5, 0.33-

0.62 g, SD ± 0.113), and 0.95g (n = 3, 0.6-1.26 g,

SD ± 0.332) respectively. A single O. scrobiculosus

measuring 2 cm in length weighed 0.08 g, and 2 A.

crassicauda measuring 8 cm each had an average weight of 3.33g (n = 2, 2.68-3.98 g). We also collected 4 L. quinquestriatus measuring 2 cm (n = 1) and 6 cm (n = 3, 1-1.98 g, SD ± 0.494), with an average weight of 0.33 g and 1.53 g, respectively, and 4 S. m. palmatus with body length ranging from 3-7 cm, weighing 0.36 g (3 cm, n = 1), 2.05 g (6 cm, n = 2, 1.47-2.6 g, SD ± 0.841), and 2.22 g (7 cm, n = 1).

Population structure of B. tadmorensis

Buthacus tadmorensis was the most common species, allowing us to study variation in size in order to determine population structure. Scorpions measuring 2 cm constituted the highest percentage of the population (34.1%), and larger scorpions reaching about 6 cm constituted 4.9% of the population (Figure 4). Th e observed diff erences among collected specimens were probably due to diff erent instars or sex of the arthropod. One of the 6-cm long specimens was a gravid specimen of B. tadmorensis, which gave birth to 21 young during 24 July 2008.

Size groups of collected B. tadmorensis were similar among the 3 study sites (Figures 4 and 5). Small scorpions measuring 2 cm in length were the most common in all study areas. Th e number of small scorpions increased dramatically over the study period. Large scorpions measuring 5 cm increased to a maximum in July and the largest size recorded in this study was 6 cm in length, although their numbers were very low throughout the study.

AB

Figure 3. Predators of scorpions. A) Lizard, Acanthodactylus boskianus feeding on B. tadmorensis. B) Th e Northern Bald Ibis, Geronticus

eremita feeding on B. tadmorensis.


339

Discussion

Diff erent methods have been evaluated for

sampling various terrestrial arthropods, including

pitfall traps, active searching of quadrats, wet cloths,

and drive transects (Tigar and Osborne, 1997;

Druce et al., 2004). In general, scorpions were most

eff ectively sampled using pitfall traps. Our results

are in agreement with Druce et al. (2004), for whom

pitfall trapping resulted in a higher number of

recovered samples than searching under stones. Even

if active searching at night using ultraviolet light

is considered the most eff ective sampling method

(Sissom et al., 1990; Warburg, 1997), this method is

time and eff ort consuming and requires several fi eld

biologists, so we did not test it.

Th e majority of the scorpion species recovered

during this study were collected using pitfall traps,

while O. scrobiculosus and S. m. palmatus were

recovered during April by searching under stones.

Th is is perhaps due to the mild temperature during

April, compared with the hot summer months when

both species retreat to their deep burrows (Amr

and Al-Oran, 1994). Rutin (1996) found that S. m.

palmatus was active during April, excavating soil for

burrow preparation.

Relative abundance of scorpions was studied

among some North American species using pitfall

traps (Bradley and Brody, 1984). It was found that

Paruroctonus utahensis was the dominant species in fl at grassland habitat, along with other species in low numbers. Other species (Vejovis coahuilae and Vejovis russelli) were associated with rocky cliff habitat. In our study, Site 3 had the highest number of scorpions, and B. tadmorensis was the most dominant species. Th is is perhaps due to the dominant loose soil, while the other 2 sites (Site 1 and Site 2) have harder and drier soil textures.

Höfer et al. (1996) studied the surface abundance of the scorpion Brotheas amazonicus, using pitfall traps. Th ey found that capture rates in the primary plateau forest and in the primary forest on white sand soil were higher than in disturbed areas. Th ey concluded that high structural diversity, ultimately refl ecting the availability of hiding places, is important for B. amazonicus. Our study sites were located within the arid Irano-Turanian biotope, with variations in soil hardness and vegetation cover. Structural habitat diversity in the study sites are low, and consist of burrows, scattered stones, and hiding places under bushes.

It seems that emergence from burrows at night is related to decreasing levels of ambient temperature and insect abundance. Th is opinion by shared with Polis (1980). In the study areas, several species of terrestrial arthropods were also trapped in the pitfall traps, with the highest number of various arthropods recorded from Site 3.

0

5

10

15

20

25

30

35

40

45

2 3 4 5 6Length in cm

Site 1

Site 2

Site 3

No.

of s

corp

ion

0

5

10

15

20

25

30

35

40

April May June July

No.

of s

corp

ions

2 cm

3 cm

4 cm

5 cm

6 cm

Figure 4. Comparison between body length of collected B.

tadmorensis from the 3 study sites. Combined data

from under-stone collecting and pitfall traps.

Figure 5. Body length of collected B. tadmorensis from all the

study sites using under-stone collecting and pitfall

traps.


340

Many species of vertebrates and invertebrates were reported to feed on scorpions. Polis et al. (1981) listed about 150 species that feed on scorpions. Among vertebrates, birds and lizards were the most signifi cant predators. Th ese studies indicated predation of scorpions by the desert monitors, Varanus sp. (Millot and Vachon, 1949; Cisse, 1972), the Common Kestrel, Falco tinnunculus, and the Woodchat Shrike Lanius senator, in Eurasia and Africa (Lourenco and Dekeyser, 1976). Th is study reports, for the fi rst time, predation by Acanthodactylus boskianus on B. tadmorensis. Ptyodactylus spp. was found to feed on scorpions both naturally and under laboratory conditions (Zlotkin et al., 2003). It seems that Acanthodactylus boskianus can tolerate the venom of B. tadmorensis, which is less potent for humans than L. quinquestriatus.

Th e Little Owl and the Northern Bald Ibis also predated on this species. Owls were reported to feed on scorpions in the Middle East; pellets of the Desert Eagle Owl, Bubo ascalaphus were found to include remains of A. crassicauda, Compsobuthus werneri, and S. m. palmatus (Rifai et al., 2000). Shehab (2004) found that scorpions constituted 9.22% of prey items of the Eagle Owl in the Syrian Desert, while the Little Owl fed on other unidentifi ed scorpions (Obuch and Krištìn, 2004). Th e diet of other birds such as the Houbara Bustard (Chlamydotis macqueenii) included

Buthacus yotvatensis nigroaculeatus and Vachoniolus spp. in Abu Dhabi (Tigar and Osborne, 2000). Vachon (1952) reported predation of scorpions by the Northern Bald Ibis, Geronticus eremta, in Southeastern Europe, Asia and Northern Africa.

Scorpions will capture almost any prey that they can overcome. Th ey feed primarily on several groups of insects, arachnids, and other arthropods (Williams, 1987). Intraspecies predation was reported among many scorpion species in North America (Polis, 1979). We report here predation of L. quinquestriatus on B. tadmorensis and cannibalism among B. tadmorensis. Maternal cannibalism in B. tadmorensis was observed, although only in captivity.

Acknowledgments

We would like to thank Professor Adel Safar, the Minister of Agriculture and Agrarian Reform, Engineer Ali Hamoud, the Director General of the General Commission of Al Badia Development and Management, and the General Commission for Scientifi c Agricultural Research for facilitating the fi eld work, and for their encouragement and continuous support. We also wish to express our gratitude to BirdLife International for supporting this study.

References

Amr, Z. S. and Al-Oran, R. 1994. Systematics and distribution of

scorpions (Arachnida, Scorpionida) in Jordan. Bollettino di

Zoologia. 61: 185-190.

Bradley, R. A. and Brody, A. J. 1984. Relative abundance of three

vaejovid scorpions acr zoss a habitat gradient. Journal of

Arachnology. 11: 437-440.

Cissé, M. 1972. Th e diet of Varanids in Senegal. Bull. Inst. Fond. Afr.

Noir. 34: 503-515.

Druce, D., Hamer, M. and Slotow, R. 2004. Sampling strategies for

millipedes (Diplopoda), centipedes (Chilopoda) and scorpions

(Scorpionida) in savanna habitats. African Zoology. 39: 293-

304.

Höfer, H., Wollscheid, E. and Gasnier, T. 1996. Th e relative

abundance of Brotheas amazonicus (Chactidae, Scorpiones) in

diff erent habitat types of a Central Amazon Rainforest. Journal

of Arachnology. 24: 34-38.

Kabakibi, M. M., Khalil, N. and Amr, Z. S. 1999. Scorpions of

southern Syria. Zoology in the Middle East. 17: 79-89.

Kovařík, F. 2005. Taxonomic position of species of the genus

Buthacus Birula, 1908 described by Ehrenberg and Lourenço,

and description of a new species (Scorpiones: Buthidae).

Euscorpius. 28:1-13.

Kovařík, F. and Whitman, S. 2004. cataloghi del museo di storia

naturale dell’università di Firenze-sezione di zoologia «la

specola» xxii. arachnida scorpiones. tipi. addenda (1988-2004)

e checklist della collezione (euscorpiinae esclusi). Atti Soc.

Tosc. Sci. nat., Mem., Serie B. 111: 103-119.

Lourenço, W. R. 2006. Further considerations on the genus Buthacus

Birula, 1908 (Scorpiones, Buthidae), with a description of

one new species and two new subspecies. Boletín Sociedad

Entomológica Aragonesa. 38: 59-70.

Lourenço, W. R. and Dekeyser, P. 1976. Two bird predators on

scorpions. Oiseau Rev. Fr. Ornithol. 46: 167-172.

Millot, J. and Vachon, M. 1949. Ordre des scorpions. Pp. 386-436,

in Traite de Zoologie (P. P. Grasse, ed.), Paris, Masson et Cie,

Vol. 6.


341

Obuch, J. and Kristín, A. 2004. Prey composition of the little owl

Athene noctua in an arid zone (Egypt, Syria, Iran). Folia

Zoologica. 53: 65-79.

Polis, G.A. 1979. Diet and prey phenology of the desert scorpion,

Paruroctonus mesaensis Stahnke. Journal of Zoology (London).

188: 333-346.

Polis, G. A. 1980. Seasonal patterns and age-specifi c variation in the

surface activity of a population of desert scorpions in relation

to environmental factors. Th e Journal of Animal Ecology. 49:

1-18.

Polis, G. A., Sissom, W. D. and McCormick, S. J. 1981. Predators

of scorpions: Field data and a review. Journal of Arid

Environments. 4: 309-327.

Rifai, L. B., Al-Melhim, W. N., Gharaibeh, B. M. and Amr, Z. S. 2000.

Th e diet of the Desert Eagle Owl, Bubo bubo ascalaphus, in the

Eastern Desert of Jordan. Journal of Arid Environments. 44:

369-372.

Rutin, J. 1996. Th e burrowing activity of scorpions (Scorpio maurus

palmatus) and their potential contribution to the erosion of

Hamra soils in Karkur, central Israel. Geomorphology. 15: 159-

168.

Shehab, A. H. 2004. Diet of the Eagle Owl, Bubo bubo, in Syria.

Zoology in the Middle East. 33: 21-26.

Sissom, W. D., Polis, G. A. and Watt, D. D. 1990. Field and laboratory

methods. In: Th e Biology of Scorpions, G. A Polis (ed.),

Stanford University Press, Stanford. pp. 445-461.

Stathi, I. and Lourenço, W. 2003. Description of a new scorpion

species of the genus Birulatus Vachon, 1974 (Scorpiones,

Buthidae) from Syria. Zoology in the Middle East. 30: 105-110.

Tigar, B. J. and Osborne, P. E. 1997. Patterns of arthropod abundance

and diversity in an Arabian desert. Ecography. 20: 550-558.

Tigar, B. J. and Osborne, P. E. 2000. Invertebrate diet of the Houbara

Bustard Chlamydotis [undulata] macqueenii in Abu Dhabi

from calibrated faecal analysis. Ibis. 142: 466-475.

Vachon, M. 1952. Études sur les Scorpions. Institut Pasteur d’Algérie,

Alger. 482 pp.

Warburg, M. R. 1997. Biogeographic and demographic changes

in the distribution and abundance of scorpions inhabiting

the Mediterranean region in northern Israel. Diversity and

Conservation. 6: 1377-1389.

Warburg, M. R. 2000. Intra- and interspecifi c cohabitation of

scorpions in the fi eld, and the eff ect of density, food and shelter

on their interactions. Journal of Ethology. 18: 59-63.

Warburg, M. R., Goldenberg, S., and Ben-Horin, A. 1980. Scorpion

species diversity and distribution within the Mediterranean and

arid regions of northern Israel. Journal of Arid Environments.

3: 205-213.

Williams, S. C. 1987. Scorpion bionomics. Annual Review of

Entomology. 32: 275-95.

Zlotkin, E., Milman T., Sion G. and Werner, Y.L. 2003. Predatory

behaviour of gekkonid lizards, Ptyodactylus spp., towards the

scorpion Leiurus quinquestriatus hebraeus, and their tolerance

of its venom. Journal of Natural History. 37: 641-646.

ecology and biology of scorpions in palmyra, syria

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