1. Introduction

2. Materials and Methods

Figure 1. Maps of Israel and adjacent countries. (a) Scheme of biogeographical units in Israel and adjacent countries (adopted from Klein 1988): 1. Upper Galilee; 2. Lower Galilee; 3. Carmel Ridge; 4. Northern Coastal Plain; 5. Jezreel Valley; 6. Samaria; 7. Jordan Valley and Sea of Galilee; 8. Central Coastal Plain; 9. Southern Coastal Plain; 10. Judea Foothills; 11. Judean Hills; 12. Judean Desert; 13. Dead Sea Area; 14. ‘Arava Valley; 15. Northern Negev; 16. Southern Negev; 17. Central Negev; 18. Golan Heights; 19. Mount Hermon; 20-22. Sinai (Egypt): (Northern Sinai; Central Sinai Foothills; Sinai Mountains). (b) Map of biogeographic sub-regions in Israel and adjacent countries, that was used by us in this publication (numbers in parentheses relate to biogeographical units in Figure 1a): 1. Hermon (19) 2. Golan (18) 3. Galilee (1 & 2) 4. Hula Valley (included in 1) 5. Coastal Plain (4, 8 & 9) 6. Jezreel Valley (5) 7. Central Mountains (Carmel, Samaria, Judea hills and Judea foothills; 3, 6, 10 & 11) 8. Dead Sea and Jordan Valley (7 & 13) 9. Judean Desert (12) 10. Negev Desert (Northern Negev, Central Negev, Southern Negev; 15, 16 & 17) 11. ‘Arava Valley (14).

Figure 1. Maps of Israel and adjacent countries. (a) Scheme of biogeographical units in Israel and adjacent countries (adopted from Klein 1988): 1. Upper Galilee; 2. Lower Galilee; 3. Carmel Ridge; 4. Northern Coastal Plain; 5. Jezreel Valley; 6. Samaria; 7. Jordan Valley and Sea of Galilee; 8. Central Coastal Plain; 9. Southern Coastal Plain; 10. Judea Foothills; 11. Judean Hills; 12. Judean Desert; 13. Dead Sea Area; 14. ‘Arava Valley; 15. Northern Negev; 16. Southern Negev; 17. Central Negev; 18. Golan Heights; 19. Mount Hermon; 20-22. Sinai (Egypt): (Northern Sinai; Central Sinai Foothills; Sinai Mountains). (b) Map of biogeographic sub-regions in Israel and adjacent countries, that was used by us in this publication (numbers in parentheses relate to biogeographical units in Figure 1a): 1. Hermon (19) 2. Golan (18) 3. Galilee (1 & 2) 4. Hula Valley (included in 1) 5. Coastal Plain (4, 8 & 9) 6. Jezreel Valley (5) 7. Central Mountains (Carmel, Samaria, Judea hills and Judea foothills; 3, 6, 10 & 11) 8. Dead Sea and Jordan Valley (7 & 13) 9. Judean Desert (12) 10. Negev Desert (Northern Negev, Central Negev, Southern Negev; 15, 16 & 17) 11. ‘Arava Valley (14).

Figure 2. Israel’s biogeographic sub-regions A. Hermon B. Golan C. Galilee D. Hula Valley E. Coastal Plain F. Jezreel Valley G. Central Mountains H. Jordan Valley I. Dead Sea J. Judean Desert K. Negev Desert L. Arava Valley. Photos by I. Armiach Steinpress.

Figure 2. Israel’s biogeographic sub-regions A. Hermon B. Golan C. Galilee D. Hula Valley E. Coastal Plain F. Jezreel Valley G. Central Mountains H. Jordan Valley I. Dead Sea J. Judean Desert K. Negev Desert L. Arava Valley. Photos by I. Armiach Steinpress.

3. Results

3.1. Systematics and Taxonomy

Based on all specimens deposited at the NNHC, HUJ (over 1600, as well as fourteen Type specimens) (Table S1), published records and the World Pseudoscorpiones Catalog [1], 61 pseudoscorpion species of 28 genera in fourteen families are now recorded from Israel. The list of the fourteen families, arranged systematically following Benavides et al. [19], is presented in Table 2. Two families are reported here for the first time from Israel: Syarinidae Chamberlin, 1930 and Cheiridiidae Hansen, 1894. Syarinidae is a widely distributed cosmopolitan family with 125 species known worldwide, most of which occur in America. Yet, several relict species are found sporadically and rarely in Europe and the Mediterranean region, with most of them being troglobionts. Such is the one we report here, which was collected in a cave in Samaria, in the Central Mountains (Figure 1b, sub-region 7). Cheiridiidae is a widely distributed cosmopolitan family with 79 species known worldwide, that has common representatives in the East Mediterranean region including Turkey (three species), Italy and Bulgaria (two species in each) and Greece, Croatia and Montenegro (one species in each), and was expected to occur also in Israel. The species we report here was collected from a cave in the Judean Desert (Figure 1b, sub-region 9).

3.1.1. Figures and Tables

Based on the results presented in the following tables and figures (Table 3 and Figure 3, Figure 4 and Figure 5), we provide hereafter information on each recorded family, followed by a taxonomic key to all families.

Table 3. List of the pseudoscorpion species in Israel arranged by alphabetical order of family, genus and species, their record from Israel (in [ ]), chorotype classification, chorological complexes, and global zoographic category (based on Harvey [18] and WPC [1]). The chorotype classifications and respective codes, as well as their attribution to chorological complexes are detailed in Table 1. New records for Israel are marked with an asterisk *, ^e indicates species endemic to Israel, ^T indicates Type specimen deposited at the NNHC, HUJ and ^¢ indicates that no specimens are deposited at the HUJ.

Table 3. List of the pseudoscorpion species in Israel arranged by alphabetical order of family, genus and species, their record from Israel (in [ ]), chorotype classification, chorological complexes, and global zoographic category (based on Harvey [18] and WPC [1]). The chorotype classifications and respective codes, as well as their attribution to chorological complexes are detailed in Table 1. New records for Israel are marked with an asterisk *, ^e indicates species endemic to Israel, ^T indicates Type specimen deposited at the NNHC, HUJ and ^¢ indicates that no specimens are deposited at the HUJ.

Family	Species	Global Zoogeographic category	Chorotype	Chorological Complex
Atemnidae
	Atemnus sp. *	Palearctic and Afrotropic	OLD	Widely distributed
	Atemnus syriacus (Beier, 1955) ¢ [16]	Palearctic	EMED	Mediterranean
Cheiridiidae *
	Apocheiridium sp. *	Palearctic	COS	Widely distributed
Cheliferidae
	Chelifer cancroides (Linnaeus, 1758) [15]	Palearctic and Afrotropic	COS	Widely distributed
	Chelifer cancroides cancroides (Linnaeus, 1758) ¢ [1]	Palearctic and Afrotropic	COS	Widely distributed
	Dactylochelifer kussariensis (Daday, 1889) [15]	Palearctic	CWA	Asian
	Dactylochelifer kussariensis kussariensis (Daday, 1889) ¢ [1]	Palearctic	CWA	Asian
	Dactylochelifer pallidus Beier, 1963 e, T [15]	Endemic (Palearctic origin)	END	Endemic
	Hysterochelifer cyprius (Beier, 1929) [15]	Palearctic	EMED	Mediterranean
	Hysterochelifer distinguendus (Beier, 1929) e [39]	Endemic (Palearctic origin)	END	Endemic
	Hysterochelifer gracilimanus Beier, 1949 ¢ [17]	Palearctic	EMED	Mediterranean
	Hysterochelifer tuberculatus(Lucas, 1849) [15]	Palearctic	MED	Mediterranean
	Hysterochelifer tuberculatus tuberculatus (Lucas, 1849) ¢ [1]	Palearctic	MED	Mediterranean
	Rhacochelifer corcyrensis (Beier, 1930) ¢ [17]	Palearctic	MED	Mediterranean
	Rhacochelifer corcyrensis bicolor Beier, 1963 T [15]	Palearctic	LEV	Mediterranean
	Rhacochelifer corcyrensis corcyrensis (Beier, 1930) ¢ [17]	Palearctic	EMED	Mediterranean
	Rhacochelifer lobipes (Beier, 1929) [15]	Palearctic	EMED	Mediterranean
	Rhacochelifer longeunguiculatus Beier, 1963 T [15]	Afrotropic	APA	Afrotropic
	Rhacochelifer maculatus (L. Koch, 1873) [17]	Palearctic	MED	Mediterranean
	Rhacochelifer peculiaris (L. Koch, 1873) [1]	Palearctic	SEM	Mediterranean
	Rhacochelifer peculiaris latissimus Beier, 1963 T [15]	Palearctic	EMED	Mediterranean
Chernetidae
	Allochernes masi (Navás, 1923) [15]	Palearctic	MED	Mediterranean
	Chernes rhodinus Beier, 1966 [16]	Palearctic	EMED	Mediterranean
	Chernes sp. *	Palearctic	HOL	Widely distributed
	Lamprochernes nodosus (Schrank, 1803) [15]	Palearctic and Afrotropic	OLD	Widely distributed
	Lamprochernes nodosus nodosus (Schrank, 1803) ¢ [1]	Palearctic and Afrotropic	OLD	Widely distributed
	Lamprochernes savignyi (Simon, 1881) [15]	Afrotropic	COS	Widely distributed
	Lasiochernes turcicus Beier, 1949 [15]	Palearctic	EMED	Mediterranean
	Nudochernes spalacis Beier, 1955 e, T [41]	Endemic (Afrotropic origin)	END	Endemic
	Pselaphochernes scorpioides (Hermann, 1804) ¢ [16]	Palearctic	HOL	Widely distributed
Chthoniidae
	Ayyalonia dimentmani Ćurčić, 2008 e [13]	Endemic (Afrotropic origin)	END	Endemic
	Chthonius jonicus Beier, 1931 ¢ [16]	Palearctic	MED	Mediterranean
	Chthonius shulovi Beier, 1963 e ¢ [15]	Endemic (Palearctic origin)	END	Endemic
	Chthonius sp. *	Palearctic	COS	Widely distributed
	Ephippiochthonius sacer (Beier, 1963) e, T [15]	Endemic (Palearctic origin)	END	Endemic
	Ephippiochthonius tetrachelatus (Preyssler, 1790) [15]	Palearctic	COS	Widely distributed
	Ephippiochthonius spp. *	Palearctic	COS	Widely distributed
Garypidae
	Garypus beauvoisii (Audouin, 1826) [15]	Palearctic	MED	Mediterranean
	Garypus levantinus Navás, 1925 [15]	Palearctic	MED	Mediterranean
Garypinidae
	Garypinus asper Beier, 1955 [16]	Palearctic	EMED	Mediterranean
	Garypinus dimidiatus (L. Koch, 1873) [15]	Palearctic	EMED	Mediterranean
Geogarypidae
	Geogarypus pulcher Beier, 1963 e, T [15]	Endemic (Palearctic origin)	END	Endemic
	Geogarypus shulovi Beier, 1963 T [15]	Palearctic	CWA	Asian
Hesperolpiidae
	Calocheirus atopos Chamberlin, 1930 T [15]	Afrotropic	APA	Afrotropic
	Cardiolpium stupidum (Beier, 1963) T [15]	Palearctic	EMCA	Euro-Asia
Menthidae
	Paramenthus shulovi Beier, 1963 e, T [15]	Endemic (Afrotropic origin)	END	Endemic
Neobisiidae
	Neobisium (Neobisium) validum (L. Koch, 1873) [15]	Palearctic	EMCA	Euro-Asia
	Neobisium sp. *	Palearctic	EMCA	Euro-Asia
Olpiidae
	Calocheiridius libanoticus Beier, 1955 [15]	Palearctic	EMWA	Mediterranean
	Halominniza aegyptiaca (Ellingsen, 1910) ¢ [1]	Palearctic	EMED	Mediterranean
	Halominniza aegyptiaca litoralis (Beier, 1963) [15]	Palearctic	LEV	Mediterranean
	Minniza babylonica Beier, 1931 [15]	Palearctic and Afrotropic	CWAP	Asian
	Minniza babylonica babylonica Beier, 1931 ¢ [1]	Palearctic and Afrotropic	CWAP	Asian
	Minniza lindbergi Beier, 1957 [15]	Palearctic	EMED	Mediterranean
	Olpium kochi Simon, 1881 [26]	Palearctic	EMED	Mediterranean
	Olpium pallipes (Lucas, 1849) [39,40]	Palearctic	MCA	Mediterranean
	Olpium pallipes balcanicum Beier, 1931 [15]	Palearctic	EMED	Mediterranean
	Olpium pallipes pallipes (Lucas, 1849) ¢ [1]	Palearctic	MCA	Mediterranean
Syarinidae *
	Hadoblothrus sp. *	Palearctic	EMED	Mediterranean
Withiidae
	Nannowithius wahrmani (Beier, 1963) e, T [15]	Endemic (Afrotropic origin)	END	Endemic

Figure 3. The geographic distribution of the pseudoscorpion families (percentage of specimens) in each of the eleven biogeographic sub-regions of Israel (Figure 1b) (data of 1548 specimens in the collection, see Supplementary Materials Table S2).

Figure 3. The geographic distribution of the pseudoscorpion families (percentage of specimens) in each of the eleven biogeographic sub-regions of Israel (Figure 1b) (data of 1548 specimens in the collection, see Supplementary Materials Table S2).

Figure 4. Geographic distribution maps of the pseudoscorpion species in Israel. Localities of 1397 identified specimens deposited in the collection (circle) (Supplementary Materials Table S3), and localities specified in: Beier [15], Halperin & Mahnert [17], Mahnert [16] and Šťáhlavský et al. [26], (square). Endemic species are indicated by a star symbol. (a) Cheliferidae; (b) Cheliferidae; (c) Olpiidae; (d) Cheliferoidea (Atemnidae, Chernetidae and Withiidae); (e) Neobisioidea (Neobisiidae and Syarinidae); (f) Chthoniidae; (g) Cheiridiidae and Garypinidae; (h) Garypoidea (Garypidae, Geogarypidae, Hesperolpiidae and Menthidae).

Figure 4. Geographic distribution maps of the pseudoscorpion species in Israel. Localities of 1397 identified specimens deposited in the collection (circle) (Supplementary Materials Table S3), and localities specified in: Beier [15], Halperin & Mahnert [17], Mahnert [16] and Šťáhlavský et al. [26], (square). Endemic species are indicated by a star symbol. (a) Cheliferidae; (b) Cheliferidae; (c) Olpiidae; (d) Cheliferoidea (Atemnidae, Chernetidae and Withiidae); (e) Neobisioidea (Neobisiidae and Syarinidae); (f) Chthoniidae; (g) Cheiridiidae and Garypinidae; (h) Garypoidea (Garypidae, Geogarypidae, Hesperolpiidae and Menthidae).

Figure 5. Seasonality, habitat, and collection methodology of pseudoscorpions deposited at the HUJ collection. (A) Seasonal distribution of the pseudoscorpion families in Israel (winter: December-February; spring: March-May; summer: June-August; autumn: September-November) (based on data of 1614 specimens in the collection, Supplementary Materials Table S4). (B) Habitats occupied by pseudoscorpion families in Israel (according to data of specimens in the collection). Data of the same specimen can contribute to more than one category, e.g. desert and under a stone; or cave and related to animals (Supplementary Materials Table S5). (C) Collection method (according to data of 583 specimens in the collection, Supplementary Materials Table S6).

Figure 5. Seasonality, habitat, and collection methodology of pseudoscorpions deposited at the HUJ collection. (A) Seasonal distribution of the pseudoscorpion families in Israel (winter: December-February; spring: March-May; summer: June-August; autumn: September-November) (based on data of 1614 specimens in the collection, Supplementary Materials Table S4). (B) Habitats occupied by pseudoscorpion families in Israel (according to data of specimens in the collection). Data of the same specimen can contribute to more than one category, e.g. desert and under a stone; or cave and related to animals (Supplementary Materials Table S5). (C) Collection method (according to data of 583 specimens in the collection, Supplementary Materials Table S6).

3.1.2. The pseudoscorpion families

Cheliferidae is the largest and most common family deposited in the NNHC HUJ collection. Four genera and eighteen species and subspecies are documented from Israel (Table 3, Figure 4a,b). Only one species is Afrotropic, and the rest are Palearctic, with eleven Mediterranean, two Asian, two cosmopolitan and two endemic species (Table 3). Cheliferids were collected all over the country, in all regions except the Hermon (Figure 3, Figure 4a,b) and all year round. Most cheliferids were collected in spring and summer, and fewer individuals were collected in autumn and winter (Figure 5A). Cheliferids were collected using all methods. They were often found under bark and in leaf litter (Figure 5B) usually in habitats with trees. The subspecies Rhacochelifer corcyrensis bicolor Beier, 1963 was found almost exclusively in the desert, mainly on Acacia trees (Vachellia Wight & Arn) using trunk traps (Figure 4a,b, Figure 5A,C). The cosmopolitan species Chelifer cancroides (Linnaeus 1758) is phoretic and often found in beehives. The genus Dactylochelifer Beier, 1932 (the species D. kussariensis (Daday1889) and the endemic species D. pallidus Beier, 1963) was typically found near wetlands (swamps and riverbanks) (Figure 4a, Figure 5A). The other endemic cheliferid Hysterochelifer distinguendus (Beier, 1929) was found only in the Judea foothills (Figure 4a). Several specimens of Rhacochelifer longeunguiculatus Beier 1963 as well as unidentified species were collected from cave entrances but no troglobiont species were recorded from Israel thus far (Figure 6A).

Olpiidae is the second largest and most common family deposited in the NNHC HUJ collection, both in number of specimens and number of species and subspecies. Four genera and ten species are documented from Israel (Table 3, Figure 4c). Although globally Olpiidae species are usually found in xeric habitats (desert and semi-desert), the species in Israel are mostly Palearctic, with eight species from the Mediterranean and two species from Central and Western Asia and the Arabian Peninsula, thus from both Palearctic and Afrotropic regions (Table 3). Olpiids were collected all over the country, in all regions except the Hula Valley, but were significantly more common in the desert, where more than 75% of the specimens were found, while less than 10% were found in all the northern regions of Israel and the coastal plain (Supplementary Materials Tables S2-S3). They were collected all year, but the highest number was collected in spring while very little were collected in winter (Figure 5A). Specimens were collected by visual search under stones, sifting and in pitfall traps (Figure 5C). None of the olpiid species are endemic to Israel, yet Halominniza aegyptiaca litoralis (Beier, 1963) is endemic to the southern Levant (Israel, Jordan and Sinai), and was found in Israel only in the desert near the Dead Sea and the Red Sea, in association with sea (salty) water (Figure 4c). The Asian species Minniza babylonica Beier, 1931, which is found in both Palearctic and Afrotropic regions, is typically found in Israel in the desert (Figure 4c). This family has not been recorded from caves or moist epigean habitats (Figure 6B).

Chernetidae is the third largest and most common family deposited in the NNHC HUJ collection in both number of specimens and number of species, with six genera and eight species, in addition to undescribed species from caves. Five of these species are widely distributed and three are Mediterranean whereas one is endemic from an Afrotropic origin (Table 3, Figure 4d). Most of the specimens (80%), were collected in the mountains of central Israel and Jezreel valley, while hardly any were collected in the desert (Figure 4d, Supplementary Materials Tables S2-S3). Chernetids were found evenly throughout the year (Figure 5A) and usually by visual search (Figure 5C). Species of Chernetidae are often phoretic on rodents or insects (Figure 5B) and tend to aggregate in their nests as evident by 80% of the family’s specimens in Israel: Lamprochernes savignyi (Simon, 1881) is phoretic on flies, Lasiochernes turcicus Beier, 1949 (Figure 6C) is found in nests of Apodemus Kaup, 1829, and Nudochernes spalacis Beier, 1955 in nests of Nannospalax species. Nearly half of the specimens collected from caves in Israel (44%) belong to the Chernetidae family, being a quarter of the family’s specimens collected in Israel, from the species Allochernes masi, Chernes rhodinus, Lasiochernes turcicus and other unidentified species (Figure 5B). They were found mainly in caves with bats, both frugivorous and insectivorous, and L. turcicus also in an Apodemus nest.

Neobisiidae is the fourth most common family deposited in the NNHC HUJ collection, in terms of number of specimens; however, it is represented in Israel with only one genus and one species throughout the country, the Palearctic Neobisium validum (L. Koch, 1873) (Figure 6D), which is found in the East-Mediterranean as well as Central and Western Asia (Table 3, Figure 4e), with the type locality in Syria. Only 4% of the specimens collected from the desert, while 93% were collected in the mountains of north and central Israel and Jezreel valley (Figure 4e, Supplementary Materials Tables S2-S3). The remaining 3% were collected in the coastal region. Preliminary DNA sequencing (not presented here) indicates a species complex, which needs to be examined further. As a hygrophilous family, Neobisiidae specimens were collected four times more in winter than in spring and autumn, while none were collected during the summer (Figure 5A). They were usually found by sifting soil and leaf litter, visual search under stones or using pitfall traps (Figure 5B,C). One specimen of a presently unidentified troglobitic species was found in a cave.

Chthoniidae is represented in Israel by five species from three genera in addition to several undescribed species from caves. Three species are endemic, one Mediterranean and one cosmopolitan (Table 3, Figure 4f). Chthoniids were found exclusively in the mountains of north and central Israel (Figure 3, Figure 4f, Supplementary Materials Tables S2-S3) or in caves (Figure 5B). Half of the pseudoscorpion specimens we found in Israel’s caves belong to the Chthoniidae, representing 75% of the family’s specimens collected in Israel. Being a family with high short-range endemism [6], we expect several new species to science among the unidentified cave specimens (Figure 6E). As a hygrophilous family, only a few Chthoniidae specimens were collected in the driest season in Israel, autumn (Figure 5A). Chthoniids were collected mostly by visual search under stones, and on bats guano in caves (Figure 5B,C), but they also occur in leaf litter, and some were collected by sifting and pitfall traps. Ayyalonia dimentmani Ćurčić, 2008 is an example of an endemic relict species found only in a single unique chemoautotrophic cave. Another endemic species, Chthonius shulovi Beier, 1963 was found only once in one locality in Wadi Qelt [15]. Both species were never found again. However, the endemic species Ephippiochthonius sacer (Beier, 1963) seems to be more widespread, and was found several times in the Mediterranean region of north and central Israel.

Garypinidae is represented in Israel by two species of Garypinus Dadday, 1889 that have an eastern Mediterranean to Palearctic distribution (Table 3, Figure 4g). Both were found exclusively in the mountains of north and central Israel (Figure 3, Figure 4g, Supplementary Materials Tables S2-S3), mostly in spring and summer (Figure 5A), often by sifting leaf litter or visual search under stones (Figure 5B,C).

Garypidae is represented in Israel by two species of Garypus L. Koch, 1873 that are halophilous and widespread in seashore habitats of the Mediterranean (Table 3, Figure 4h). Both were found in all seasons but winter (Figure 5A), exclusively in the salty habitat of the Mediterranean coast (Figure 3, Figure 4h, Figure 5B). Some specimens were found in algae debris at the coastline.

Geogarypidae is represented in Israel by two species from one genus: Geogarypus shulovi Beier, 1963 is Palearctic from Central and Western Asia, and Geogarypus pulcher Beier, 1963 is endemic to Israel (Table 3, Figure 4h). Most of the specimens were found in the mountains of north and central Israel, but some also in the coastal plain and Hula valley (Figure 3, Figure 4h, Figure 5B, Supplementary Materials Tables S2-S3). They were usually found by sifting leaf litter, mostly in spring (Figure 5B,C).

Hesperolpiidae is represented in Israel by two species from two genera, Calocheirus atopos Chamberlin, 1930 is Afrotropic from the Arabian Peninsula and Sudan, and Cardiolpium stupidum (Beier, 1963) is Palearctic from East-Mediterranean and Central-West Asia (Table 3, Figure 4h). They are found mostly in spring (Figure 5A), in small numbers but in a wide distribution along Israel (Figure 3, Figure 4h, Figure 5B).

Menthidae is an infrequently collected family and represented in our collection by only three specimens; all belong to the endemic species Paramenthus shulovi Beier, 1963; an Afrotropic genus. Each of the specimens was collected from a different sub-region of the country; the Negev desert, the Central Mountains, and the Galilee (Table 3, Figure 4 h).

Atemnidae is represented in our collection by only two specimens, each from a different species and a different region: Atemnus syriacus (Beier, 1955) in the north of Israel, and an undescribed species from the Judean desert (Table 3, Figure 4d) .

Withiidae is represented in our collection by the endemic species Nannowithius wahrmani (Beier, 1963), which belongs to the Afrotropic genus Nannowithius Beier. It is a small myrmecophilous species that was found in nests of the ant Messor semirufus (André, 1883) in the Negev desert, under a stone near the sea of Galilee, and phoretic on the myrmecophilous scorpion Birulatus israelensis which lives only in Messor ebeninus Santschi, 1927 nests (Figure 4d, Figure 6f) [27].

Cheiridiidae is represented in our collection by a single specimen that was collected in a cave in the Judean desert (Figure 4g).

Syarinidae is represented in our collection by a single specimen that was collected in a cave in the Central Mountains (Figure 4e).

Figure 6. Habitus of live pseudoscorpion representatives from Israel. A. Hysterochelifer cyprius (Cheliferidae). B. Olpium pallipes balcanicum (Olpidae). C. Lasiochernes turcicus (Chernetidae). D. Neobisium validum (Neobisiidae). E. Ephippiochthonius sp. (Chthoniidae). F. Nannowithius wahrmani (Withiidae) phoretic on Birulatus israelensis. Photos A, B, D, E, F by S. Aharon, C by J. A. Ballesteros.

Figure 6. Habitus of live pseudoscorpion representatives from Israel. A. Hysterochelifer cyprius (Cheliferidae). B. Olpium pallipes balcanicum (Olpidae). C. Lasiochernes turcicus (Chernetidae). D. Neobisium validum (Neobisiidae). E. Ephippiochthonius sp. (Chthoniidae). F. Nannowithius wahrmani (Withiidae) phoretic on Birulatus israelensis. Photos A, B, D, E, F by S. Aharon, C by J. A. Ballesteros.

3.1.3. A dichotomous key to the pseudoscorpion families and genera of Israel (applying to adults) (in parentheses no. of genera and no. of species in the family).

1. a. Legs with different number of tarsal segments: legs I and II with one tarsal segment and legs III and IV with two tarsal segments; pedipalpal chelal fingers slender and without venom apparatus; two trichobothria (ib and isb) medially on dorsal face of pedipalpal chela hand; coxal spines present (Figure 7A) . . . . . . . . . . . . . . . . . . . . . . . . . suborder Heterosphyronida . . . . . . . . . . . . . . . . . . . . . superfamily Chthonioidea . . . . . . . . . family Chthoniidae (3, 7) 2

1. b. All legs with same number of tarsal segments; at least one pedipalpal chelal finger with venom apparatus (Figure 7B); trichobothria ib and isb usually located on the lateral face of the fixed chelal finger (occasionally ib is on the dorsal surface); coxal spines absent (Figure 7C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .suborder Iocheirata 4

2. a. Coxae III without coxal spines; only coxae II each with 11 fine elongated coxal spines, middle spines longest; apex of pedipal pal coxa with three setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ayyalonia

2. b. Coxae II and III with coxal spines; apex of pedipal pal coxa with two setae . . . . . . . . . . Chthonius-related genera 3

3. a. Teeth of pedipalpal chelal fingers pointed, aligned, erect and spaced; pedipal pal hand with dorsal step-like outline (dorsodistal saddle-shaped constriction) (Figure 7D) between the groups of trichobothria ib-isb ang eb-esb-ist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ephippiochthonius

3. b. Teeth of pedipalpal chelal fingers blunt, dense and distinctly reclined backwards; pedipal pal hand with evenly rounded weakly curved dorsal outline (Figure 7E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chthonius

4. a. Rectangular carapace (anterior and posterior margins similar in length); robust chelicerae, their bases almost as wide as posterior side of carapace; all legs with two tarsal segments; two pairs of eyes (epigean species) or none (caves); setae present on the apical margin of pedipalp coxa (Figure 7F,G) . . . . . . . . . . . . . . . .infraorder Hemictenata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . superfamily Neobisioidea 5

4. b. Carapace not rectangular (anterior margin shorter than posterior margin); width of chelicerae bases smaller than posterior margin of carapace (less than half its length) (Figure 7H); the legs have either all one tarsal segment or all two tarsal segments; either no eyes or one or two pairs of eyes or eyespots; setae on the apical margin of pedipalp coxa may be present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .infraorder Panctenata 6

5. a. Apex of pedipalpal coxa rounded and bears three or more setae (Figure 7F) . . . . . . . . . . . family Neobisiidae (1, 3)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobisium

5. b. Apex of pedipalpal coxa not rounded and bears two long subequal setae (Figure 7G) . . . . family Syarinidae (1, 1)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hadoblothrus

6. a. All legs with two tarsal segments (Figure 7I); two pairs of eyes with lenses; usually elongated carapace (longer than wide); pedipalpal femur with or without trichobothria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7

6. b. All legs with one tarsal segment (Figure 7J); one pair of eyes or eye-spots, or eyes absent; carapace sub-triangular; pedipalpal femur without trichobothria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

Figure 7. A. Ephippiochthonius tetrachelatus with coxal spines present. B. Geogarypus pulcher with venom apparatus on both pedipalpal chelal fingers. C. Neobisium validum with no coxal spines. D. Ephippiochthonius sp. with dorsal step-like outline of pedipalpal hand. E. Cthonius sp. with evenly rounded weakly curved dorsal outline of pedipalpal hand. F. Neobisium validum with five setae on rounded pedipalp coxa apex. G. Hadoblothrus sp. pedipalp coxal apex not rounded and bears two long subequal setae. H. Garypinus sp. with the width of the chelicerae bases smaller than the width of the posterior margin of carapace. I. Garypinus sp. with two tarsal segments in all legs. J. Hysterochelifer tuberculatus with one tarsal segment in all legs.

Figure 7. A. Ephippiochthonius tetrachelatus with coxal spines present. B. Geogarypus pulcher with venom apparatus on both pedipalpal chelal fingers. C. Neobisium validum with no coxal spines. D. Ephippiochthonius sp. with dorsal step-like outline of pedipalpal hand. E. Cthonius sp. with evenly rounded weakly curved dorsal outline of pedipalpal hand. F. Neobisium validum with five setae on rounded pedipalp coxa apex. G. Hadoblothrus sp. pedipalp coxal apex not rounded and bears two long subequal setae. H. Garypinus sp. with the width of the chelicerae bases smaller than the width of the posterior margin of carapace. I. Garypinus sp. with two tarsal segments in all legs. J. Hysterochelifer tuberculatus with one tarsal segment in all legs.

7. a. Arolia bifurcate (Figure 8A); at least a few tergites and sternites are divided. . . . . . . . . .superfamily Garypinoidea

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . family Garypinidae (1, 2)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Garypinus

7. b. Arolia simple (Figure 8B) (not bifurcate); tergites and sternites can (in certain families) be all undivided . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . superfamily Garypoidea . . . . . . . . . . . . . . . . . .8

8. a. Venom apparatus present only in the fixed chelal finger; fixed chelal finger with 11 trichobothria, of which two are on the dorsal surface of the hand; specialised joint present between coxae II and III . . . . . . family Menthidae (1, 1)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramenthus shulovi

8. b. Venom apparatus present in both chelal fingers; fixed chelal finger with 8 or (rarely) 7 trichobothria, including 3 on internal face; dorsal surface of the hand without trichobothria; articulation furrow between coxae II and III absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. a. All tergites and sternites undivided (Figure 8C,D); abdomen oblong, not much broader than the carapace; the carapace sub-rectangular and slightly tapering orally; eyes positioned near anterior margin of carapace, not sitting on mounds; arolia longer than claws; vestitural bristles of pedipalps rather long, erect, and thinly pointed; pleural membrane of abdomen with smooth and even longitudinal striation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

9. b. At least a few tergites and sternites divided (Figure 8E); abdomen broadly oval, distinctly broader than the carapace; carapace triangular and strongly tapering orally; eyes protruding, sitting on mounds, removed from anterior margin of the carapace; arolia can be longer or shorter than the claws; vestitural bristles of pedipalps very short, fine, bent, and abruptly pointed; pleural membrane of abdomen granulate or with short wavy undulating striation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15

10. a. Palpal fingers distinctly bent; palpal femur with or without two fairly long dorsal tactile bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . family Olpiidae (4, 10) . . . . . . . .11

10. b. Palpal fingers straight; palpal femur with a single fairly long dorsosubbasal tactile bristle . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .family Hesperolpiidae (2, 2) . . . . . . . 14

11. a. Carapace without submedian transverse furrow, longer than broad maximum 1.5 times . . . . . . . . . . . . . . . . . . . .12

11. b. Carapace with a submedian transverse furrow; either just slightly or 1.7 to 1.8 times longer than broad. . . . . . . 13

12. a. Carapace and pedipalps dark brown; palpal femur 0.6 to 0.7 mm long; fingers about as long as the hand with pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Olpium

12. b. Carapace and pedipalps light yellowish-brown; palpal femur 0.9 to 1 mm long; fingers much longer than hand with pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halominniza

13. a. Body length about 2 to 3 mm; carapace 1.7 to 1.8 times longer than broad; femur, tibia and chela of pedipalps dark brown, with pedicel and tips reddish, rarely entirely reddish; desert species with worm-shaped abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minniza

13. b. Body length 1.5 to 1.7 mm; carapace but slightly longer than broad; femur and tibia of pedipalps light brownish-yellow, chela blackish-olivebrown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Calocheiridius

14. a. Body length 2.3 to 2.8 mm; eyes about half their diameter distant from one another; carapace with a weak transverse furrow; palpal hand very broad, abruptly constricted medially and laterally towards the base of the fingers; fingers at least as long as the femur; trichobothria est situated near the middle of fixed finger; three trichobothria ist, it and et form a group near tip of finger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calocheirus

14. b. Body length 1.2 to 1.9 mm; eyes touching, the anterior eyes slightly excavated behind, the posterior eyes conical and flattened; carapace without a transverse furrow; palpal hand not very broad, medially obliquely narrowed towards the base of fingers; fingers shorter than femur; trichobothria est situated rather basally, proximal of the middle, near trichobothria isb, close to the group formed of eb-esb-ib-isb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cardiolpium

15. a. Carapace triangular and medially concave (Figure 8E); maxilla of palpal coxae with developed shoulder; coxal area not expanding posteriorly: coxa IV approximately same width as coxa I (broad and short); body 2 to 2.5 mm long; arolia longer than the claws; tergites usually with dark spots; occur in leaf litter and under stones or bark in the Mediterranean shrubland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . family Geogarypidae (1, 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Geogarypus

15. b. Carapace triangular but not medially concave (Figure 8F); maxillar shoulder of palpal coxae not developed; coxal area expanding posteriorly: i.e. coxa IV distinctly longer and narrower than coxa I; body 5 to 6 mm long; arolia shorter than claws; tergites usually with dark bands; occur only in littoral or supralittoral zones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . family Garypidae (1, 2)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Garypus

16. a. Small body size (< 2 mm); carapace triangular and coarsely granulate with one medial deep transverse furrow (Figure 8G); one pair of small eyes, if not absent, distinctly removed from anterior margin; femur and patella of all legs fused, with suture between them hardly visible; coxa IV much wider than coxa I; pedipalps with reduced number of trichobothria: at most seven on fixed chelal finger and one or two on moveable finger. . .superfamily Cheiridioidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .family Cheiridiidae (1, 1)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocheiridium

16. b. Body size larger than 1.5 mm; eyes (if not absent) positioned near the anterior margin of the carapace (Figure 8H); femur and patella of legs not fused; coxa IV not wider than coxa I; pedipalps usually with eight trichobothria on fixed chelal finger and four on moveable finger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . superfamily Cheliferoidea. . . . 17

17. a. Venom apparatus present in both chelal fingers; one pair of eyes or eye spots; males with or without coxal sacs or ram’s horn organs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18

17. b. Venom apparatus absent from one chelal finger; no eyes or one pair of eyes or eye spots; males without coxal sacs or ram’s horn organs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22

18. a. Articulation of femur and patella of leg I and II narrow and transverse, therefore these joints scarcely movable; male (and occasionally females) abdominal posterior sternites with discrete patches of glandular sensory setae; hind coxae of male without coxal sacks and ram’s horn organs absent; very small myrmecophilous specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .family Withiidae (1, 1)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nannowithius wahrmani

18. b. Articulation of femur and patella of leg I and II well developed (Figure 8I), great and oblique and therefore well movable; male abdominal sternites without patches of sensory setae; hind coxae of male excavate caudally, male genitalia with coxal sacks and ram’s horn shaped organs (Figure 8I); not myrmecophilous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . .family Cheliferidae (4, 18) . . . . . . . . . . . . . . . .19

19. a. Carapace with dense granulation and greater bristle-bearing granules; abdominal tergites of male with lateral carinae and elongate hind-angles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19. b. Carapace with dense granulation without greater granules; abdominal tergites of male without lateral carinae, their hind-angles not elongate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

20. a. Pedipalpal femur with coarse bristle-bearing medial granules; subbasal bristle of chelicera present . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hysterochelifer

20. b. Pedipalpal femur without coarse medial granules; no subbasal bristle of chelicera . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Chelifer

21. a. At least submedian transverse furrow of the carapace deeply incised; pedipalps slender. . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylochelifer

21. b. Both transverse furrows of carapace shallow; pedipalps clumsy and strong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Rhacochelifer

22. a. No eyes or eye spots (Figure 8J); venom apparatus developed only in movable chelal finger; carapace with grooves; robust (Figure 8J); abdomen broadly oval, distinctly broader than the carapace . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . family Chernetidae (6, 9) . . . . . . . . . . .23

22. b. One pair of eyes or eye spots; venom apparatus developed only in fixed chelal finger; longitudinal grooves may be present; slender and elongate abdomen, not broader than the carapace . . . . . . . . . . . . . . . . . . family Atemnidae (1, 2)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Atemnus

23. a. Body and pedipalp setae long, thin and pointed; carapace almost smooth; subbasal transverse furrow on carapace indistinct; tibia and hand of pedipalps with fairly long lateral pseudotactile setae; often phoretic on flies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Lamprochernes

23. b. Body and pedipalp setae short, stout, dentate or clavate; carapace granulate; tibia and hand of pedipalps without lateral pseudotactile setae; not phoretic on flies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

24. a. Tarsus of leg IV with long erect tactile seta near the middle, distinctly longer than width of tarsus . . . . . . . . . . . 25

24. b. Tarsus of leg IV without erect tactile seta near the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

25. a. Carapace rather coarsely granulated, with two distinct transverse furrows, of which the posterior is flatter than the anterior; body and pedipalp setae always slightly but clearly clavate, relatively long; male pedipalp without dense long setation; palpal femur at most 0.5 mm long; small 1.5-2 mm; in ground litter and detritus, in moist places . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pselaphochernes

25. b. Carapace with dense and fine granulation with posterior transverse furrow situated scarcely nearer to hind border than to the anterior furrow; body setae dentate, but not clavate; palpal femur longer than 0.7 mm; body length 2-4 mm; in nests of small mammals and in caves with bats . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . .26

26. a. Tactile bristle near the middle of the hind tarsus long and simple; body setae rather long serrated to slightly truncate not clavate, those of the pedipalps only serrated; setae on femur and tibia of the pedipalps of the male very long and dense like a mane; fairly large and robust: body length 4 mm, palpal femur at least 1 mm long; in nests of Apodemus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lasiochernes

26. b. Tactile bristle near the middle of the hind tarsus dentate and scarcely twice as long as the other bristles and about one third as long as width of tarsus; body setae very short; medial bristles of the palpal femur of the male but scarcely more dense than in the female; body length 2-3 mm, femur length 0.69-0.74 mm; in nests of the rodent Nannospalax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nudochernes

27. a. Carapace quite coarsely granulated; body and pedipalp setae serrated and strongly culled usually quite short; slightly clavate; pedialps are moderately slender, granulated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allochernes

27. b. Carapace moderately coarsely granulated, sometimes partially reticulated; body and pedipalp setae dentate and more or less truncate; pedialps robust, granulated; pedipalp coxae usually not granulated . . . . . . . . . . . . . . . . . . Chernes

Figure 8. A. Bifurcate arolia. B. Simple arolia. C. Minniza babylonica with all tergites undivided. D. Minniza babylonica with all sternites undivided. E. Geogarypus sp. with five tergites divided. F. Geogarypus sp. with carapace triangular and medially concave. G. Garypus beauvoisi with carapace triangular but not medially concave. H. Cheiridiidae sp. with one medial deep transverse furrow in the coarsely granulate and triangular carapace. I. Hysterochelifer tuberculatus with eyes positioned near the anterior margin of the carapace. J. Hysterochelifer tuberculatus with articulation of femur and patella of leg I and II well developed and male genitalia with ram’s horn shaped organs. K. Chernetidae sp. robust with two grooves in the carapace and without eyes or eye spots.

Figure 8. A. Bifurcate arolia. B. Simple arolia. C. Minniza babylonica with all tergites undivided. D. Minniza babylonica with all sternites undivided. E. Geogarypus sp. with five tergites divided. F. Geogarypus sp. with carapace triangular and medially concave. G. Garypus beauvoisi with carapace triangular but not medially concave. H. Cheiridiidae sp. with one medial deep transverse furrow in the coarsely granulate and triangular carapace. I. Hysterochelifer tuberculatus with eyes positioned near the anterior margin of the carapace. J. Hysterochelifer tuberculatus with articulation of femur and patella of leg I and II well developed and male genitalia with ram’s horn shaped organs. K. Chernetidae sp. robust with two grooves in the carapace and without eyes or eye spots.

4. Discussion

• Distribution and Zoogeography

• Endemism

• Seasonality

• Habitats and Collection methods

References

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