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a) Species richness of earthworms in the Levant
We define the Levant bioprovince as a stretch of land about 150 km wide, wedged between the Mediterranean sea and the Syrio–Arabian deserts, stretching from the Taurus Mountains in the north to the Isthmus of Suez in the south (Pavlíček et al., 2006).
The earthworm fauna of the Levant had long been neglected and our knowledge was limited to the sporadic collections done by European scientists. The work done by Rosa (1893) was certainly the most important one.
A regular earthworm survey started in the Levant only in the mid-1990s. The survey has been conducted by Dr Tomaš Pavlíček (Institute of Evolution, University of Haifa, Israel) and Professor Csaba Csuzdi, Hungarian Natural History Museum, Budapest, Hungary, and Department of Zoology, Eszterházy Károly University, Eger, Hungary. The research on earthworms had been supported by Prof. E. Nevo, the former director of the Institute of Evolution, University of Haifa.
Altogether, 50 species of earthworms were recorded in the Levant bioprovince (Table 2), including the 12 species described as new to science (Table 1).
Please click on the above links to access to the data if the different regions.
As a result of our survey in the Levant, the following, twelve species new to science have been recorded and two species were re-established from synonymy (Table 1).
| Species | Recorded in |
| Dendrobaena alexandrii Szederjesi et al. 2013 | Szederjesi et al., 2013b |
| Dendrobaena mahunkai Csuzdi et al., 2007 | Csuzdi et al., 2007 |
| Dendrobaena negevis Csuzdi and Pavlíček, 1999 | Csuzdi and Pavlíček, 1999 |
| Dendrobaena nevoi Csuzdi and Pavlíček, 1999 | Csuzdi and Pavlíček, 1999 |
| Dendrobaena omodeoi Csuzdi et al., 2007 | Csuzdi et al., 2007 |
| Dendrobaena orientalis karak Csuzdi and Pavlíček, 2005 | Csuzdi and Pavlíček, 2005b |
| Dendrobaena rothschildae Csuzdi and Pavlíček, 1999 | Csuzdi et al., 1999 |
| Dendrobaena transjordanica Szederjesi et al., 2013 | Szederjesi et al., 2013b |
| Healyella jordanis Csuzdi and Pavlíček, 1999 | Csuzdi and Pavlíček, 1999 |
| Perelia galileana Csuzdi and Pavlíček, 2005 | Csuzdi et Pavlíček, 2005a |
| Perelia makrisi Szederjesi et al., 2016 | Szederjesi et al., 2016a |
| Perelia shamsi Csuzdi and Pavlíček, 2005 | Csuzdi and Pavlíček, 2005a |
| Species re-established from synonymy | |
| Dendrobaena succinta (Rosa, 1905) | Szederjesi et al., 2019 |
| Perelia phoeba (Cognetti, 1913) | Szederjesi et al., 2016b |
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| Allolobophora nematogena | |||
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| Octodrilus complanatus | Dendrobaena veneta | Dendrobaena semitica | Dendrobaena pentheri |
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b) Characteristic features of the earthworm fauna in the Levant
The above mentioned survey allows to make some conclusions about the earthworm fauna in the Levant.
This fauna (except for some introduced species) is exclusively Palearctic and does not show a transitory character between the Ethiopic and Eurasian regions. Since ancient genera are missing (e.g. Anatolian genus Spermophorodrilus), it seems that this fauna originated later than faunas of some nearby Palearctic zoogeographic regions such as Anatolia, Caucasus and the Balkans. Autochthonous in the Levant are the Palearctic family Lumbricidae and possibly the family Criodrilidae (represented in the Levant by one species, Criodrilus lacuum, Hoffmeister, 1845). Families Ocnerodrilidae, Acanthodrilidae and Megascolecidae are represented only by peregrine species.
On the species level, about 69% of all recorded species are autochthonous (Table 2). As a result of the adaptive radiation that took place inside the genera Dendrobaena, Healyella and Perelia, as well as due to partial isolation of the province by mountains and deserts about 50% of the autochthonous earthworm species are endemic in the Levant.
| Species | Recorded in | Expected origin |
| Family Lumbricidae | ||
| Allolobophora chlorotica (Savigny, 1826) | Rosa, 1893; Csuzdi and Pavlíček et al., 2005a | P |
| Aporrectodea caliginosa (Savigny, 1826), including the form trapezoides | Michaelsen, 1892; Rosa, 1893; Zicsi, 1985; Michalis, 1993; Pavlíček et al., 1997; Csuzdi and Pavlíček, 2005a, b; Pavlíček and Csuzdi, 2005, 2006, 2012; Csuzdi et al., 2007; Szederjesi et al., 2013a, b; 2016b | P |
| Aporrectodea georgii (Michaelsen, 1890) | Rosa, 1893; Michaelsen, 1926 | A |
| Aporrectodea jassyensis (Michaelsen, 1891) | Rosa, 1893; Michalis, 1993; Csuzdi and Pavlíček, 2005a; Csuzdi et al., 2007; Pavlíček and Csuzdi, 2012; Szederjesi et al., 2013a; 2016 | A |
| Aporrectodea rosea (Savigny, 1826) | Rosa, 1893; Zicsi 1985; Michalis, 1993; Pavlíček et al., 1997; Csuzdi and Pavlíček, 2005a, b; Pavlíček and Csuzdi, 2005, 2006, 2012; Csuzdi et al., 2007; Szederjesi et al., 2013a, b | P |
| Bimastos parvus (Eisen, 1874) | Csuzdi and Pavlíček, 2005a; Pavlíček and Csuzdi, 2005; Szederjesi et al., 2013a | P |
| Dendrobaena alexandrii Szederjesi et al., 2013 | Szederjesi et al., 2013b | A |
| Dendrobaena byblica byblica (Rosa, 1893) | Rosa, 1893; Zicsi, 1985; Csuzdi and Pavlíček, 2002, 2005b; Pavlíček and Csuzdi, 2006; Csuzdi et al., 2007; Szederjesi et al., 2013a, b | A |
| Dendrobaena fedtschenkoi (Michaelsen, 1900) | Michaelsen, 1926 | A |
| Dendrobaena hauseri Zicsi, 1973 | Zicsi, 1985; Szederjesi et al., 2013a | A |
| Dendrobaena kervillei (Michaelsen, 1910) | Rosa, 1893; Michaelsen, 1910; Zicsi, 1985; Csuzdi and Pavlíček, 1999, 2002; Szederjesi et al., 2013a | A |
| Dendrobaena mahunkai Csuzdi et al., 2007 | Csuzdi et al., 2007 | A |
| Dendrobaena negevis Csuzdi and Pavlíček, 1999 | Csuzdi and Pavlíček, 1999, 2002; Pavlíček and Csuzdi, 2005; Szederjesi et al., 2013a, b | A |
| Dendrobaena nevoi Csuzdi and Pavlíček, 1999 | Csuzdi and Pavlíček, 1999; Szederjesi et al., 2013a | A |
| Dendrobaena omodeoi Csuzdi et al., 2007 | Csuzdi et al., 2007 | A |
| Dendrobaena orientalis Cernosvitov, 1940 | Cernosvitov, 1940; Zisci, 1985; Pavlíček et al., 1996; Csuzdi and Pavlíček, 1999, 2002; Csuzdi et al., 2007; Szederjesi et al., 2013a | A |
| Dendrobaena orientalis karak Csuzdi and Pavlíček, 2005 | Csuzdi and Pavlíček, 2005b | A |
| Dendrobaena pantaleonis (Chinaglia, 1913) | Michalis, 1993 | A |
| Dendrobaena pentheri (Rosa, 1905) | Pavlíček and Csuzdi, 2006; Pavlíček et al., 2006; Csuzdi et al., 2007; Szederjesi et al., 2016 | A |
| Dendrobaena rothschildae Csuzdi and Pavlíček, 1999 | Csuzdi et al., 1999 | A |
| Dendrobaena samarigera (Rosa, 1893) | Rosa, 1893; Stephenson, 1922; Pavlíček et al., 1996; Csuzdi and Pavlíček, 1999; Szederjesi et al. , 2013a | A |
| Dendrobaena semitica (Rosa, 1893) | Rosa, 1893; Michaelsen, 1910; Zicsi, 1985; Pavlíček et al., 1996; Csuzdi and Pavlíček, 1999, 2002, 2005b; Pavlíček and Csuzdi, 2005, 2006, 2012; Csuzdi et al., 2007; Szederjesi et al., 2013a, b; 2016 | A |
| Dendrobaena succinta (Rosa, 1905) | Pavlíček and Csuzdi, 2006; Szederjesi et al. 2019 | A |
| Dendrobaena transjordanica Szederjesi et al., 2013 | Szederjesi et al. 2013b | A |
| Dendrobaena veneta veneta (Rosa, 1886) | Rosa, 1893; Zicsi, 1985; Michalis, 1993; Pavlíček et al., 1996; Csuzdi and Pavlíček, 1999, 2002, 2005b; Pavlíček and Csuzdi, 2005, 2006, 2012; Csuzdi et al., 2007; Szederjesi et al., 2013a, b; 2016b | A |
| Dendrodrilus rubidus subrubicundus (Eisen, 1874) | Pavlíček and Csuzdi, 2012; Szederjesi et al., 2016b | P |
| Eisenia fetida (Savigny, 1826) | Csuzdi and Pavlíček, 2005a, b; Pavlíček and Csuzdi, 2012; Szederjesi et al., 2016b | P |
| Eiseniella neapolitana (Örley, 1885) | Rosa, 1893; Csuzdi and Pavlíček, 2005a, b; Pavlíček and Csuzdi, 2006; Csuzdi et al., 2007; Szederjesi et al., 2013a, b, 2016b | A |
| Eiseniella tetraedra (Savigny, 1826) | Rosa, 1893; Michaelsen, 1926; Cernosvitov, 1938; Pavlíček et al., 1997; Csuzdi and Pavlíček, 2005a, b; Pavlíček and Csuzdi, 2006; Csuzdi et al., 2007; Szederjesi et al., 2013a, b, 2016b | P |
| Healyella jordanis Csuzdi and Pavlíček, 1999 | Csuzdi and Pavlíček, 1999, 2002; Szederjesi et al., 2013a | A |
| Healyella syriaca (Rosa,1893) | Zicsi, 1985; Pavlíček et al., 1996; Csuzdi and Pavlíček, 1999, 2002; Csuzdi et al., 2007; Szederjesi et al., 2013a, b | A |
| Helodrilus patriarchalis (Rosa, 1893) | Rosa, 1893, Csuzdi and Pavlíček, 2005a, b; Pavlíček and Csuzdi, 2006; Csuzdi et al., 2007; Szederjesi et al., 2013a, b | A |
| Murcheiona minuscula (Rosa, 1905) | Michalis, 1893; Csuzdi and Pavlíček, 2002; Pavlíček and Csuzdi, 2006; Csuzdi et al., 2007; Szederjesi et al., 2013a | A |
| Octodrilus complanatus (Antoine Dugès, 1828) | Rosa, 1893; Pavlíček and Csuzdi, 2006; Csuzdi and Pavlíček, 2005a | A | Octodrilus transpadanus (Rosa, 1884) | Csuzdi and Pavlíček, 2005b; Csuzdi et al., 2007; Szederjesi et al., 2013b | A |
| Octolasion cyaneum (Savigny, 1826) | Pavlíček and Csuzdi, 2006 | P |
| Perelia aharonii (Stephenson, 1922) | Stephenson, 1922; Csuzdi and Pavlíček, 2005a | A |
| Perelia galileana Csuzdi and Pavlíček, 2005a | Csuzdi and Pavlíček, 2005 | A |
| Perelia hatayica Csuzdi et al. 2007 | Csuzdi et al., 2007 | A |
| Perelia makrisi Szederjesi et al., 2016 | Szederjesi et al., 2016b | A |
| Perelia nematogena (Rosa, 1903) | Csuzdi and Pavlíček, 2005a; Pavlíček and Csuzdi, 2006 | A |
| Perelia phoeba (Cognetti, 1913) | Szederjesi et al., 2016b | A |
| Perelia shamsi Csuzdi and Pavlíček, 2005 | Csuzdi and Pavlíček, 2005a; Szederjesi et al., 2013a | A |
| Proctodrilus tuberculatus (Cernosvitov, 1935) | Pavlíček and Csuzdi, 2006 | P |
| Family Acanthodrilidae | ||
| Dichogaster bolaui (Michaelsen, 1891) | Csuzdi et al., 2008 | P |
| Microscolex dubius (Fletcher, 1887) | Pavlíček et al., 1996; Csuzdi and Pavlíček, 2005b; Pavlíček and Csuzdi, 2005, 2012 | P |
| Microscolex phosphoreus (Antoine Dugès, 1837) | Cernosvitov, 1940; Michalis, 1993; Csuzdi et al. 2007 | P |
| Family Criodrilidae | ||
| Criodrilus lacuum Hoffmeister, 1845 | Rosa, 1893; Michaelsen, 1926; Cernosvitov, 1938; Csuzdi and Pavlíček, 2005a; Csuzdi et al. 2007 | A |
| Family Megascolecidae | ||
| Metaphire californica (Kinberg, 1867) | Pavlíček and Csuzdi, 2005 | P |
| Family Ocnerodrilidae | ||
| Ocnerodrilus occidentalis Eisen, 1878 | Cernosvitov, 1940 | P |
| Total: 50 species | 36 A + 14 P | |
References
Cernosvitov, L., 1938. Report of the Percy Sladen expedition to Lake Hulech, A contribution to the study of the fresh waters of Palestine. Oligochaeta. Ann. Mag. Nat. Hist. 2: 535-550.
Cernosvitov, L., 1940. On some Oligochaeta from Palestine. Ann. Mag. Nat. Hist. 11: 438–447.
Csuzdi, Cs. and Pavlíček, T., 1999. Earthworms from Israel. I. Genera Dendrobaena Eisen, 1874 and Bimastos Moore, 1891 (Oligochaeta: Lumbricidae). Isr. J. Zool. 45: 467-486. Abstract
Csuzdi, Cs. and Pavlíček, T. , 2005. Earthworms from Israel. II. Remarks on the genus Perelia Easton, 1983 with description of a new genus and two new species. Acta Zool. Acad. Sci. Hungaricae 51: 75–96. Abstract
Csuzdi, Cs. and Pavlíček, T. , 2002. Murchieona minuscula (Rosa, 1906), a new recorded earthworm from Israel, and distribution of genera Dendrobaena and Bimastos in Israel (Oligochaeta, Lumbricidae). Zool. Middle East 25: 105–114. Abstract
Csuzdi, C. & Pavlíček, T., 2005. The earthworms (Oligochaeta) of Jordan. Zool. Middle East 34: 71–78. Abstract
Csuzdi, Cs., Pavlíček, T. and Mısırlıoğlu, M., 2007. Earthworms (Oligochaeta: Lumbricidae, Criodrilidae, Acanthodrilidae) of the Hatay Province, Turkey, with description of three new lumbricids. Acta zool. hung. 53: 347-361. Abstract
Csuzdi, Cs., Pavlíček, T. and Nevo, E., 1999. A new earthworm species, Dendrobaena rothschildae sp. n. from Israel, and comments on the distribution of Dendrobaena species in the Levant (Oligochaeta: Lumbricidae). Opusc. Zool. Budapest 31: 25–32. Abstract
Csuzdi, C., Pavlíček, T. & Nevo, E., 2008. Is Dichogaster bolaui (Michaelsen, 1891) the first domicole earthworm species? Eur. J. Soil Biol. 44: 198–201. Abstract
Michaelsen, W., 1892. Terricolen der Berliner Zoologischen Sammlung II, Arch. f. Naturgesch.. 1:, 209-261.
Michaelsen, W., 1910. Oligochäten von verschiedenen Gebieten. Mitteilungen aus dem Naturhistorischen Museum Hamburg. 27: 47-169.
Michaelsen, W., 1926. Note sur les Oligochètes rapportés par M. Henri Gadeau de Kerville de son voyage zoologique en Syrie. (Avril-Juin 1908). Voyag. Zool. Gadeau Kerville, Syrie 1: 351–354.
Michalis, K. A., 1993. Contribution to the study of the Oligochaetofauna of Cyprus. Acta zoologica cracoviensia, 36: 23–28.
Pavlíček, T., & Csuzdi, Cs., 2012. Chapter 23 : Clitellata : Oligochaeta. In The Wildlife of Cyprus (D.J. Sparrow and E. John, eds.) pp 587-–599, Terra Cypria. Abstract
Pavlíček, T. and Csuzdi, Cs., 2005. Earthworm fauna of Jordan — A review. In: Advances in Earthworm Taxonomy II (Annelida: Oligochaeta). Proceedings of the 2nd International Oligochaeta Taxonomy Meeting dedicated to Victor Pop. 4–8 September 2005, pp. 183–188.
Pavlíček, T., and Csuzdi, Cs., 2006. Species richness and zoogeographic affinities of earthworms in Cyprus. European Journal of Soil Biology, 42, S111–S116. Abstract
Pavlíček, T., Csuzdi, Cs. and Nevo, E., 2006: Biodiversity of earthworms in the Levant. Isr. J. Ecol. Evol. 52: 461-466. Abstract
Pavlíček, T., Csuzdi, Cs. and Nevo, E., 1997. The first recorded earthworms from the Negev and Sinai Deserts. Isr. J. Zool. 43: 1–3. Abstract
Pavlíček, T., Csuzdi, Cs., Smooha, G., Beiles, A. and Nevo, E., 1996. Biodiversity and microhabitat distribution of earthworms at" Evolution Canyon," a Mediterranean microsite, Mount Carmel, Israel. Isr. J. Zool. 42: 449–454. Abstract
Rosa, D., 1893. Viaggi del Dr. E. Festa in Palestina nel Libano e regioni vicine. II. Lumbricidi. Boll. del Musei di Zool. ed Anat. Comp. della R. Univ. di Torino 8: 1–14.
Stephenson, J., 1922. VIII.— On some earthworms from India and Palestine belonging to the British Museum. Ann. Mag. Nat. Hist. 9: 129–136.
Szederjesi, T., Pavlíček, T., and Csuzdi, Cs., 2016. Description of the first endemic earthworm species from Cyprus (Oligochaeta: Lumbricidae). Zoology in the Middle East, 62: 158–163. Abstract
Szederjesi T., Pavlíček, T., Csuzdi, Cs., 2013a. New data to the earthworm fauna of Israel (Oligochaeta, Lumbricidae). Opuscula zoologica Budapest 44: 201-209. Abstract
Szederjesi, T., Pavlíček, T. and Csuzdi, Cs., 2013b. New earthworm species from Jordan (Oligochaeta, Lumbricidae). Acta Zool. Acad. Sci. Hungaricae 59: 391–400. Abstract
Szederjesi, T., Pavlíček, T., Márton, O., Krízsik, V., and Csuzdi, Cs., 2019. Integrative taxonomic revision of Dendrobaena veneta (Rosa, 1886) sensu lato with description of a new species and resurrection of Dendrobaena succinta (Rosa, 1905) (Megadrili: Lumbricidae). Journal of Natural History 53: 301–314. Abstract
Zicsi, A. 1985. Regenwürmer (Oligochaeta: Lumbricidae) aus Israel und den benachbarten Ländern. Rev. Suisse Zool. 92: 323–331.
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