Research Article ISSN 2336-9744 (online) | ISSN 2337-0173 (print) The journal is available on line at www.biotaxa.org/em Arachnogeographical comparison between West Palearctic and Afrotropical Areas PETAR BERON National Museum of Natural History, Tsar Osvoboditel Bld 1, 1000 Sofia, Bulgaria e-mail: [email protected] Received 5 September 2016 │ Accepted 25 November 2016 │ Published online 28 November 2016. Abstract Comparison is made between the Arachnofaunas of Western Palearctic (Europe, SW Asia and North Africa to 20oN) and Afrotropical (from 20oN to the line Zambezi – Kunene) areas. This border (actually transition zone) divides two Kingdoms – Holarctic and Paleotropic. What concerns Arachnida so far such comparison has never been made (for all the orders). Outlined are the endemic taxa and it is clear that the difference is due mostly to climatic difference (for the warm preferring groups like Schizomida, Amblypygi, Solifugae, Scorpiones, Ricinulei). Some groups in tropical Africa are relict (suborder Paleoamblypygi). In the Western Palearctic such groups (endemic and relics) are the scorpions of the genera Belisarius and Akrav (with Southamerican affinities). In our time the vast deserts like Sahara and the Arabian desert are a very important barrier for many groups. Key words: Palearctic, Afrotropical, Arachnogeography. Introduction The northern part of Africa is included in the Palearctic Region and the boundary (usualy at 20oN) is separating two Kingdoms – Holarctic and Paleotropical. It is common knowledge that a biogeographical boundary is not a line, but rather a more or less wide transitional zone. We have also to take into account the historical changes during the last several thousand years. Part of these changes is due to climatic factors, other – to human activities (also causing changes in the climate), like deforestation, diverting of rivers, etc. The traditional zoogeographic regions are delimitated mainly on the distribution of terrestrial vertebrates. The difference between the regions and kingdoms are usually on much higher taxonomical level when we consider vertebrates, than with the Arachnida. Differences between Afrotropical and Neotropical regions, or between Holarctic and Paleotropical are between orders or suborders and very much between families. Historically it was not as it is today. In green Sahara and Mediterranean Africa lived elephants, crocodiles, giraffes, in Atlas mountains there were until recently ostriches, lions and bears. They were exterminated by Man. We do not speak about geological periods when these animals (plus Tubulidentata, Marsupialia, tapirs and many others) lived in Europe. The present day biogeographical subdivision should consider the picture of the last several thousand years. We don‘t know what was the distribution of many Arachnids over the green Sahara. It was not influenced by the rare human population during the millenia, as it certainly was in Europe and North America with the disappearing of forests and their replacement with anthropogenic landscape. Important were also new crops like maiz and potatoes, chemization and other kinds of human intruding into the ecosystems. Ecol. Mont., 7, 2016, 464-506 BERON If we follow the classical subdivision (Africa south of Sahara and the southernmost Arabia form the Afrotropical Region), north of this ―line‖ is the Palearctic (different subregions). This ―line‖ (actually transitional zone) is separing two kingdoms, so differences should be substantial. Let us check how the known distribution of Arachnida fits into the classical scheme. Geography and General Zoogeography of Western Palearctic Here Western Palearctic is understood as embracing Europe, Africa North of 20oN and SW Asia (from Bosphorus to Sind). Included is also Siberia west of Yenisey (Johanssen‘s line). Europe is divided into Eurosiberian and Mediterranean Subregions. North Africa near Mediterranean Sea has also Mediterranean fauna, to the South it changes into Saharo-Sindian desert fauna. Neither Europe, nor North Africa and Sahara have always been as they are today. The glaciations changed the older fauna of Europe, than came the destructive human activities. The forests have been very much reduced, and this brought a total desappearence of many invertebrate communities. Such was also the case of the deforestation of circummediterranean territory. South of the barren Atlas mountains once there was a green Sahara, gradually desiccated (Bodenheimer, 1935, Darlington, 1957, de Lattin, 1967, Geptner, 1936, Udvardy, 1975). Analyzing the Zoogeography of the Levant, Por (1975) reached to the conclusion that the Levant province is a peculiar and complex ―subtraction-transition zone‖ (following the expression of Darlington, 1957). The Levant is ―a stretch of land about 150 km wide, wedged in between the [Mediterranean] sea and the Syrio-Arabian desert, stretching from the mouth of the River Orontes and the Amanus and Taurus mountain ranges in the north, to the Isthmus of Suez in the south‖(POR, 1975). This ―subtraction-transition zone‖ between the Palae-arctic and Ethiopian Regions, born by the desertification, is called Palaeo-eremic region. Usually the biota of the Levant is considered to be Palearctic (Mediterranean), but, according to Por (1975), the inclusion of the Old World deserts in the Holarctis is due to a certain ―Europa-centrism‖ of the scientists. ―According to this scheme, the Ethiopis begins only south of the Sahara, in the Savannas, where the last of the typical Palaearctic animals fade out. However, if at all, the connections of of the Palaeo-eremic faunal inventory are much closer to the Ethiopis than to Palaearctis….Our [Israel‘s] typical desert animals, such as scorpions, agamas, gerbils, sand grouse and gazelles, have Ethiopian rather than Holarctic connections, even if some species are so-called ―Mediterraneans‖ (Por, op. cit.). According to the systems of Bodenheimer (1935) and Por (1975), in the Levant could be distinguished four faunal elements: Palaearctic, Palaeo-eremic, Ethiopian and Oriental, with clear prevalence (especially in the north) of the Palaearctics in most of the animal groups. The line along the foothills of Northern Galilee and the Golan Heights was called by Por (1975) ―Nehring Line‖. South of it follows a ―transitional zone‖, where Palaearctic elements mingle with Palaeo-eremic elements. More to the south Por (1975) outlines a ―Bodenheimer Line‖, which is the end of the transitional zone. Follow Ramon Mountains and the mountains of Sinai. The ―Ethiopian element does not prevail in any area. The Oriental species are even more scattered, without showing any geographical pattern. The endemic species among the animals are chiefly of Palaeo-eremic, Ethiopian and also of Oriental origins.…This suggests a higher age of the tropical element, and perhaps a younger and more expansive character of the Holarctic species‖ (Por). In conclusion, ―The Levant province is a meeting place and transitional area between the Palaearctic, Oriental and Ethiopian zoogeographic regions. The broad ―Palaeo-eremic desert belt serves as a filtering barrier between the three faunal regions‖. The islands of Macaronesia (Canaries, Madeira, Azorean Island) are here treated as part of the Palearctic Region, Cabo Verde islands as part of the Afrotropical Region. Africa between 20oN and Zambezi – Kunene (tropical, or intertropical Africa) Geography, General Zoogeography and Paleogeography The sub-Saharan Africa is situated between the usually accepted southern border (actually transition belt) of the Palearctic (Holarctic), a frontier between one kingdom and another (Paleotropica). South of the Sahara follow the belts of semidesert (Sahel) and the different types of savanna. In the described area there are also high mountains with afroalpine vegetation and mountain rainforest. Tropic rain forests are tropical moist forests of semi-deciduous varieties distributed across nine West African countries. In the first half of the 1980s, an annual forest loss of 7200 square kilometers was note down along the Gulf of Guinea, a figure equivalent to 4-5 percent of the total remaining rain forest area. By Ecol. Mont., 7, 2016, 464-506 465 ARACHNOGEOGRAPHICAL COMPARISON BETWEEN WEST PALEARCTIC AND AFROTROPICAL REGION 1985, 72 percent of West Africa's rainforests had been transformed into fallow lands and an additional 9 percent had been opened up by timber exploitation. It is generally believed that firewood provides 75 per cent of the energy used in sub-Sahara Africa. With the high demand, the consumption of wood for fuel exceeds the renewal of forest cover. The rain forests which remain in West Africa now merely are how they were hardly 30 years ago. In Guinea, Liberia and the Ivory Coast, there is almost no primary forest cover left unscathed; in Ghana the situation is much worse, and nearly all the rain forest are cut down. Guinea-Bissau loses 200 to 350 km² of forest yearly, Senegal 500 km² of wooded savanna, and Nigeria 6,000,050,000 of both. Liberia exploits 800 km² of forests each year. Tropical Africa is about 18% of the world total covering 20 million km² of land in West and Central Africa. Recent estimates show that the annual pace of deforestation in the region can vary from 150 km² in Gabon to 2900 km² in Ivory Coast. Remaining tropical forest still cover major areas in Central Africa but are abridged by patches in West Africa. The tropical environment is rich in terms of bio-diversity. Tropical African forest is 18 per cent of the world total and covers over 3.6 million square kilometers of land in West, East and Central Africa. This total area can be subdivided to 2.69 million square kilometers (74%) in Central Africa, 680,000 square kilometers (19%) in West Africa, and 250,000 square kilometers (7%) in East Africa. In West Africa, a chain of rain forests up to 350 km long extends from the eastern border of Sierra Leone all the way to Ghana. In Ghana the forest zone gradually dispels near the Volta river, following a 300 km stretch of Dahomey savanna gap. The rain forest of West Africa continues from east of Benin through southern Nigeria and officially ends at the border of Cameroon along the Sanaga river. The variety of the African rain forest flora is also less than the other rain forests. This lack of flora has been credited to several reasons such as the gradual infertility since the Miocene, severe dry periods during Quaternary, or the refuge theory of the cool and dry climate of tropical Africa during the last severe ice age of about 18000 years ago (Jeannel, 1961, Moreau, 1952, 1955, 1963, Rabinowitz, Coffin & Falvey, 1983) What concerns the insects, it is worth reading Jeannel (1961, ―La Gondwanie et le peuplement d‘Afrique‖): ‖Le peuplement entomologique de l‘Afrique gonwanienne peut être envisagé comme échelonné sur deux périods. Au Jurassique et au Crétacé, c‘est l‘histoire des lignées autochtones qui ont pris naissance sur cette portion d‘Inabrésie. Au Tertiaire, c‘est l‘histoire des migrations des lignées indo-malaises qui se repandent en Afrique, realisant le type de distribution gondwanienne orientale; c‘est aussi l‘histoire des lignées sudamadiennes qui se repandent dans l‘Afrique intertropicale. Alors que le peuplement de l‘Afrique de l‘ère Secondaire a été seulement realisé par des lignées autochtones, selui de l‘ère Tertiaire se complique par des apports venus de l‘Est et du Sud‖. This analyses of the prominent French entomologist are based mostly on the distribution and the affinities of Pselaphids and some other groups of Coleoptera. When studying the distribution of Arachnida, it seems useful to compare the conclusions of Jeannel with the data extracted of the recent profound research on many groups of Arachnids. It is clear, for exemple, that such group as Ricinulei in Africa (only in the western part) suggest former very old ―Afrobrazilian‖ distribution. There are similar exemples also by other groups (the family Neogoweidae – Opiliones, Cyphophthalmi). Jeannel (loc.cit.) indicates that the equatorial forest in Central Africa was established in the Pliocene and is only a residu of a much more extensive forest, coming from the Malaisian area and broken by the wast steppic and desert regions of India, Arabia and East Africa, deprived of forest by climatic changes and human activities. So, when defining a group, family or genus of Arachnids (for exemple in Madagascar) as having African origin, we should think of a former origin of these (now East African) elements from South East Asia. Some of the elements in the present day intertropical fauna of Africa actually originate of the southern ―sudamadian‖ areas. Paleogeography and Paleoclimatology Laurasia The name of the northern supercontinent, which was detached from Pangaea by 200 Ma, came from the fusion of the names of Laurentia (the North American craton) and Eurasia. The supercontinent consists roughly by Laurentia, Siberia, Baltica, Kazakhstania, and the North China and East China crаtons. Laurasia 466 BERON is considered as a Mesozoic phenomenon. Europe separated from Greenland in the Paleocene (ca. 60 million years ago). Laurentia (North America) was detached from Eurasia around Cretaceous. Table 1. Comparison between the orders, suborders and families of Arachnida in the Afrotropical and Palearctic Regions. Group Region Region Afrotropical Palearctic Order Palpigradi present present Fam. Eukoeneniidae present present Order Ricinulei present absent Fam. Ricinoididae present absent Order Solifugae present present Fam. Galeodidae present present Fam. Karschiidae absent present Fam. Daesiidae present present Fam. Solpugidae present present Fam. Ceromidae present (southern Africa) absent Fam. Melanoblossiidae present (southern Africa) absent Fam. Gylippidae present (southern Africa) present Fam. Hexisopodidae present (southern Africa) absent Fam. Rhagodidae absent present Order Scorpiones present present Fam. Bothriuridae present (Namibia) Indian Himalaya Fam. Buthidae present present Fam. Pseudochactidae absent present Fam. Euscorpiidae absent present Fam. Scorpiopidae absent present Fam. Troglotayasicidae absent ? present Fam. Iuridae absent present Fam. Diplocentridae absent present Fam. Hemiscorpiidae absent present Fam. Hormuridae present present Fam. Scorpionidae present present Fam. Akravidae absent present Fam. Lisposomidae present (South Africa) absent Order Schizomida present absent Fam. Hubbardiidae present absent Order Uropygi present (relict) present Fam. Hypoctonidae present present Order Amblypygi present present Suborder Neoamblypygi present present Fam. Charinidae present present Fam. Phrynichidae present present Suborder Paleoamblypygi present absent Fam. Paracharontidae present absent ..continued on the next page Ecol. Mont., 7, 2016, 464-506 467 ARACHNOGEOGRAPHICAL COMPARISON BETWEEN WEST PALEARCTIC AND AFROTROPICAL REGION Table 1. (Continued) Order Opiliones present present Suborder Cyphophthalmi present present Fam. Sironidae present present Fam. Ogoveidae present absent Fam. Neogoveidae present absent Fam. Pettalidae present (South Africa) absent Suborder Eupnoi present present Fam. Caddidae present (South Africa) present (Japan) Fam. Phalangiidae present present Fam. Sclerosomatidae present present Fam. Neopilionidae present (South Africa) absent Suborder Dyspnoi absent present Fam. Ischyropsalididae absent present Fam. Sabaconidae absent present Fam. Dicranolasmatidae absent present Fam. Trogulidae absent present Fam. Nemastomatidae absent present Fam. Nipponopsalididae absent present (Japan, Korea) Suborder Laniatores present present Fam. Cladonychiidae absent present (Holoscotolemon) Fam. Travuniidae absent present Fam. Triaenonychidae present (South Africa) present (Japan, Korea) Fam. Assamiidae present absent Fam. Biantidae present Nepal, India Fam. Epedanidae absent Japan, India, Taiwan, Fam. Phalangodidae absent present Fam. Podoctidae present present (Japan, India) Fam. Pyramidopidae present absent Fam. Samoidae present (Mozambique) absent Fam. Zalmoxidae Seychelles absent Order Pseudoscorpiones present present Suborder Epiocheirata present present Fam. Chthoniidae present present Fam. Tridenchthoniidae present, India, Seychelles, Japan, Bhutan, Nepal, Fam. Pseudotyrannochthon. absent present Fam. Lechytiidae present present Fam. Feaellidae present India Suborder Iocheirata present present Fam. Ideoroncidae present present Fam. Hyidae absent India, Fam. Gymnobisiidae present (South Africa) present Fam. Neobisiidae present present Fam. Syarinidae present present ..continued on the next page 468 BERON Table 1. (Continued) Fam. Garypidae present present Fam. Geogarypidae present present Fam. Larcidae absent present Fam. Cheiridiidae present present Fam. Pseudochiridiidae present India, Nepal Fam. Olpiidae present present Fam. Garypinidae present (South Africa) present Fam. Menthidae present (Socotra) present (Israel) Fam. Sternophoridae present India Fam. Withiidae present present Fam. Cheliferidae present present Fam. Atemnidae present present Fam. Chernetidae present present Order Araneae present present Suborder Mesothelae absent present Fam. Liphistiidae absent present Suborder Orthothelae present present Infraorder Mygalomorphae present present Fam. Microstigmatidae present (South Africa) absent Fam. Hexathelidae (Macrothelinae) present present Fam. Dipluridae (Euagrinae) present present Fam. Nemesiidae present (southern Africa) present Fam. Theraphosidae present present Fam. Atypidae present present Fam. Antrodiaetidae absent present (Japan) Fam. Cyrtaucheniidae present present Fam. Idiopidae present present Fam. Ctenizidae present (southern Africa) present Fam. Migidae present absent Infraorder Araneomorphae present present Fam. Archaeidae present (southern Africa) absent Fam. Hypochilidae absent present Fam. Austrochilidae present (southern Africa) absent Fam. Filistatidae present present Fam. Drymusidae present (South Africa) absent Fam. Scytodidae present present Fam. Sicariidae present present Fam. Leptonetidae absent present Fam. Ochyroceratidae present ? present (China) Fam. Telemidae present present Fam. Pholcidae present present ..continued on the next page Ecol. Mont., 7, 2016, 464-506 469 ARACHNOGEOGRAPHICAL COMPARISON BETWEEN WEST PALEARCTIC AND AFROTROPICAL REGION Table 1. (Continued) Fam. Caponiidae present absent Fam. Tetrablemmidae present India Fam. Dysderidae present absent Fam. Oonopidae present present Fam. Orsolobidae present (southern Africa) absent Fam. Segestriidae present present Fam. Eresidae present present Fam. Hersiliidae present present Fam. Oecobiidae present present Fam. Palpimanidae present present Fam. Mimetidae present present Fam. Deinopidae present present Fam. Uloboridae present present Fam. Anapidae present present Fam. Araneidae present present Fam. Cyatholipidae present present Fam. Linyphiidae present present Fam. Sinopimoidae absent China (doubtfull status) Fam. Symphytognathidae present present (1 sp. in Japan) Fam. Synaphridae absent (but pres. on Madagascar) present Fam. Tetragnathidae present present Fam. Nephilidae present present Fam. Theridiidae present present Fam. Theridiosomatidae present present Fam. Ctenidae present present Fam. Lycosidae present present Fam. Oxyopidae present present Fam. Pisauridae present present Fam. Psechridae absent present Fam. Trechaleidae absent present Fam. Zoridae present present Fam. Zorocratidae present absent Fam. Zoropsidae present (South Africa) present Fam. Agelenidae present present Fam. Amaurobiidae present present Fam. Anyphaenidae present (South Africa) present Fam. Cybaeidae absent present Fam. Desidae present present Fam. Dictynidae present present Fam. Hahniidae present present Fam. Sparassidae present present Fam. Selenopidae present present ..continued on the next page 470 BERON Table 1. (Continued) Fam. Zodariidae present present Fam. Chummidae present (South Africa, end.) absent Fam. Clubionidae present present Fam. Miturgidae present present Fam. Phyxelididae present present Fam. Titanoecidae absent present Fam. Ammoxenidae present (southern Africa) absent Fam. Cithaeronidae present present Fam. Gallieniellidae present absent Fam. Gnaphosidae present present Fam. Prodidomidae present present Fam.Trochanteriidae present present Fam. Philodromidae present present Fam.Thomisidae present present Fam. Salticidae present present Fam. Corinnidae present present Fam. Liocranidae present present Order Opilioacarida present present Fam. Opilioacaridae present present Order Holothyrida absent (only Seychelles) absent Order Ixodida present present Fam. Argasidae present present Fam. Ixodidae present present Fam. Nuttalliellidae present (southern Africa) absent Order Mesostigmata present present Order Sarcoptiformes present present Order Trombidiformes present present Analysis and comments` The Afrotropical Region is bordering only the western part of the huge Palearctic Region (Saharo-Sindian Province). For many higher taxa is marked ―present‖ for both Regions, but actually they may be distributed in tropical Africa and, say, Japan or Korea. Sometime inbetween there is a gap of thousands kilometers and the difference between the arachnofauna of tropical Africa and Europe is much bigger. Some comments on the distributions of the various taxa are needed. General Arachnology and Arachnogeography The northern boundaries of several orders cross the described area. Uropygi do not live in W. Palearctic (one sp. in West Africa). The northern limit of Amblypygi is Morocco, Egypt, Israel, south Asia Minor, and the islands Rhodes and Cos. The scorpions barely leave the Mediterranean climate in Europe. Schizomida and Ricinulei are missing in the whole Palearctic. Solifugae (3 fam.) are limited to the south of Balkan peninsula, in Iberian Peninsula and Sicily. In Europe Opilioacarida are represented only in the southernmost areas of Italy and Greece and some islands. Palpigradi are distributed up to France, Austria, Slovenia, Hungary and Romania. The opilions Laniatores in Europe live mostly in the southern caves (considered by some as relicts). Similar is the situation of another suborder of Opiliones – Cyphophthalmi, well represented in Iberian and Balkan peninsulas, but almost entirely missing in the North (one species in Poland). The other orders (Araneae, Pseudoscorpions, Opiliones ―Palpatores‖, Acariformes and Parasitiformes) are better Ecol. Mont., 7, 2016, 464-506 471 ARACHNOGEOGRAPHICAL COMPARISON BETWEEN WEST PALEARCTIC AND AFROTROPICAL REGION represended in Northern Europe, but many families are missing. Entirely lacking in Palearctic and in continental Africa is the order Holothyrida (Parasitiformes) (Beron, 2000, 2008b, Griffin, 1998, 2000, Haddad, 2004, Harvey, 2003, Newlands, 1978), Vachon, 1950). All orders of Arachnida are represented in tropical Africa, some of them by endemic families and even suborders, like Paleoamblypygi – one sp., considered a ―living fossil‖). Between 20oN and Kunene- Zambezi live six families of Solifugae, three fam. of Amblypygi (one endemic), Uropygi are represented by one endemic genus (Etienneus) in West Africa, in the whole of Africa there are four genera of one family of Schizomida (in one family; one subfamily endemic for southern Africa), 71 families of spiders, 15 fam. of Opiliones, 20 fam. of Pseudoscorpiones, 3 end. genera of Opilioacarida, numerous mites and ticks (Acaridida, Oribatida, Prostigmata, Mesostigmata, Ixodida). There are 8 described species of Palpigradi, one genus (Allokoenenia Silv.) is endemic. Lacking are the suborders Dyspnoi (Opiliones), Mesothelae (Araneae) Special attention should be paid to the cave and high mountain Arachnids. The numerous caves of Europe, Caucasus, Lebanon and Asia Minor harbour many endemic genera and species of Arachnida (mostly Araneae, Pseudoscorpiones, Opiliones). North of the Alps the caves fauna is much poorer and the troblobites are almost non existing (presumably due to the glaciations). Some cave Arachnida are known also from Morocco and Algeria. Palpigradi In Europe (including Madeira) are registered 28 species of Palpigradi, all belonging to one genus – Eukoenenia Börner (fam. Eukoeneniidae). They are recorded (without counting the subspecies) from France (9), Italy (12), Greece (3), Bulgaria (1), Hungary (1), Austria (2), Bosna and Herzegovina (1), Croatia (2), Malta (2), Portugal (2), Romania (4), Slovakia (1), Slovenia (1), Spain (4) – 14 countries (Bertrand, 1980, Christian, 1998, Condé, 1951, 1956, 1979a, 1979b, 1980, 1990, 1996, Hansen, 1926, Mayoral & Barranco, 2002b, Peyerimhoff, 1902, 1908, Remy, 1948, 1952, 1957, Rowland & Sissom, 1980). Christian et al. (2014) described two species from Italy (Eukoenenia lanai Christian and E. roscia Christian) and concluded that ―Our area [the Alps of SW Piedmont] is part of the most prominent Pleistocene refugium of the SW Alps…. It lies south of the Holdhaus line, which roughly marks the northern range boundaries of troglobiotic beetles and other subterranean arthropods with similarly limited means of dispersal… Though the area overlaps the border of maximum Würm glaciation (Casazza et al. 2008), subterranean habitats for palpigrades may have existed continuously since late Neogene times‖. Christian (2014) described also the new troglobite species Eukoenenia vargowitshi from Abhazia. In Iberian Peninsula (all from Spain and from the two slopes of the Pyrenees) have been recorded Eukoenenia bouilloni Condé, 1980, E. brolemanni (Hansen, 1926), E. draco zariquieyi (Condé, 1951), E. hispanica (Peyerimhoff, 1908), E. pyrenaella Condé, 1990, E. pyrenaica (Hansen, 1926), E. gadorensis Mayoral et Barranco, 2002, E. mirabilis (Grassi et Calandruccio, 1885)(Barranco & Mayoral, 2007). Remy (1949) reported from Corsica Eukoenenia mirabilis and E. berlesei. Eukoenenia patrizii Condé, 1956 (endemic) was described from Sardinia, E. mirabilis was recorded from Sardinia by Roewer (1953). From Sardinia Condé & Heurtault (1995) described a second troglobitic (endemic ?) species – E. grafittii. The first known species of the order E. mirabilis (Grassi et Calandruccio, 1885) is known from Sicily. As a whole, six species of Palpigradi have been recorded from seven islands of the Mediterranean: Sicily, Sardinia, Majorca, Iraklia nr. Naxos, Kithira, Corfu and Malta. The only species known from Malta is Eukoenenia christiani Condé from a cave (Condé, 1988). Barranco & Mayoral (2007) described from the cave of Kef Aziza in Morocco the new species Eukoenenia maroccana – the third Moroccan Palpigradi, after E. mirabilis (Grassi et Calandruccio, 1885) and E. hanseni (Silvestri, 1913). Very few Palpigradi have been described from tropical Africa: Eukoenenia pauli Condé, 1979 (Gabon), E. angolensis (Remy, 1956), E. machadoi (Remy, 1950) (Angola), E. hesperia (Remy, 1953)(Ivory Coast), E. kenyana Condé, 1979 (Kenya), Koeneniodes notabilis Silvestri, 1913, Leptokoenenia scurra Monniot, 1966 (Congo), and Allokoenenia afra Silvestri, 1913 (Guinea) (Remy, 1950, 1953, 1956, Monniot, 1966, Condé, 1979, Silvestri, 1913). The genus Allokoenenia is endemic for tropical Africa. This number does not reflect the real picture of the distribution of Palpigradi between 20oN and Kunene-Zambezi. These tiny and fragile creatures are rarely collected, mostly by the few specialists on them in person. 472 BERON Eukoenenia lawrencei Remy, 1957 is the only species of this order, known from Southern Africa. Solifugae The list of European Solifugae of Blick (2004) contains 18 sp., but geographically half of them live on territories outside Europe (Rhodes, Asiatic Turkey, Cyprus, Canaries). On the Balkan Peninsula (continental part, mostly in Greece), are recorded members of three families of Solifugae (Alexiou, 2014): Fam. Galeodidae – Galeodes Olivier: G. graecus C.L. Koch (Greece, Bulgaria), G. elegans Roewer (Rep. Macedonia), G. hellenicus Roewer (Greece). Fam. Karschiidae – Barrussus furcichelis Roewer, Eusimonia nigrescens Kraepelin (Greece). Fam. Daesiidae – Biton ehrenbergi Karsch (Greece). The northern boundary of Solifugae in eastern Europe (Galeodes graecus C.L. Koch, G. araneoides Pallas) runs through SW Bulgaria (G. graecus C.L. Koch), Rep. Macedonia (G. elegans Roewer) and Ukraine (Map 1). On Iberian Peninsula is known Gluvia dorsalis (Latreille)(Daesiidae)(Spain, Portugal). In the Canary Islands live Eusimonia wunderlichi Pieper (Karschiidae). Only one species is known in Ukraine: Galeodes araneoides (Pallas) (Galeodidae). From Mediterranean islands only on Sicily are known two species of Daesiidae (Biton ehrenbergi Karsch and B. velox Simon). In North Africa are distributed Solifugae from 19 genera and the families Galeodidae, Karschiidae, Daesiidae, Solpugidae, and Rhagodidae. A numerous genus is Galeodes Olivier with 25 sp. in N. Africa. Endemic genus of Solifugae for North Africa is Barrus Simon, 1880 – Egypt (1 sp.) Figure 1. Distribution of Solifugae in Europe. Ecol. Mont., 7, 2016, 464-506 473
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