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.

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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

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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

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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

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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

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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

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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

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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

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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.

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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.

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Table 2. Distribution of Solifugae in North Africa.

Country Mauritania Morocco Algeria Tunisia Libya Egypt No of sp. 4 25 30 23 19 27 Fam. Daesiidae + + + + + + Biton Karsch - 1 6 5 5 4 Blossia Simon 1 5 1 1 - 2 Gluviopsilla Roewer - - 1 - - - Gnosippus Karsch - - - - - 1 Tarabulida Roewer - - - - 1 - Fam. Galeodidae + + + + + + Galeodes Olivier 1 9 7 9 7 1 Galeodopsis Birula - - - - 1 - Othoes Hirst 1 - 1 - - - Paragaleodes Kraepelin 1 1 2 1 - 2 Fam. Karschiidae - + + - + + Barrus Simon - - - - - 1 Eusimonia Kraepelin - 4 3 - 1 1 Fam. Rhagodidae - + + + + + Rhagodes Pocock - - 1 1 1 3 Rhagoditta Roewer - - 2 1 - 1 Rhagodira Roewer - - 2 1 - 1 Rhagodeya Roewer - - - - 1 - Fam. Solpugidae - + + + + + Oparba Roewer - 1 - - - - Oparbella Roewer - 2 4 4 1 2 Solpuga Lichtenstein - - 1 - 1 - Zeria Simon - 1 1 - - -

Tropical Africa Non of the 6 families is endemic or typical for tropical Africa.

Endemic genera for this area: Blossiana Roewer (Ethiopia), Bitonota Roewer (Ethiopia), Bitonupa Roewer (Kenya), Solpugistella Turk (Kenya), Rhagodalma Roewer (Sudan), Rhagodippa Roewer (Djibuti). Benoit (1960) enumerated 29 species from the former Belgian Congo and Ruanda-Urundi. They all live far from the dense tropical forest of Congo and are concentrated mainly in Katanga. In tropical Africa (between 20oN and the line Kunene – Zambezi) are recorded 41 genera of Solifugae of 6 families: Ceromidae, Daesiidae, Galeodidae, Rhagodidae, Hexisopodidae and Solpugidae. The richest countries are Somalia (48) and Ethiopia (46)(Harvey, 2013c). Southern Africa (South Africa, Lesotho, Namibia, Swaziland, Botswana, Zimbabwe, southern part of Mozambique) is extremely rich in Solpugids. Here are represented six of the 12 families of Solifugae, incl. the endemic family Hexisopodidae and the endemic genera Hexisopus Karsch (15 sp.), Chelypus Purcell (9 sp.). With 127 sp. of Solifigae, 47 (37%) of which endemics and nine genera occurring only in Namibia, this country appears to be the richest in the world in Solifugae (GRIFFIN, 1998). Even more Solifugae (163 sp.) live in in South Africa. This makes their absence in Madagascar even more enigmatic.

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Table 3. Solifugae in the countries of Afrotropical Region.

Countries Ethiopia Eritrea Somalia Djibuti Uganda Rwanda Number of species 46 5 48 8 6 1 Fam. Ceromidae + - - - + - Ceroma Karsch 1 - - - 2 - Fam. Daesiidae + + + + - - Biton Karsch 7 2 8 2 - - Bitonota Roewer 1 - - - - - Blossia Simon - - 2 - - - Blossiana Roewer 1 - - - - - Eberlanzia Roewer - - 1 - - - Gluviopsilla Roewer - - 1 - - - Gluviopsis Roewer 1 - 5 1 - - Gnosippus Karsch - 1 - - - - Hemiblossia Roewer 1 - - - - - Triditarsula Roewer 1 - - - - - Fam. Galeodidae + - + + - - Galeodes Olivier 7 - 5 1 - - Gluviema Caporiacco - - 1 - - - Paragaleodes Kraepelin 2 - - - - - Fam. Rhagodidae + + + + + - Rhagodes Pocock - - 6 - - - Rhagodessa Roewer - 1 - - - - Rhagodia Roewer 1 - - - - - Rhagodinus Roewer 1 - - 1 - - Rhagodippa Roewer - - 1 - - Rhagoditta Roewer 1 1 1 - - - Rhagodoca Roewer 5 - 5 1 2 - Rhagoduna Roewer 1 - - - Fam. Solpugidae + - + + + - Ferrandia Roewer - - 2 - - - Solpuga Lichtenstein 1 - - - - - Solpugassa Roewer 2 - 1 2 - - Solpugeira Roewer 1 - - - - 1 Solpugella Roewer ------Solpugyla Roewer 1 - 1 - - - Zeria Simon 8 - 3 - 2 - Zeriassa Pocock 3 - 5 - - - Guinea Countries Sudan Senegal Gambia Guinea Chad Mali Niger Bissau Number of sp. 21 1 1 1 5 1 2 6 Taxa Fam. Daesiidae + - - - + - - + ..continued on the next page

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Table 3. (Continued) Biton Karsch 4 ------1 Blossia Simon 1 ------Gluviopsis Roewer ------1 Gnosippus Karsch - - - - 1 - - - Fam. Galeodidae + - - + + + + + Galeodes Olivier 5 1 - - 1 1 1 2 Othoes Hirst 1 ------Fam. Rhagodidae + - + - - - - - Rhagodalma Roewer 1 ------Rhagodessa Roewer 3 ------Rhagodeya Roewer 1 ------Rhagodolus Roewer - - 1 - - - - - Rhagoduna Roewer 1 ------Fam. Solpugidae + - - + + - - + Oparbella Roewer ------1 Solpugassa Roewer 1 ------Zeria Simon 2 - - 1 2 - - - Zeriassa Pocock 1 ------

Camero- Countries Nigeria Togo Malawi Ruanda Kenya Tanzania on Number of species 4 9 2 4 1 37 33 Taxa Fam. Ceromidae - - - + - + + Ceroma Karsch - - - 2 - 1 4 Fam. Daesiidae - + - - - + + Biton Karsch - 2 - - - 3 - Bitonupa Roewer - - - - - 1 - Blossia Simon - - - - - 2 3 Hemiblossia Roewer - - - - - 1 - Fam. Galeodidae + + + - - + - Galeodes Olivier 1 2 1 - - 1 - Fam. Rhagodidae + - - - - + - Rhagodes Pocock ------2 Rhagodoca Roewer - - - - - 7 - Rhagodolus Roewer 1 ------Fam. Solpugidae + + + + + + + Oparba Roewer 1 3 - - - - - Oparbella Roewer - 1 - - - - - Solpuga Licht. - - - - - 1 - Solpugassa Roewer ------1 Solpugella Roewer - - - - 1 - - ..continued on the next page

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Table 3. (Continued) Solpugiba Roewer - - - - - 1 - Solpugistella Turk - - - - - 1 - Solpugylla Roewer ------4 Zeria Simon 1 1 1 2 - 11 15 Zeriassa Pocock - - - - - 7 3 DR Mozamb- Countries Congo Angola Zambia Congo ique Number of sp. 1 30 31 4 13 Taxa Fam. Ceromidae - + + - - Ceroma Karsch - 1 1 - -

Fam. Hexisopodidae - - + + -

Chelypus Purcell - - 1 1 - Hexisopus Karsch - - 1 - -

An older, but most reliable source of information on the group is the check list of Lawrence (1955), listing from this area 181 species of 33 genera and 6 families. Wharton (1981) outlined the Solpugids of Namibia. Important papers on the subject, used for preparing the text, are (Benoit, 1960, Birula, 1912, 1917, Borelli, 1924, Chemini, 1995, Dippenaar-Schoeman & González Reyes, 2006, Drensky, 1936, El-Hennawy, 1998, 2007, 2008, Hewitt, 1919, Kraepelin, 1899, Lawrence, 1955, 1960, 1966, Levy & Shulov, 1964, Panouse, 1957, Purcell, 1903, Roewer, 1934, 1941, Simonetta & Delle Cave, 1968, Werner, 1925). The families Gyllipidae and Rhagodidae are shared with Asia, Solpugidae – only with the rest of Africa, Ceromidae – with southern part of tropical Africa and only Daesiidae is shared also with South America. Fossil Ceromid is recorded from Brazil.

Endemic genera of Solifugae south of Zambezi – Kunene: Fam. Ceromidae – Ceromella Roewer, Toreus Purcell. Fam. Gyllipidae – Bdellophaga Wharton, Trichotoma Lawrence. Fam. Hexisopodidae – Hexisopus Karsch, Chelypus Purcell. Fam. Daesiidae – Namibesia Lawrence. Fam. Melanoblossiidae (Melanoblossiinae endemic. in Southern Africa) – Daesiella Hewitt Lawrencega Roewer, Melanoblossia Purcell, Microblossia Roewer, Unguiblossia Roewer. Fam. Solpugidae – Solpugiba Roewer, Solpugista Roewer, Solpugema Roewer, Prosolpuga Roewer, Solpuguna Roewer, Metasolpuga Roewer.

Ricinulei All Ricinulei in Africa are known only from West Africa (11 sp., all of genus Ricinoides Ewing). The order is missing in Palearctic (Hansen, 1921, Hansen & Soerensen, 1904, Judson & Hardy, 2001, Legg, 1976, 1978, Tuxen, 1974).

Scorpiones From Europe, Asia Minor, Cyprus, Malta and Caucasus are recorded Scorpions of the families Euscorpiidae (Euscorpius Thorell), Buthidae (Buthus Leach, Mesobuthus Vachon ), Iuridae (Iurus Thorell, Protoiurus Soleglad et al., Calchas Birula, Neocalchas Yagmur et al.).

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Table 4. Scorpions in Northern Africa.

Country Morocco Algeria Tunis Libya Egypt Number of Species 50 30 17 16 28 Fam. Buthidae + + + + + Ananteroides Borelli - - - - - Androctonus Ehrenberg 7 7 4 3 5 Buthacus Birula 4 5 2 3 2 Butheoloides Hirst 5 1 - - - Buthiscus Birula - 1 1 1 - Buthus Leach 15 4 4 3 6 Cicileiurus Teruel 1 - - - - Cicileus Vachon - 1 - - - Compsobuthus Vachon 2 2 - 1 4 Egyptobuthus Lourenço - - - - 1 Hottentotta Birula 3 3 - 2 - Isometrus Ehrenberg - 1 - - - Leiurus Ehrenberg - - - - 1 Lissothus Vachon - 1 - 1 - Mauritanobuthus Qi et Lour. - - - - - Microbuthus Kraepelin 1 - - - 1 Orthochirus Karsch 3 2 1 1 2 Parabuthus Pocock - - - - 2 Pseudolissothus Lourenço - 1 1 - - Saharobuthus Lour. et Duhem 1 - - - - Fam. Euscorpiidae + + + + + Euscorpius Thorell 1 2 3 - - Fam. Scorpionidae + + + + + Nebo Simon - - - - 1 Scorpio L. 6 2 2 1 1 Fam. Hemiscorpiidae - - - - + Hemiscorpius Peters - - - - 1

Endemic genera for the area are: Mauritanobuthus Qi et Lourenço, 2007 – Mauritania (M. geniezi). Saharobuthus Lourenço et Duhem, 2009 – Western Sahara.

According to the lists of Vachon (1952) and Dupré (2013) (actualized), from the Northafrican countries are recorded scorpions of 24 genera and 4 families. From Tassili n‘Ajjer Vachon (1958) identified 10 species of scorpions of 7 genres and the families Buthidae and Scorpionidae. According to this author, in the ―saharo-mountainious‖ region (Tassili n‘Ajjer, Air, Adrar Iforas and Tibesti) is seen one altitudinal separation of species of Palearctic origin, having ―climbed‖ above 1000 m with the establishment of drought in North Africa, and species, widespread in Sahara.

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Figure 2. Distribution of scorpions in Europe. ■ – Euscorpius. O – Iurus. X – Mesobuthus. ♦ - Buthus. ! - Belisarius.

This list has been completed by Lourenço (2009) with the description of Compsobuthus tassili. From Tunisia Vachon (1951) identified 7 sp. of scorpions of the genera Euscorpius, Scorpio, Buthus, Androctonus, Buthacus and Buthiscus. Among the scorpions of Tunisia this author distinguishes two species of European origin (―Buthus” occitanus and Euscorpius sicanus) and five Northafrican elements. The scorpions of Egypt (12 sp.) have been listed as early as 1910 by E. Simon. Now the species in this country are 28. Graham et al. (2012) indicate that ―Molecular clock estimates suggest an ancient disjunction across the Mediterranean Sea‖(mean = 5.56 Mya, about the sudden refilling of the Mediterranean Sea after it had evaporated during the Messinian salinity crisis). The list of Morrocan scorpions (50 sp.) is more recent – of Nickel, Hornung & Koch (2009 a, b). Other important papers on the subject, used while preparing the text: El-Hennawy (1992a, 2009), Fet (1988), Fet, Sissom, Lowe & Braunwalder (2000), Kovařik (2001, 2003, 2009), Lamoral (1979), Lamoral & Reynders (1975), Levy & Amitai (1980), Lourenço (2003, 2007), Lourenço & Duhem (2007), Lourenço, Duhem & Cloudsley-Thompson (2010), Prendini (2011), Probst (1973), Qi Jian-Xin & Lourenço (2007), Simon (1910), Stockmann & Ythier (2010), Vachon (1941, 1966, 1968).

Endemic genus for the area is: Mauritanobuthus Qi et Lourenço, 2007 – Mauritania (M. geniezi).

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Table 5. Scorpions in Near East and palearctic Middle Asia Asia Country Minor Syria Lebanon Jordan Israel Iraq Kuwait Iran Afgan. Uzb. Kaz. Kir. Tadz. Number of sp. 26 20 10 19 19 20 6 55 29 8 5 2 6 Taxa Fam. Pseudochactidae ------+ + - - + Pseudochactas Gromov ------1 1 - - 1 Fam. Akravidae - - - - + ------Akrav Levy - - - - 1 ------Fam. Buthidae + + + + + + + + + + + + + Afghanobuthus Lour. ------1 - - - - Androctonus Ehr. 1 3 3 4 3 1 1 1 2 - - - - Anomalobuthus Kr. ------1 - 1 1 - - Apistobuthus Finn. ------1 1 - - - - - Birulatus Vachon - 1 - 1 1 - - - 1 - - - - Buthacus Birula - 3 1 1 2 2 1 1 1 1 - - - Buthus Leach - - - 2 1 ------Compsobuthus Vach. 2 5 1 5 4 2 2 9 3 - - - - Hottentotta Birula 2 1 1 1 1 3 - 6 5 - - - 1 Iranobuthus Kovarik ------Isometrus Ehr. - 1 ------Kraepelinia Vachon ------1 - - - - - Leiurus Ehrenberg 1 2 1 2 1 ------Liobuthus Birula ------1 - 1 1 - - Mesobuthus Vachon 4 2 1 - - 3 - 6 2 2 2 2 2 Odontobuthus Vachon - - - - - 2 - 5 - - - - - Orthochirus Karsch 1 - - 1 1 2 - 9 9 2 1 - 1 Polisius Fet et al. ------1 - - - - - Psammobuthus Birula ------1 Razianus Farzanpay ------1 - - - - - Sassanidiothus Far. ------2 2 - - - - Vachoniolus Levy et al. ------1 ------Fam. Euscorpiidae + - - - - (+) - + - - - - - Euscorpius Thorell 6 - - - - (+) ------Scorpiops Peters ------2 - - - - - Fam. Hemiscorpiidae - - - - - + + ------Hemiscorpius Peters - - - - - 1 6 ------Fam. Scorpionidae - + + + + + + + - - - - - Nebo Simon 1 1 1 1 - 1 1 ------Scorpio L. - 1 1 1 1 1 1 ------Fam. Iuridae + - - - - + ------Calchas Birula 4 - - - - 2 ------..continued on the next page

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Table 5. (Continued) Iurus Thorell 1 ------Neocalchas Yagmur et al. 1 ------

Protoiurus Soled.et al. 3 ------Country Saudi Turkm. Qatar UAE Bahrain Yemen Oman Arabia Number of sp. 7 29 6 14 4 37 39 Taxa Fam. Buthidae + + + + + + + Androctonus Ehr. - 4 1 1 1 2 2 Anomalobuthus 1 ------Kraepelin Apistobuthus Finn. - 1 1 1 - 1 1 Babycurus Karsch - - - - - 1 1 Buthacus Birula - 3 2 3 2 - 1 Butheolus Simon - 4 - 1 - 2 2 Buthus Leach - - - - - 1 - Compsobuthus Vachon - 5 1 4 - 6 4 Femtobuthus Lowe ------1 Hottentotta Birula - 2 - 2 - 4 4 Kraepelinia Vachon 1 ------Leiurus Ehrenberg - 4 - - - 3 2 Liobuthus Birula 1 ------Mesobuthus Vachon 1 ------Microbuthus Kraep. - - - - - 1 3 Odontobuthus Vach. ------2 Orthochiroides Kov. - - - - - 2 - Orthochirus Karsch 2 2 - - - 1 4 Parabuthus Pocock - 1 - - - 1 - Pectinibuthus Fet 1 ------Picobuthus Lowe ------2 Vachoniolus Levy et al. - 1 - 1 - - 3 Fam. Euscorpiidae - - - - - (+) - Euscorpius Thorell - - - - - (+) - Fam. Hemiscorpiidae - + - + - + + Hemiscorpius Peters - 1 - 1 - 3 3 Fam. Scorpionidae - + + - - + + Heteronebo Pocock - - - - - 2 - Nebo Simon - - - - - 4 3 Pandinus Thorell - - - - - 1 - Scorpio Linnaeus - 1 1 - - 1 -

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Endemic genera for this area are: Fam. Pseudochactidae Pseudochactas Gromov, 1998 (Turkmenistan, Afghanistan). Fam. Buthidae Neohemibuthus Lourenço, 1996 (Iran). Paraorthochirus Lourenço et Vachon, 1995 (Iran).

Fam. Akravidae Akrav Levy, 2007 – Israel.

Table 6. Scorpions in Western and Central Tropical Africa.

Guinea Burk. DR Countries Senegal Gambia Guinea Bissau Liberia Ghana Faso Mali Congo Congo Gabon Number of species 11 2 15 3 3 5 5 8 18 14 9 Fam. Buthidae + + + + + + + + + + + Ananteroides Borelli - - 1 1 - - - - 1 1 - Androctonus Ehrenberg 1 ------2 4 4 4 Babycurus Karsch 2 - 3 - - 3 - - 1 - - Buthacus Birula 2 ------1 Butheoloides Hirst 1 - - - - 1 1 1 2 1 1 Buthus Leach 2 - 1 - - - - - 1 - - Compsobuthus Vachon - - - - - 1 1 1 - - - Hottentotta Birula 1 1 1 1 - - 1 1 2 2 - Uroplectes Peters - - - - 1 ------Fam. Scorpionidae + + + + + + + + - - 1 Pandinus Thorell 1 1 4 1 1 1 1 2 + + + Scorpio L. 1 - - - - - 1 - - - - Fam. Hormuridae - - + - + - - - 2 3 1 Hormurus Thorell - - 2 - - - - - + + + Liocheles Sundevall - - 1 - - - - - 1 - - Opisthacanthus Peters - - 2 - - - - - 1 1 2 DR Centr Countries Congo Gabon Angola Congo AR Number of sp. 18 14 9 4 24 Taxa Fam. Buthidae + + + + + ..continued on the next page

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Table 6. (Continued) Akentrobuthus 1 1 - - - Lam. Babycurus Karsch 4 4 4 1 3 Congobuthus Lour. 1 - - - - Darchenia Vachon - - 1 - - Hottentotta Birula 2 1 1 2 1 Isometrus Ehrenb. 1 - - - - Leiurus Ehrenberg - - - 1 - Lychas C. L. Koch 2 2 - - 1 Parabuthus - - - - 5 Pocock Uroplectes Peters - - 1 1 - Fam. + + + + + Scorpionidae Opistophthalmus - - - - 5 Koch Pandinus Thorell 2 3 1 1 -

Schizomida The order is missing in the Palearctic. In Afrotropical Region are known four genera: Artacarus Cook, 1898 (1 sp.), endemic to West Africa (Liberia and Ivory Coast), Lawrencezomus Armas (1 sp.)(Liberia), Kenyazomus Armas (1 sp.), endemic for Kenya, and Megaschizomus Lawrence, 1955 (Southern Africa, endemic subfamily Megaschizominae) (Armas, 2010, Harvey, 2003, Lawrence, 1955, Reddell & Cokendolpher, 1995).

Figure 3. Distribution of Schizomida in Africa.

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Amblypygi From Turkey and the Near East there are known only six species of Amblypygi: Fam. Charinidae Charinus ioanniticus (Kritscher, 1959) – Turkey, Israel, the Dodecanese islands Kos and Rhodes (known also from Egypt). Charinus omanensis Delle Cave, Gardner et Weygoldt, 2009 – Oman (cave). Charinus dhofarensis Weygoldt, Pohl et Polák, 2002) – Oman. Fam. Phrynichidae Phrynichus dhofarensis Weygoldt, Pohl et Polák, 2002 – Oman. Ph. jayakari Pocock, 1894 – Hadramaut. These are among the species outlining the northern limit of Amblypygi in the Old World. Actually, Yemen, Oman and it‘s province Dhofar are part of the south of Arabian Peninsula, considered usually as part of the Afrotropical Region. Musicodamon atlanteus Fage, 1939 (Phrynichidae) is endemic genus and species from Morocco and Algeria. El-Hennawy & Hisham (2002) recorded from Egypt Charinus ioanniticus (Charinidae), known also from Turkey, Israel and the Greek islands Kos and Rhodes. Charinidae and Phrynichidae are the only families of Amblypygi, known from the Palearctic Region. The first list of African Amblypygi (Lawrence, 1969) contained 16 sp. Without the North African Musicodamon and the island Charinus spp. from Socotra, the Seychelles and Madagascar remain from tropical Africa 13 sp. Since several nomenclatorial changes took place. The new genus Euphrynichus Weigoldt, 1995 has been described, Titanodamon Pocock was synonymised with Damon C.L. Koch, Myodalis Simon – with Phrynichus Karsch. Paracharon caecus Hansen should be added to the list. “Hemiphrynus” machadoi Fage is now Xerophrynus Weigoldt, 1996 (endemic). Now (2016) in tropical Africa are known 20 sp. of Amblypygi of 6 genera and 3 families (Charinidae, Phrynichidae, Paracharontidae).

Fam. Charinidae Charinus Simon – widespread in the world, in tropical Africa known from Tanzania, Guinea, Ecuatorial Guinea, São Tome, and Somalia (6 sp.) Fam. Phrynichidae Damon C.L. Koch – widespread and endemic in tropical Africa (11 sp., 8 in tropical Africa north of Kunene - Zambezi). Euphrynichus Weygoldt – Kenya, Tanzania, Angola, Malawi, Zimbabwe, 2 sp. (E. amanica (Werner, 1916), E. bacillifer (Gerstaecker, 1873). Phrynichus Karsch – Eritrea, Somalia, Djibouti, Kenya, Cameroon, Rwanda, Tanzania, Congo (7 sp.). Xerophrynus Weigoldt – Angola (1 sp. X. machadoi with unclear position), Namibia. Fam. Paracharontidae Paracharon Hansen - P. caecus Hansen, 1921.

The most interesting Amblypygid in tropical Africa is Paracharon Hansen from Guinea - Bissau (1 sp.), only member of the family Paracharontidae and the suborder Paleoamblypygi (―living fossil‖) (Delle Cave, 1986, Delle Cave, Gardner & Weygoldt, 2009, Fage, 1939, 1954, Hansen, 1921, Harvey, 2003, 2013c, Kovařík & Vlasta, 1996, Kritscher, 1959, Lawrence, 1958, Pocock, 1895, Prendini, Weygoldt & Wheeler, 2005, Weygoldt, 1995, 1996, 2000a, 2000b), Weygoldt, Pohl & Polák, 2002, Whittick, 1941).

Uropygi (Thelyphonida) The enigmatic Etienneus africanus (Hentschel, 1899) from Senegal, Gambia, Guinea and Guinea-Bissau is now considered autochthonous and relict (Huff & Prendini, 2009). Nowhere else in Africa live Uropygi. They are missing also in the Palearctic (Harvey, 2013, Cooke & Shadab, 1973, Hentschel, 1899, Heurtault, 1984).

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Pseudoscorpiones The Pseudoscorpions of Europe have been summarized by Beier (1963), but many new species, genera and even families have been added since. According to Lissner (2014), in Europe are known 73 genera of Pseudoscorpiones, belonging to 14 families (Chthoniidae, Neobisiidae, Bochicidae, Larcidae, Syarinidae, Olpiidae, Garypidae, Geogarypidae, Garypinidae, Atemnidae, Cheiridiidae, Cheliferidae, Chernetidae, Withiidae) – nearly half of the Pseudoscorpiones families in the World. ―The European pseudoscorpion fauna counts to 861 species in 73 genera and 16 families‖ (Harvey 2013). These numbers include species and genera from Asian parts of Russia and Turkey (perhaps 20 species?). The fauna is much more diverse in the southern parts of Europe. The highest number of species is found in Italy (226) followed by Spain (196). Blick, Muster & Duchác (2004) list 102 pseudoscorpion species for Central Europe. Species number decline to about 20 in southern Scandinavia (Beier, 1963). None, except a few introduced synanthropic species, are found at higher latitudes than ca. 65°N (Beier, 1963)‖. According to Dashdamirov & Schawaller (1992), in Caucasus have been recorded 66 sp. of Peudoscorpions, belonging to 29 genera and 10 families. In Armenia are known 19 sp. of 12 genera and the families Chthoniidae, Neobisiidae, Garypidae, Cheiridiidae, Olpiidae, Cheliferidae, Chernetidae. In Georgia are known 46 sp. In Azerbaidjan are known 48 sp. of 21 genera and 8 families (Dashdamirov, 1990). In this paper is done a zoogeographical analysis with 11 zoogeographical complexes. Asia-Minor – Caucasian elements are 23.0% of species; endemic Caucasian complex includes 6 species (17.9%); Palearctic type of areal comprises 15.4%; Mediterranean elements are 10.3%, European complex – 10.3%. Two species are cosmopolytic. Some other important papers on the subject: Beier (1944, 1955, 1959, 1961, 1967, 1972, 1979), Callaini (1983c, 1988), Christophoryová, Štahlavský, Krumpál & Fedor (2012), Šurţiš (1974), Šurţiš, Dimitrieviš & Legakis (2004), Daday (1889), El-Hennawy (1988), Gardini (1994, 2000), Harvey (1990, 1992, 2003, 2011, 2013a), Heurtault (1974, 1983, 1990), Leclerc (1989), Legg (1988), Legg & O‘Connor (1997), Lessert (1911), Mahnert (1981, 1982a, 1982b, 1982c, 1983a, 1983b, 1985, 1988, 2004), Meinertz (1962), Petrov (1997), Rafalski (1967), Redikorzev (1924, 1926, 1930), Schawaller (1983), Schawaller & Dashdamirov (1988), Telnovs (2002a), Vachon (1954).

Endemic genera of Pseudoscorpiones in Europe: Fam. Chthoniidae Neobalkanella Šurţiš, 2013 - Serbia (1 sp.). Troglochthonius Beier, 1939 - Bosnia and Herzegovina, cave (1 sp.); ? Sardinia. Fam. Neobisiidae Insulocreagris Šurţiš, 1987 – Croatia (Vis), Herzegovina (2 sp.) Protoneobisium Šurţiš, 1988 – Croatia (1 sp.). Archeoroncus Šurţiš et Rada, 2012 – Croatia. Ernstmayria Šurţiš et Dmitrijeviš, 2006 – Crete (1 sp.). Fam. Syarinidae Arcanobisium Zaragoza, 2010 – Spain (A. comasi Zaragoza, 2010). Hadoblothrus Beier, 1952 – Italy, Greece (2 sp.). Pseudoblothrus Beier, 1931 – Italy, Crimea, France, Switzerland. Microcreagrella Beier, 1961 – Azores, Madeira. Fam. Bochicidae Titanobochica Zaragoza et Reboleira 2010 – Portugal (T. magna Zaragoza et Reboleira, 2010). Troglobisium Beier, 1939 – Spain [T. racovitzai (Ellingsen, 1912)].

From North Africa (Egypt, Libya, Tunisia, Algeria, Morocco and Mauritania and Tibesti in Northern Chad) are known 26 genera and 10 families (Chthoniidae - Chthonius, Garypidae – Garypus, Garypinidae - Garypinus, Geogarypidae - Geogarypus, Neobisiidae – Neobisium, Roncus, Olpiidae – Minniza, Olpium, Serianus, Solinus, Atemnidae – Atemnus, Diplotemnus, Chernetidae – Allochernes, Chernes, Dendrochernes, Lamprochernes, Goniochernes, Pselaphochernes, Cheliferidae – Chelifer, Dactylochelifer, Hysterochelifer, Rhacochelifer, Mesochelifer, Lophochernes, Withiidae – Plesiowithius, Withius). General papers of the pseudoscorpions have been published by Callaini on Algeria (1983) and Morocco (1988). Among these genera 21 live also in Europe and 18 - in Africa south of Sahara.

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According to Harvey (online), the number of species known from these countries is: Egypt - 13; Libya - 15; Tunisia - 25; Algeria - 44; Morocco - 38; Mauritania - 2; Tibesti in N. Chad (after Heurtault, 1970 a,b,c, 1971).

Endemic species are: Chthonius kabylicus Callaini, 1983 – Algeria. Neobisium (Blothrus) peyerimhoffi Heurtault, 1990 – Algeria. N. dumitrescoae Heurtault, 1990 – Algeria. Roncus numidicus Callaini, 1983 – Algeria. Roncus (Parablothrus) comasi Mahnert, 1983 – Tunisia. Roncus (Parablothrus) gardinii Heurtault, 1990 – Algeria. Rhacochelifer massylicus Callaini, 1983 – Algeria. Geogarypus mirei Heurtault, 1970 – Chad (Borcou). Goniochernes vachoni Heurtault, 1970 – Tibesti.

Endemic genus for this (mostly desert) area is only Plesiowithius Vachon, 1954 – Mauritania (1 sp.)(Withiidae) In tropical Africa (the described area) have been recorded pseudoscorpions of genera and 16 families: Chthoniidae, Tridenchthoniidae, Feaellidae, Garypidae, Geogarypidae, Olpiidae, Ideoroncidae, Neobisiidae, Syarinidae, Cheiridiidae, Pseudochiridiidae, Sternophoridae, Chernetidae, Cheliferidae, Atemnidae, Withiidae. There are no endemic families in tropical Africa.

Endemic genera of pseudoscorpions are: Fam. Chthoniidae Congochthonius Beier, 1959 – DR Congo (1 sp.). Fam. Neobisiidae Afroroncus Mahnert, 1981 – Kenya (2 sp.). Nannoroncus Beier, 1955 – Kenya, Uganda (1 sp.). Fam. Withiidae Aisthetowithius Beier, 1967 – Kenya, Tanzania (1 sp.). Cryptowithius Beier, 1967 – Kenya (1 sp.). Pogonowithius Beier, 1979 – DR Congo (1 sp.). Trichotowithius Beier, 1944 – Ethiopia, Kenya (2 sp.). Fam. Cheliferidae Chamberlinarius Heurtault, 1983 – Ivory Coast (1 sp.). Fam. Atemnidae Synatemnus Beier, 1944 – Tanzania (2 sp.).

Opiliones In Europe (incl. Caucasus) are known Opiliones from ten families: Sironidae - Cyphophthalmus, Cladonychiidae – Holoscotolemon, Phalangodidae – Lola, Ausobskya, Paralola, Scotolemon, Ptychosoma; Travuniidae – Abasola, Arbasus, Buemarinoa, Dinaria, Kratochviliola, Travunia, Trojanella, Nemastomatidae – Carinostoma, Histricostoma, Mitostoma, Nemastoma, Paranemastoma, Trogulidae – Anarthrotarsus, Anelasmocephalus, Calathocratus, Trogulocratus, Trogulus, Dicranolasmatidae - Dicranolasma, Ischyropsalididae – Ischyropsalis, Phalangiidae – Odiellus, Opilio, Phalangium, etc. Sclerosomatidae – Astrobunus, Gyas, Leiobunum, Nelima. In North Africa are registered Opiliones of four families: Phalangiidae – Amilenus, Dasylobus, Dicranopalpus, Egaenus, Lacinius, Lophopilio, Megabunus, Metaphalangium, Metaplatybunus, Mitopus, Odiellus, Oligolophus, Opilio, Phalangium, Platybunus, Rilaena, Zacheus; Sclerosomatidae – Gyas; Trogulidae – Trogulus, Anelasmocephalus, Calathocratus, Trogulocratus; Phalangodidae – Ptychosoma. In Egypt (Cokendolpher, 1990) are known 6 sp. of Opiliones: Trogulidae – Trogulus gypseus; Phalangiidae – Metaphalangium cirtanum, M. orientalis, Phalangium aegyptiacum, Ph. savignyi, Ph. copticum.

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Cyphophthalmi In Europe Cyphophthalmi (Sironidae) are represented mainly in the southern peninsulae (in Iberian Peninsula the genera Iberosiro, Odontosiro, Paramiopsalis, Parasiro; in the Balkan Peninsula – Siro, Cyphophthalmus = ¿Tranteeva). Siro is known also from Turkey, Italy, Poland and Slovakia. The suborder is not known from North Africa.

Tropical Africa Cyphophthalmi From tropical Africa are known three genera: Ogovea Hansen et Soerensen, 1914 from the Gulf of Guinea (Cameroon, Congo, Bioko, 3 sp.), Parogovia Hansen, 1921 – Bioko, Gabon, Sierra Leone (3 sp.), and the aberrant genus Marwe Shear, 1985 from Kenya (1 sp.). The family Ogoveidae (Ogovea Hansen et Soerensen) is endemic for the Gulf of Guinea. Fam. Neogoveidae has one genus in Africa (Parogovia Hansen), the other genera live in South America. Marwe belongs (?) to Sironidae. Otherwise the Cyphophthami of Europe and tropical Africa belong to different families, without transition between them. The European taxa (Temperophthalmi) are closer to the Cyphophthalmi of North America.

Eupnoi According to the Catalogues of Staręga (1984, 1992, with suppl.), in tropical Africa are recorded Eupnoi from two families. Fam. Phalangiidae (Phalangiinae) - Camerobunus Staręga et Snegovaya (1 sp.), Cristina Loman (13 sp.), Rhampsinitus Simon (47 sp.), Guruia Loman, Coptophalangium Staręga, Odontobunus Roewer, Dacnopilio Roewer, Megistobunus Hansen, ?Hindreus Kauri. Fam. Sclerosomatidae Endemic (known only from one country) are the genera: Fam. Phalangiidae Camerobunus Staręga et Snegovaya, 2008 – Cameroon. Coptophalangium Staręga, 1984 – Ethiopia.

Dyspnoi – not recorded.

Laniatores According to the Catalogues of Staręga (1984, 1992, with suppl.), in the described area are recorded Laniatores from 5 families. Fam. Assamiidae – Aberdereca Goodnight et Goodnight (1 sp.), Bambereca Kauri (1), Bwitonatus Roewer (1), Comereca Roewer (1), Ereca Soerensen (23), Erecella Roewer (9), Erecula Roewer (7), Eusidama Roewer (1), Hypoxestus Loman (10), Metereca Roewer (= Leleupereca Roewer), Lygippulus Roewer (5), Metarhagdopygus Roewer, Randilea Roewer (1), Sesostris Soerensen (5), Sesostrellus Roewer (3), Spinixestus Roewer (6) Fam. Pyramidopidae – Guinea-Bissau, Gambia, Equatorial Guinea, DR Congo, Tanzania, Ghana, Togo, Ivory Coast, São Tomé and Príncipe, Fuerteventura (Canary Islands) (ca.40 sp.). Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Erecanana Strand; other genera on the Seychelles and in Mauritius. Fam. Biantidae – Hinzuanius Karsch, Metabiantes Roewer, Monobiantes Lawrence, Proconomma Roewer. Fam. Samoidae – Microconomma Roewer (Cameroon), Tetebius Roewer (Mozambique). Other genera on the Seychelles. Other important papers used in the text: Bivort & Giribet (2010), Blick & Komposh (2004), Chemini (1995), Chevrizov (1979), Delfosse (2004), Delfosse & Iorio (2007), Farzalieva & Esyunin (2000), Giribet (2000), Giribet & Prieto (2003), Goodnight & Goodnight (1959), Hadži (1973), Hansen (1921), Hansen & Soerensen (1914), Kauri (1985), Komposch (2004), Komposch & Gruber (2005), Kury (2003, 2011a, 2011b), Kury & Cokendolpher (2000), Kury & Mendes (2007), Lawrence (1931), Legg (1990), Lotz (2009), Klimeš (2000), Martens (1978), Mheidze (1964), Novak (2004, 2005a), Novak, Delakorda & Novak (2006), Ozimec (2000), Pack-Beresford (1926), Pinto-da-Rocha, Machado & Giribet (eds.)(2007), Prieto (2003, 2008), Rafalski (1960, 1961), Rafalski & Staręga (1997), Rambla (1967), Roewer (1923, 1927, 1951),

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Sankey & Savory (1974), Santos & Prieto (2010), Schönhofer (2013), Sharma, Prieto & Giribet (2011), Shear (1985), Staręga (1976a, 1976b, 1978, 1989, 2000), Staręga & Snegovaya (2008b), Stol (1993, 2007), Szalay (1968), Šilhavý (1956), Telnovs (2002b), Tumšs (1963), Vanhercke (1999), Wijnhoven (2005), Winiarska (2008).

Araneae In Europe (including Madeira and Azorean Islands) until the end of 2011 have been registered 4892 species and subspecies of spiders (van Helsdingen, 2012). They include 4491 ―true‖ indigenous spiders, 99 introduced species and 302 nomina dubia. The category of unique records /endemics contains 2041 species (45.4%). These spiders belong to the families Atypidae, Ctenizidae, Nemesiidae, Filistatidae, Sicaridae, Scytoditidae, Leptonetidae, Pholcidae, Dysderidae, Segestriidae, Oonopidae, Palpimanidae, Mimetidae, Eresidae, Oecobiidae, Uloboridae, Nesticidae, Theridiosomatidae, Anapidae, Mysmenidae, Araneidae, Linyphiidae, Tetragnathidae, Theridiidae, Lycosidae, Agelenidae, Pisauridae, Oxyopidae, Zoropsidae, Cybaeidae, Argyronetidae, Desidae, Hahniidae, Amaurobiidae, Dictynidae, Titanoecidae, Anyphoenidae, Clubionidae, Liocranidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae, Corinnidae, Zodariidae, Prodidomidae, Zoridae, Sparassidae and others. The most numerous families (figures of 2011) are Linyphiidae (1366), Gnaphosidae (480), Salticidae (400), Dysderidae (331), Lycosidae (303), Theridiidae (258), Agelenidae (200), Thomisidae (193), Araneidae (150), Zodariidae (111), Philodromidae (106). The best represented what concerns the endemics are Linyphiidae (534, 42.4%), Agelenidae (93, 50%) and most of all Dysderidae (227, 70.5%). Since 2011 the number of spider species in Europe is more than 4500. The actualized checklist of all Turkish spiders (Bayram, Kunt & Danisman, 2014), on line, contains 1013 sp. of 330 genera and 53 families. The Araneae of Northern Africa, well studied, especially in Morocco, seems very similar in families and genera to the Southeuropean. According to Scharff (1990), in Africa south of Sahara are considered valid 365 species of Linyphiidae of 67 genera. Some other important papers on Araneae, used in this text: Aakra & Hange (2000), Agnarsson (1996), Blick, Bosmans et al. (2004), Benoit (1960), Bosmans (2009), Bosmans & Vanytven (Internet), Buchar & Ružiţka (2002), Cardoso & Crespo (2008), Cardoso & Morano (2010), Denis (1961, 1968), El- Hennawy (1987, 1990, 1992b, 2000, 2002, 2006), Eskov (1994), Gajdos, Svaton & Sloboda (1999), Griswold (1991), Holm (1962), Huber (2012 ), Huber & Warui (2012), Komposch (2011), Koponen (2005), Koponen & Fritzén (2013), Kronestedt (Version 2001), Larsen & Scharff (2003), Le Peru (2007), Marinu & Verneau (2002), Merrett, Locket & Millidge (1985), Merrett & Millidge (1992), Merrett & Murphy (2000), Mikhailov (1997, 1998, 1999, 2000), Miloševiš (2002), Roberts (1995), Thaler (1976, 1980, 1988), Thaler & Buchar (1994, 1996), Nentwig, Blick, Gloor, Hänggi & Kropf (2013), Newlands (1978), Platnick (2014), Proszynski & Staręga (1971), Samu, Szinetar (1999), Scharff (1992), Topcu, Demir & Seyyar (2005), Van Helsdingen (2012), Varol (2003), Vilkas (1992), Weiss & Urak (2000). In southern Africa are recorded spiders of 64 families. Endemic spider supraspecific taxa in Southern Africa: Fam. Chummidae – endemic family – Chumma Jocqué. Fam. Microstigmatidae – Microstigmata Strand (the other genera are known from South America). Fam. Gallienellidae – Austrachelas Lawrence, Drassodella Hewitt. Fam. Orsolabidae – Afrilobus Griswold et Platnick, Azanialobus Griswold et Platnick. Fam. Nemesiidae – Lepthercus Purcell, Pionothele Purcell.

Opilioacarida The small order Opilioacarida (37 sp., Beron, 2014, with suppl.) is represented in Western Palearctic only by two species of genus Opilioacarus With, described from South Europe (Italy, Greece) and Algeria and endemic for Western Palearctic. In the Afrotropical Region are known Adenacarus Van der Hammen (Yemen), Panchaetes Naudo (Angola, Ivory Coast), Phalangiacarus Coineau et Van der Hammen (Gabon) and Salfacarus Van der Hammen (South Africa, Tanzania, Madagascar). The first three genera are endemic for the Afrotropical Region, Salfacarus is shared with Madagascar (Van der Hammen, 1966, 1977, Coineau & Van der Hammen, 1979, Naudo, 1963, Vásquez & Klompen, 2010).

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Holothyrida Holothyrids do not exist in Palearctic Region and on the African continent. They are known from Seychelles and Mascarene Islands (Beron, 2014).

Trombidiformes Prostigmata Fam. Erythraeidae Lomeustium Haitlinger, 2006 – 1 sp. (Togo, Ghana, Benin)(larval, end. genus).

Conclusion 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 factors (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 endemic and relict groups 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.

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