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Proceedings of the Tenth International Symposium on Neuropterology. Piran, Slovenia, 2008. Devetak, D., Lipovšek, S. & Arnett, A.E. (eds). Maribor, Slovenia, 2010. Pp. 257–266.

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Patterns of distribution of the (: Myrmeleontidae: ) in the northern half of Africa. Faunal transitions and regional overlaps

André Prost1 1F- 39320 Loisia, France ; E-mail: [email protected]

Abstract. The Afrotropical Region encompasses the African continent south of latitude 16°N. This definition leaves most of the Sahara within the Palaearctic Region, together with the Mediterranean basin. This paper describes the geographical distribution of Palparini in the northern part of the Afrotropical Region, i.e. between the equator and 16°N. Distribution maps are presented. The presence of 25 species in the region is confirmed. A new synonymy tigris (Dalman, 1823) = Nosa adspersa Navás, 1914 is proposed. The striking feature is the apparent absence of species that extend over the entire Afrotropical domain. The equatorial forest is a biological barrier. In contrast to this situation, four species that belong to the Eastern Mediterranean arid ecosystems penetrate deeply into this region of Africa. As a consequence of the ecological barriers of the region, with a desert to the north and an equatorial forest belt in the south, a high degree of endemicity in Neuroptera has developed. Three species are recorded uniquely from Ethiopia/Somalia, while P. zebroides is known exclusively from desert locations in Chad and Niger; and all except one specimen of S. arenosus originate from the Bamako district, Mali. Other species seem to depend on the degree of aridity/humidity of the African savannah, where the majority are distributed along elongated narrow bands from the Atlantic Ocean to the Nile River, with very limited northern or southern expansion. The fragmentation of the African rain forest into a western bloc (Sierra Leone to Ivory Coast) and an equatorial bloc (Cameroon to Congo) has isolated Neuroptera populations but was not conducive to the differentiation of endemic characteristics, as was the case for other groups.

Key words: Myrmeleontidae, Palparinae, Palparini, , Africa

Background

In 1995, a preliminary revision of the Palparini of West Africa set the total number of species at 18, based upon examination of about 450 specimens deposited in European museums and private collections (Prost, 1995). Later, further information was added by Michel (1999) and Whittington (2002), the genus Palparellus was revised by Mansell (1996), a catalogue of the west Palaearctic fauna was published by Aspöck et al. (2001), a synopsis of species of the World was published on the internet (Oswald, 2008), and nomenclatural changes were introduced by the “Catalog of World’s Antlions” (Stange, 2004). I have further examined collections in the National Museums of Scotland in Edinburgh (courtesy of Andrew Whittington), in the Russian Academy of Sciences in St Petersburg (courtesy of Viktor Krivokhatsky), and in Cornell University, Ithaca, New York (courtesy of Richard Hoebeke).

The scope of the 1995 revision was limited to West Africa, which is not a meaningful concept in zoogeography. As pointed out by Aspöck et al. (2001), the equatorial forest is a barrier that divides the continent into two unconnected savannah areas. The only communication is through the East African corridor along the Rift Valley, a narrow path with so many geological and climatological features that it is hardly a route for dispersal. Therefore, the present paper considers the distribution of Palparini in Africa north of the equatorial divide, from 0° to 16°N, which is the conventional limit between the Afrotropical and the Palaearctic zoological regions.

ISBN 978-961-6657-16-7 © 2010 Faculty of Natural Sciences and Mathematics UM

Taxonomy

A major clarification effort was made by Stange (2004) in the “Catalog of World’s Antlions”. However, a few conclusions regarding Palparini are not substantiated in that work.

For example, the name Nosa tigris established by Navás (1912a) is not a new combination, nor can the merger already established by Banks (1913) of the subspecies N. tristis niansanus and N. tristis brevifasciatus into Nosa tristis be considered new synonyms. Even more difficult is the fact that Stange accepts the synonyms Nosa tigris = N. leonina and digitatus = P. pobeguini on the basis of old literature. The revision of West African taxa (Prost, 1995) had clearly established that major differences in male genitalia lead to the recognition of the four taxa as valid species, and allowed the conclusion that synonymies should be revoked.

The case of Palpares percheronii/tessellatus deserves a short discussion. The revision (Prost, 1995) considered that the name tessellatus (Rambur, 1842) had priority over the name percheronii introduced by Guérin- Méneville in a publication issued as successive folders between 1829 and 1844. An analysis of the situation by Cowan (1971) concluded indeed that the text had been issued in one block in September 1844 and that “it is quite certain that valid publication under the International Code of Nomenclature did not take place until August or September 1844”. Insect plates of course had been circulated since 1828, especially to accompany the second edition of Cuvier’s “Règne ” which completely lacked illustrations. Unfortunately, only 44 of the 110 insect plates can be assigned a date, none of which include Myrmeleo percheronii. Meanwhile, Cuvier’s “Animal Kingdom” was in high demand in English-speaking countries: seven illustrated English editions were provided to the public between 1824 and 1840, five in London, one in Edinburgh, and one in New York. Stange (2004) correctly made the decision to recognize Griffith’s 1831 publication in New York of the “Animal Kingdom” as the first valid publication and illustration of the name M. percheronii. This opinion is definitive, unless it is established that Griffith’s London edition issued in 16 volumes from 1824 to 1835 takes precedence, or unless an earlier date can be assigned to a plate issued in France. Lastly, Guérin changed his name to Guérin-Méneville for obscure reasons in 1836 (Cowan, 1971). Therefore, the correct citation for the species should be Palpares percheronii (Guérin, 1831) in Cuvier, G., “Animal Kingdom”, New York: Griffith, 1831.

A minor point refers to Tomatares clavicornis which Stange attributes to “Latreille, 1817”. Latreille’s volume on was published as volume five of the second edition of Cuvier’s “Règne animal”, with the date 1829 printed on the front page. The first edition of Cuvier’s treaty in 1817 did not cover insects. Thus, Tomatares clavicornis (Latreille, 1829) is the correct citation for the species.

List of species and related distribution

A few published records are based upon erroneous identification, and the corresponding species are consequently not part of the fauna of Africa north of the equator. These are Palpares cataractae (Henwood’s 1977 specimen from Nigeria was revised as P. cephalotes); Palpares aegrotus (Fraser’s 1954 specimen from Guinea is P. obsoletus); and Palpares torridus misidentified from Ghana. In addition, Fraser (1950) mistakenly used the name Palpares libelluloides for specimens of P. latipennis collected in northern Niger. Examination of male genitalia has ascertained proper identification (Prost, 1995). Unfortunately, Stange’s catalogue reiterates the presence in Niger of a species which is strictly Palaearctic.

Twenty-five species are present in the sub-region. References in the following list will be given only for additions or corrections to geographical information summarized in the “Catalog of World Antlions” (Stange, 2004). Distribution maps are presented by means of Geographical Information System software and based on published valid records for each species. Multiple records within a zone appear as filled areas.

Goniocercus klugi (Kolbe, 1898): Arabian Peninsula (Saudi Arabia, Oman, Yemen) and Egypt, Ethiopia, So- malia, Sudan, Mauritania; also in Algeria and in Niger (Whittington, 2002), and in Chad (Navás, 1931) (Fig. 1).

Goniocercus walkeri (McLachlan, 1894): Iran, Israel, Saudi Arabia, Yemen, Ethiopia, Sudan, Kenya.

Lachlathetes furfuraceus (Rambur, 1842) (Fig. 2): Burkina Faso, Ghana, Mali, Niger, Nigeria, Senegal, and also Benin (Prost, 1995). The specimen from Ituri, Congo, which I examined in the Tervuren Museum (MRAC), is not P. furfuraceus (Prost, 1995) and the identification of a specimen collected on Lake Mweru, in the Katanga province of Congo (Navás, 1932) is doubtful. Both could belong to Palpares abyssinicus which Banks (1920) reported from Eastern Congo.

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Fig. 1. Distribution of Goniocercus klugi:  – single findings; dark area: multiple records.

Fig. 2. Distribution of Lachlathetes furfuraceus (black) and Lachlathetes gigas (grey).

Fig. 3. Distribution of Nosa tigris and Nosa leonina. Nosa tigris:  – a single finding; grey area: multiple records; Nosa leonina:  – single findings; striped area: multiple records.

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Lachlathetes gigas (Dalman, 1823): Sierra Leone, Guinea (Fig. 2). The presence of the species in Gabon (Navás, 1926) remains hypothetical as long as the specimen of the entomological institute in Berlin is unique. Moreover, it belongs to the collection Le Moult made of insects purchased from various collectors and a mistake in the place of origin cannot be ruled out.

Nosa tigris (Dalman, 1823): Benin, Burkina Faso, Cameroon, Central African Republic, Chad, Congo- Brazzaville, Ghana, Guinea, Ivory Coast, Liberia, Mali, Niger, Nigeria, Senegal, Sierra Leone; also in Togo (Kolbe, 1898; Stitz, 1912), in Eritrea (Esben-Petersen, 1916), in Ethiopia (Kolbe, 1898 as P. hamatus; Navás, 1914 as Nosa adspersa, NEW SYNONYM) and in Gabon (Navás, 1926) (Fig. 3). In addition, I identified a female from Sudan [Bahr el Jebel 5°11N, 3 Nov. 1948, JC Bradley] and a female from Uganda [Kabuwangasi, 18 Nov 1948, JC Bradley] both in the Entomological Department, Cornell University, Ithaca, New York.

Nosa leonina Navás, 1911 is a valid species, with male genitalia that differ significantly from N. tigris (Prost, 1995). The MRAC has numerous specimens from Congo from which it was described (Navás, 1911a). It was further recognized in eastern Central African Republic (Navás, 1926). I examined specimens from Lago enclave, southern Sudan (MRAC), and a couple from Zimbabwe in the collection of the National Museums of Scotland (published as N. tigris by Whittington, 2002). Known distribution is shown in Fig. 3.

Palparellus spectrum (Rambur, 1842): Benin, Burkina Faso, Chad, Congo-Brazzaville, Ivory Coast, Mali, Niger, Nigeria, Senegal, Togo; also in Guinea (Navás, 1912b) and in Sudan (Whittington, 2002) (Fig. 4).

Palpares abyssinicus Kolbe, 1898: Ethiopia, Congo-Kinshasa; also in Eritrea (Banks, 1920). This species is the alternate of the closely related Lachlathetes furfuraceus in the eastern part of the region. It should probably be transferred to the genus Lachlathetes.

Palpares cephalotes (Klug, 1834) : Afghanistan, Pakistan, Iran, Israel, Egypt, Libya, Tunisia, Saudi Arabia, and Oman in the Palaearctic Region ; Chad, Ethiopia, Mauritania, Niger, Senegal, Sudan ; also in Nigeria (Prost, 1995) and in Djibouti (Navás, 1911b) (Fig. 5).

Palpares digitatus Gerstaecker, 1893: Congo-Kinshasa, Guinea, Ivory Coast, Kenya; also in Ghana (Gerstaecker, 1893), southern Somalia (Banks, 1913), and Nigeria (Whittington, 2002) (Fig. 6).

Palpares incommodus (Walker, 1853): Burkina Faso, Gambia, Guinea, Mali, Nigeria, Senegal; also in Sudan (Navás, 1912c as P. costatus) (Fig. 7).

Palpares nigrescens Navás, 1913: Burkina Faso, Central African Republic, Congo-Brazzaville, Congo- Kinshasa, Ivory Coast, Mali, Senegal; I examined an additional specimen from Nigeria [Prov. Bornu, Biu division, 3 Apr 1973, K. Proud] in the collection of the National Museums of Scotland (Fig. 8).

Palpares obsoletus Gerstaecker, 1898: Angola, Benin, Congo-Brazzaville, Congo-Kinshasa, Guinea, Ivory Coast, Kenya, Mali, Namibia, Niger, Tanzania, Zambia; also in Nigeria (Navás, 1912b; Whittington, 2002 as P. longicornis), Zimbabwe (Whittington, 2002) and Ghana (Whittington, 2002 as P. berlandi) (Fig. 8).

Palpares papilionoides (Klug, 1834): Saudi Arabia, Ethiopia, Kenya, Senegal, Somalia, Sudan, Uganda, Tanzania; also in Yemen and Niger (Whittington, 2002); also I received a specimen for identification from Cameroon [Prof. Ngamo, Ngaoundere, 2007] (Fig. 9). Israel and Turkey are quoted by Stange (2004), but not by Aspöck et al. (2001) which makes the localities dubious.

Palpares percheronii (Guérin, 1831): Chad, Mali, Mauritania, Niger, Senegal; also in southern Algeria and Sudan (Whittington, 2002) (Fig. 10).

Palpares radiatus Rambur, 1842: Burkina Faso, Chad, Mali, Mauritania, Niger, Senegal; also in Nigeria (Prost, 1995) (Fig. 7). Ivory Coast mentioned by Stange with a reference to Prost (1995) is misquoted as I concluded in that paper that Ivory Coast was not a reliable record for the species.

Palpares umbrosus Kolbe, 1898: Ivory Coast, Guinea, Mali, Togo; also in Ghana (Prost, 1995; Whittington, 2002 as P. berlandi) (Fig. 6).

Palpares zebroides Fraser, 1950: Chad, Niger (Fig. 10).

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Fig. 4. Distribution of Palparellus spectrum ().

Fig. 5. Distribution of Palpares cephalotes:  – single findings; grey area: multiple records.

Fig. 6. Distribution of Palpares digitatus () and Palpares umbrosus ().

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Fig. 7. Distribution of Palpares radiatus () and Palpares incommodus ().

Fig. 8. Distribution of Palpares nigrescens and Palpares obsoletus. Palpares nigrescens:  – single findings; Palpares obsoletus: grey area: multiple records.

Fig. 9. Distribution of Palpares papilionoides:  – single findings, grey area: multiple records.

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Fig. 10. Distribution of Palpares dispar , Palpares zebroides and Palpares percheronii. Palpares dispar:  – a single finding, black area: multiple records; Palpares zebroides:  – single findings; Palpares percheronii: grey area: multiple records.

Fig. 11. Distribution of Parapalpares latipennis: grey area: multiple records.

Fig. 12. Distribution of Stenares arenosus () and Stenares hyaena ().

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Parapalpares latipennis (Rambur, 1842) (Fig. 11): Burkina Faso, Mauritania, Mali, Niger, Nigeria, Senegal, Sierra Leone and Sudan (Whittington, 2002); the presence of specimens that conform to the type has not been confirmed in any other location. When he cites records from Angola and Congo, Stange unfortunately perpetuates a confusion which I had attempted to sort out with the suggestion to assign specimens from central and southern Africa to different species (Prost, 1995). Hagen (1887) and Navás (1912a) had already expressed doubts about the identity of western and southern African populations. Handschin & Markl (1955) are in agreement when they consider the central and southern forms as a homogeneous population which differs from those of West Africa. They proposed to retain the name latipennis for this homogeneous group and to rename the West African individuals. This opinion is not congruent with the fact that the type specimens of Rambur are from Senegal. Three are still preserved: one in Paris (MNHN) and two in Brussels (IRSN). Therefore, the name P. latipennis should apply to the West African population; central and southern specimens belong to other species (among them Palpares praetor).

Parapalpares somalicus (Insom & Carfi, 1988): Somalia

Stenares arenosus Navás, 1924: Burkina Faso, Mali (Fig. 12). The quotation of Senegal by Stange is a misreading of Navás (1924), who located a specimen at “Koulouba, Haut Senegal & Niger”. Koulouba is part of the modern town of Bamako, in Mali, and was at that time the capital city of a military territory, “Haut Sénégal et Niger”, which included the French possessions in inner West Africa. Also, Stange’s mention of Sudan, with no reference, is a misunderstanding for “Soudan”, the former name of present Mali. The validity of S. arenosus as a species distinct from S. hyaena is questionable. The differences observed in male genitalia (Prost, 1995) were not confirmed by Michel (1999). Wing patterns show multiple colour grades, with the darker forms identified as S. hyaena in forest areas. It is possible that S. arenosus is a paler form of the same species that occurs in dry savannah. Nor was the re-examination of Dalman’s type of S. hyaena in Stockholm conclusive, the specimen being of intermediate coloration.

Stenares hyaena (Dalman, 1823): Central African Republic, Congo, Ivory Coast, Sierra Leone; also present in Guinea (Prost, 1995, with additional specimens in the collection of the Russian Academy of Sciences, St Petersburg), in Nigeria (Prost, 1995; Whittington, 2002) and in Sudan (Esben-Petersen, 1915) (Fig. 12). Individuals from India and Pakistan belong to another species.

Stenares completus Banks, 1915: Ethiopia.

Tomatares clavicornis (Latreille, 1829): Burkina Faso, Chad, Mali, Mauritania, Niger, Senegal; also in Nigeria (Whittington, 2002) and in Sudan [Prov. Kordofan, one specimen examined in Edinburgh, National Museums of Scotland].

Tomatares striolatus (Stitz, 1912): Ethiopia.

Palpares dispar Navás, 1912 (Fig. 10): A record of Palpares angustus McLachlan was published from Mali (Whittington, 2002). I examined the two males in Edinburgh, the wing patterns and the genitalia of which are similar to those of Palpares dispar. They were labelled P. oranensis by Waterston. The location of the catches at the northern extremity of Mali around 23°N places it outside the Afrotropical region. The record of P. dispar from Eritrea may be the result of an occasional dispersion from Yemen; and the synonym species Palpares gestroi was recorded from Libya only. Therefore I take the conservative position not to include P. dispar in the Afrotropical fauna at this stage.

Distribution patterns

A single species, Palpares obsoletus, is present in the entire Afrotropical region, from Mozambique and Namibia westward to Guinea. It confirms that the equatorial forest ecosystem is a biological barrier for most Neuroptera which prefer drier environments and are more diversified in savannah regions.

The Sahara is also a biological frontier: there is no evidence that any Western Mediterranean species can reach the southern limit of the desert. By contrast, four species which Aspöck et al. (2001) qualify as “Eremiales Faunenelements”, and which inhabit dry areas of the Middle East (Iran, Syria, Arabian Peninsula), penetrate the eastern part of the African region and can expand westward under favourable conditions. The Red Sea and the Nile Valley are breaches in the desert frontier. Palpares cephalotes is the most characteristic case: the species, centred in Egypt, and the Arabian Peninsula, from where most of the specimens originate, has expanded to Iran, Afghanistan and Pakistan, and it has firmly established itself in a narrow band known as the Sahel zone of Africa

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from Sudan to Dakar, where annual rainfall is below 300 millimetres. The same applies to Goniocercus klugi described from Sudan, which occasional records show can reach Mauritania under similar climatic conditions. Palpares papilionoides described from “Arabia” is relatively common down to Zanzibar and recent records show that it can be occasionally captured in northern Cameroon and Niger. I have not been able to identify which publication could justify the claim that it reaches Senegal; therefore the locality should not be retained. Finally Goniocercus walkeri described from Aden is well established from Sudan to Kenya.

The division of the African rain forest into a western bloc, Liberia and Sierra Leone, and a central equatorial bloc, from Cameroon to Congo, separated by a wide savannah gap, has induced in many insect groups a drift towards individual speciation. Insects in the western bloc tend to develop a high degree of endemicity. This is not the case in Palparini. Two species characterize the western rain forest, Lachlathetes gigas and Stenares hyaena, together with the more opportunistic Palpares digitatus. All three are also present in the equatorial forest. Two closely related species alternate with P. digitatus in slightly modified ecological conditions: Palpares umbrosus in the humid savannah of West Africa, and P. pobeguini in the rain forest of the Congo River basin. The three taxa constitute together a group of sister species.

Ten species represent the core group of the savannah Palparini. A single one, Nosa tigris, is common across the region. Others seem to have strict hygrometric requirements. Their distribution is shaped into narrow bands from the Atlantic Ocean to the Ethiopian plateau or even the Indian Ocean. By contrast, the north-south range does not exceed a few hundred kilometres. This pattern coincides with the meteorological maps of annual rainfall: the summer monsoon comes from the south and rapidly loses strength over the continent; within a distance of 1,500 km, annual rainfall drops from over 6 metres in Liberia to 100 millimetres in Mauritania.

Palpares umbrosus is the more hydrophilic species, limited to areas with over 1,200 mm annual rainfall; P. nigrescens, P. incommodus, L. furfuraceus, P. latipennis, and P. spectrum inhabit regions which receive annually between 800 and 1,100 mm, with a tolerance for more humid environments in the case of furfuraceus, and for much drier conditions in latipennis. The climatic zone around 700–900 mm is home to P. radiatus and T. clavicornis. The characteristic species of the Sahel is P. percheronii which is not present when rainfall exceeds 500 mm.

Three Palparini appear, from available data, to be narrowly localized. All specimens except one of Stenares arenosus come from the district of Bamako, Mali. The two known specimens of P. zebroides originate from oases in Air Mountains, Niger (Fraser, 1950) and the Tibesti Mountains, Chad (Prost, 1995), 1,200 km distant. Nosa leonina is so far confined to north- eastern Congo, with expansions into nearby areas of the Central African Republic and Sudan; it is likely to be present all along the western slope of the African Rift as indicated by the two specimens from Zimbabwe in Edinburgh.

Finally, three species have not been recorded outside Ethiopia (S. completus, T. striolatus), or Somalia (P. somalicus), a geographical area with specific conditions that may justify the isolation of insect populations.

Acknowledgements. The distribution maps were prepared by Charlotte Roudier-Daval, Department of Geography, University Paris-Nanterre, to whom I wish to express my grateful thanks for her contribution.

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Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 13655

Reference Citation: Prost, A. 2010 [2010.11.25]. Patterns of distribution of the Palparini (Neuroptera: Myrmeleontidae: Palparinae) in the northern half of Africa: faunal transitions and regional overlaps. Pp. 257-266 Devetak, D.; Lipovšek, S.; Arnett, A. E. (eds.). Proceedings of the 10th International Symposium on Neuropterology (22-25 June 2008, Piran, Slovenia). University of Maribor, Maribor, Slovenia. 307 pp.

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