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-p- -p-p p- Pp - p -p p Advan~esIn Neuropterology Proceedlng5 of the Th~rdInterndtlonal Sympos~umon Neuropterology Berg en Dal, Kruger Ndtlondl Park, R S A 1988 Mansell, M W & Aspcl\, H (Eds) Pretoria, R S A 1990 4, 181-190 . p-p p-p -- -p - -P-P Biogeography and relationships of southern African Myrmeleontidae (Insecta: Neuroptera) Mervyn W. MANSELL National Collection of Insects, Pretoria, South Afrlca ABSTRACT Southern Africa has a rich fi1~1naof Myrmeleontidae: about 170 species in 40 genera are estimated for the sub- continent. A systematic revision of this fauna is in progress. Two important aspects of the research, biogeogra- phy and the relationships of southern African ant-lions with those in other regions, are outlined. Myrmeleontidae manifest three main distributional trends. correlating with rainfall patterns, within the subregion. Local myr- nieleontids form a major component of the Afrotropical fauna and also have generic affiliations with those of the Oriental and Palaearctic Regions. Key words: Myrmeleontidae. distribution, biogeography, relationships, systematics, evolution, southern Afri- ca, Afrotropical. INTRODUCTION Southern Africa, that part of the Afrotropical Region to the south of the Cunene, Kavango and Zambezi Rivers (Fig. l), has a rich fauna of Myrmeleontidae. Approximately 150 species in 45 genera have been recorded from this subregion. Many are synonyms but present research indicates that they will be offset by discovery of new taxa, and that the actual figure is probably about 170 species in 40 genera. This compares favourably with the rich Australian fauna, where New (1985) enumerated 201 species comprising 42 genera, from an area approxin~atelytwo and one third times larger than this subcontinent. A list of genera, with approximate numbers of authentic described and undescribed spe- cies presently known from southern Africa, is presented in Table 1. Although the higher classification of Myrmeleontidae is still tenuous (see below), the listed genera are arranged into subfamilies and tribes according to Stange & Miller (1990). A key to these genera was provided by Manscll (1985). A comprehensive revision of this fauna is in progress and two important aspects of the research, the biogeography and relationships of the southern African Myrmeleontidae are outlined here with emphasis on thc dominant group, the subfamily Palparinae. Detailed distribution records will be provided in forthcoming revisions of the various taxa, the ob- ject of this contribution being to summarize available data and to illustrate biogeographi- cal trends. It also complcrncnts the contributions of Hiilzcl (1986) and New (1986) on the Palaearctic and Australian Myrmeleontidae. The two topics are introduced against a brief historical background and its implications for the present studies. and the paper is concluded with discussion of aspects of myrmeleon- tid evolution relevant to the Palparinae. Table 1. Genera of southern African Myrmeleontidae with approximate numbers of known and undescribed species. (The itatiI\ of the last eight genera I\ uncertain: most are synonyms). Subfamily MYRMELEONTINAE Nur~r~olc~onEsb.-Pcterien (l) Tribc ACANTHACLISINI Ncrnol~onNavk (5 +4) Al9i~rNavis (l) Ncurolron Navas (8) Controc.1isi.s Navis (8 + 1) Obus Navas (1 +2) Futlr.irltr Navas (1 +2) Subfam~lyPALPARINAE Syn~cncsKolbe (2) Trlbe MAULINI Tribe DENDROLEONTINI I\orlcr?luru\ E\ben-Petcr\en (1 +2) Btrt1ki.su.s Nav5s (2 + l ) Muulu Navac (l + l) Cyr11oth~11c.sGerstaeckcr (1 1) Trlbc PALPARINI 7i-icho1c.o~Esb.-Pet. (2 + 1 ) Cr~rrnhornorphu.\McLachlan (1 + l) Tribe MYRMECAELURINI Golufrus Navjs ( 1 ) Cuetcl Navas (6) Lrlthlurl~et~~McL'ichlan (1) Furg~I1(1Markl (I + 1) Prllptrre~ Rdrnbur (23 +2) Nrsol~onBanks (3) Pr;llpc~nd~u\Per~ngucy (3) Tribe MYRME12EONTINI P(rrne\~~Hagen (3 + 2) H~rgt~nott~yitrBanks (2) Torr~utnrc~\Hagen (1) Myrrnclcorz Linnaeus ( 13 + 3) Grri riov ( f4) Tribe NEMOLEONTINI Bunyutus Navlis (4 +2) Borzluc Ndvis (1) Bmt~lz~plcc~ironEsben-Petersen (I) C~rrlpcstr~tuNavii (1) C~rr~ophutzcsBanks ( 1 ) Gurzrlulur Navfii (2) Creol~onTillyard (8) Lrdruc Navai (1) E,ru~tolconKimmins ( 1 ) Mocl~u,~Navlii (1) Distolcorl Tillyard (4 + l) Nc1grc~lu.r Navk Gynznoleorz Banks ( 1 ) Notm Navii Mtic.roti~murusCosta (7) Sutrr Navh (1) HISTORICAL RESUME AND IMPLICATIONS Pulpclrp.s spcciosus was the first myrmcleontid to be docunlented from southern Africa, when it was described by Linnaeus in 1758. Subsequent to Linnaeus, 17 authors (listed in Tab. 2) have enhanced and, in some cases. confounded our knowledge of the local Myr- melcontidae. References, and details of their contributions are provided by Mansell (1985). Table 2. Historical review: authors who have contributed to the systematics of southern African Myrmeleontidae Linnaeu5 1758 Kolbe 1898 Thunberg 1784 van der Weele 1903, 1908 Olikier 1809, 181 1 Peringuey 19 10, 191 1 Bur~neister1842 Banks 1909- 1941 Rambur 1842 St~tz1912 Walker 1853, 1860 Esben-Peterien 19 12- 193 1 Hagen 1860 - 1887 Navas 1909- 1937 Gerstaecker 1863- 1894 Kimmlns 1943, 1948 McLachlan 1867 Markl 1953, 1954 Southern African Myrmeleontidae In the 23 1 years since Linnaeus' work, the only resident taxonomist to study local myr- rneleontids was PCringuey (1910, 19 1 1). All other taxonomic studies on Afrotropical Myr- meleontidae were carried out by systematists outside Africa and this has several important implications for present-day research on ant-lions. (1) All type-material, with the exception of that of PCringuey, is housed in overseas muse- ums. Much of it is very old or has been lost or destroyed, especially that of Navis, or is inaccessible to local taxonomists for political reasons. This is a major co~nplicatingfactor. (2) Many of the taxa described by thc above authors are from outside southern Africa, and have subsequeritly been recorded here. This implies that a large proportion of the local genera and species are widespread, necessitating a detailed knowledge of the whole Afrotropical fauna. (3) A large number of genera and species were inadequately described, and there are many synonyms amongst the plethora of available names for Afrotropical Myrmeleontidae. In the abscnce of type specimens, in rnany cases, and poor descriptions, the elucidation of these names and associated taxa is a daunting task. (4) As earlier workers had limited material, none of their accounts is comprehen\ive. No revision of the local fauna, or that of any other Afrotropical component, had hitherto been attempted. There is thus no convenient starting point as is the case in most other families, where the fundamental works of Tjeder (1 957- 1967) allow easy access to the group. (5) The fifth important implication is that all the earlier research was bared on preserved adult material. Nothing was known about the immature stages or biology of the local taxa. PRESENT STUDIES It was the latter two aspects especially, that stimulated the current studies on Myrmeleon- tidae by the author about 20 years ago. Initially research centred on biology, particularly the discovery and identification of larvae, but it quickly becarnc apparcnt that the sys- tematics of Afrotropical Myrmeleontidae, at all taxonomic levels, was in disarray and in urgent need of attention. A comprehensive systematic revision of the Afrotropical fauna, with enlphasis on the southern African taxa, thus became the main priority. It was decid- ed to approach these systematic studies from the species level, primarily to discover and identify the various taxa, to investigate their biology, and to use this essential information to elucidate the higher classification of Myrmeleontidae. It was deemed premature at that stage to attempt a supra-generic arrangement until the majority of species had been adequately studied, an approach vindicated by the persistent entropic state of higher classification of the group, even at subfamily level (Hiilzel 1986). The supra-generic categories thus referred to in this work follow Stange & Miller (1990), but still require clear phylogenctic sustantiation. During the course of this research, extensive field studies have been carried out in all the provinces and major biotopes of South Africa and South West AfricaINamibia, as well as Botswana, Zimbabwe, Mozambique and Malawi. A collection of over 10.000 Myr- meleontidae, in addition to other fa~niliesof Neuroptera, has been compiled and curated in the National Collection of Insects in Pretoria, and is the nlost comprehensive available for this region. Larvae of approximately one third of the known specics have also been discovered and reared. All the relevant literature has been obtained and catalogued, including the compilation of pertinent taxonomic indices. The author is in regular contact with neuropterological colleagues around the world, and most of the leading specialists have undertaken field M. W. Mansell work and collaboration with hi~nin southern Africa. Much comparative material from other regions has been exchanged, especially from America. In the process, a large amount of unpublished data on adults, larvae and biology of the Myrmeleontidae has accumulated and is presently being prepared for publication. In this regard, revisions of the three genera comprising the tribe Dendroleontini in southern Africa have already appeared (Mansell 1985, 1987, 1988). Attention is now focussed on the sub- family Palparinae which is richly represented in this region (see below). The author has much information on thls group, including long series of most Afrotropical taxa, some of which have been correlated with their larvae. The first two papers, one describing a new genus compr~singfour new ~peciesand the other rev~singPamexis, are almost com- plete. They will be followed by revisions of all the other palparine genera in turn. BIOGEOGRAPHY OF SOUTHERN AFRICAN MYRMELEONTIDAE Southern African Myrmeleontidae manifest three main distributional trends which are ap- parently influenced by increasing aridity from east to west on the subcontinent and, possi- bly, by altitude in the centre of the area (Fig. 2). There is (1) an eastern faunal component, (2) a western component and (3) a more widespread or general faunal element.

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