Biogeography and Relationships of Southern African Myrmeleontidae (Insecta: Neuroptera)

Home , Afrotropical realm, Palparinae

-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 subcontinent. A systematic revision of this fauna is in progress. Two important aspects of the research, biogeography 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 myrnieleontids 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 species 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 biogeographical 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 myrmeleontid 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 systematics 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 subfamily 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 apparently influenced by increasing aridity from east to west on the subcontinent and, possibly, 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. Eastern faunal component. This constitutes about 50 % of the total and is dominated by genera and species with a wide distribution in Africa. These taxa extend southwards to the eastern parts of the subregion, along the tropical corridor tapering into the Trans- vaal and Natal. The southern African area covered by this element includes Zimbabwe, Mozambique, northern and eastern Transvaal, eastern Swaziland and north-eastern Natal (Fig. I). Based on present knowledge, the following genera are exclusively eastern in distribution: Aviu, Bunkisus, Bunyutus, Distoleon, Gymnoleon, Hugenomyin, Lnchlathetes, Maula and Tonzatares. Two of these genera, Lnchlathetes and Tomatares do extend into the extreme northern parts of S.W. A. /Namibia and Botswana from central Africa, but are absent from the southern parts of the western region. All these genera are widespread in the Afrotropical Realm and even beyond. Aviu, Banyutus, Gymizoleotz and Mauls occur widely in central Africa, Bankisus extends through Africa into Yemen and Oman in the the Arabian Peninsula (Holzel 1983; Mansell 1985), Luchlathetes and Totnarares occur in the Oriental Region as well, and Holzel (1986) indicates that Distoleon is distributed throughout the Old World, whilst Hagenomyia is also recorded from Madagascar, the Oriental Region, Australia and North America. This eastern faunal element also includes a number of species which occur outside Africa: Banyutus lethulis (Walker), Centroclisis distincta (Rambur) C. felina (Gerstaecker), Creoleon ufricaar~us(Rambur), C. mortifrr (Walker), Cymothales bozcvieri van der Weele, Hagenomyia tristis (Walker), Myrmeleon ohscurus Rambur, M. quinyuemaculatus Hagen and Syngenes longicornis (Rambur) also live in Madagascar, whilst M. ohscurus and C. clistirzcta extend to Mauritius as well and, according to Navis (1921), C. houvieri inhabits in the Comores Islands too. Western faunal component. Secondly. there is a rich so-called arid-adapted element in- habiting the drier western parts of the subregion, characterized by a high degree of ende- mism and cornprising about 42 % of the local fauna. This component occupies approximately the territories of South West AfricaINa~nibia,Botswana, north-western Transvaal, and the western and northern Cape Province (Fig. 1). The following genera are confined to the western parts of the subregion and all but one, (Crarnborr~orphus), arc endemic to southern Africa (Tab. 3): Brcichyplcc.tron, Cupophuaes, Cramhornorplzus, Exuetolron, Furgellu, Golujrus, Isorzemurus, Obus, Pulparidius, Pamexis and an undescribed gcnus in the tribe Palparini. The majority of species which occur in the west are also endemic to that region. Southern African Myrmeleontidae

Fig. I. Southern Africa with major political and provincial boundaries mentioned in the text.

Table 3. Genera of Myrmeleontidae endemic to southern Africa.

Brachyplectron Esben-Petersen Nunnoleon Esben-Petersen Capophanes Banks Obus Navas Exaetoleon Kimmins Palparidius PCringuey Furgellu Mark1 Pumexis Hagen Golafrus Naviis Tricholeon Esben-Peterscn Isonemurus Esben-Petersen Gen. nov. (Palparini)

Some of the western species, for example Palparidius concinnus Pkringuey and Pulpares kalahariensis Stitz have been able to extend their ranges into the eastern regions along an arid zone corresponding approximately with the Limpopo R~vervalley (Fig. 2). Another feature of this component is the number of small genera which arc apparently very localized. Exuetnleon obtabilis (PCringuey) is known by four specimens from the south-western Cape. Brachyplectron capense Esben-Petersen inhabits the coastal belt from Cape Town to Port Elizabeth. Capophanes conspersa Banks is known from two localities in S.W.A./Namibia, whilst the five species of Pamexis seem to be confined to the Cape Province, extending from Worcester in the south to O'Kiep in the north and Graham- stown in the east. It is also in these dry western parts of southern Africa that the Palpari- nae have reached the zenith of their radiation, where the subfamily is represented by no fewer than six genera, four unique to the area. M. W. Mansell

Fig. 2. Southern Africa with average annual rai~ifallpattern in millimetres. Abbreviations: B - Beaufort West; C - Cape Town; D Durban; G - Gaberone; H - Harare; K - Keetmanshoop; MA - Maun; M - Messina;

PE - Port Elizabeth; P - Pretoria; U - Upington; WB - Walvis Bay; W - Windhoek.

General fauna1 component. The third element are those genera and a few species which are widespread, some even occupying the central high-lying areas of southern Africa, including the southern Transvaal, Orange Free State, Lesotho, north-eastern Cape, Natal and Transkei (Fig. l), which have a relatively poor myrmeleontid fauna. The genera in this category include Cenrroclisis, Creoleon, Cuetu, Cymothales, Fudrinu, Mucronemu- rus, Myrmeleon, Nannoleon, Nemoleon, Nesoleon, Neuroleon, pal pare^, Syngenes and Tricholeon. The majority of species within these genera however, are strictly easterly or westerly in distribution. This is particularly true of Palpures, a point elaborated below. The above genera, with the exception of Nannoleon and Tricholeon which are presently only known from this subregion (Tab. 3), all have a wide distribution in the Afrotropical Region and all, except Cyrnothules and Fudrina, occur beyond the limits of the Region. Although representatives of all 14 of these genera occur in both the eastern and western elements, very few species are distributed throughout, the most widespread taxa being: Centroclisis brachygaster (Rambur), C. distinctu (Rambur), Creoleon mort[fer (Walker), Cymothales lllustris (Navas), Myrrneleon ulcestris Banks, M. doralice Banks, M. ob~cu- rus Rambur, Nannoleon nzichuelseni Esben-Petersen, Nesoleon boschimanus (PCringuey) and Neuroleon chlorunthe (Banks).

DISTRIBUTION OF PALPAKINAE

This dominant group epitomizes the easterly and westerly distributional patterns, and not Southern African Myrmeleontidae one palparine species occurs throughout the entire subregion. Twenty genera have been proposed within this great group of ant-lions and of these, approximately 12 are still valid. Nine occur in the Afrotropical Region (exceptions: Dimares Hagen, Tomatarella Kim- ~nins,Pse~ldirnares Kimmins,) and the following seven in southern Africa (exceptions: Stenures Hagen, Echthromyrmex McLachlan): Crambomorphus, Golafrus, hchlathetes, Palpares, Palparidius, Parnexis and Tomatarrs . Three of these, Golafrus, Palparidius Pame.xis, as well as an undescribed genus, are endemic to southern Africa. There are about 55 valid Afrotropical species in the genus Palpcrres and at least 23 of these occur in this subregion, 16 being endemics. Palparine taxa which occur in the east are: kich/r~thete.smoestus (Hagen), Palpares criffer (Burmeister). P. fistivus Gerstaecker, P. inc1enrc.n.s (Walker), P. lentus NavBs, P. nyas- sanus Navas, P. sohrinus PCringuey, P. sparus McLachlan and Tomarures cirrinus (Hagen). The following palparine species belong exclusively to the western fauna1 element and all except two are endemic to the subregion ( Palpares immcwsus McLachlan and P. flavofas- ciarus McLachlan extend into southern Angola): Crambomorphus (two species), Golafius orzeili Peringucy, Palparidius (three species), Pamexis (five species), Palpares annulatus Stitz, P. tumpanai Navlis, P. darnurcnsis McLachlan, P. dubiosus PCringuey, P. elegun- tulus PCringuey, P. jlflavufusciutus, P. grariosus Navas, P. irnrnensus, P. kalahariensis Stitz, P. karrooanu.~PCringuey, Po~arnpoanu~sPCringuey, P. pulchellus Esben-Petersen, P. speciosus (Linnaeus) and three undescribed species, as well as a ncw genus with four undescribed species. The most widespread taxon in southern Africa is P. cafler which extends from the northern Transvaal southwards to the eastern Cape, and is the only palparine that has been recorded froni the high-lying central areas of southern Africa. It is replaced by the closely related P. .specio.su.s in the south-western parts of the Cape Province. The above outline of distribution pattcrns is based on available data and although details will change as distributional data accumulates, the general pattern is substantiated. These distribution patterns arc clearly manifest in the middle portion of the subcontinent, be- tween approximately 19 " and 33 " south latitude, but become less evident in the extreme north and south where rainfall is more evenly spread (Fig. 2).

RELATIONSHIPS OF SOUTHERN AFRICAN MYRMELEONTIDAE

The local fauna, as can be expected, is closely linked to that of tropical Africa. Apart from the endemic southern African genera (Tab. 3), this subcontinent shares 22 other genera with the rest of the Afrotropical Region. Of these, the following six genera are confined to the Afrotropical Region: Avia, Crambomorphus, Cynlothales, Faclrina, Gymnoleoiz and M~lula.In addition, all those mentioned below also occur in this Region. Southern Africa shares the next highest number of genera with the Oriental Region with 12: Ccntroclz,sis, Creoleon, Cuetu, Di~toleon,Hagenomyia, Macronemurus, Myrmeleon, Nesolcorz, ~Vcwroleorz,Pcrlpares, Tomarares (Ghosh 1984) and hchlathetes. The Palaearctic Region follows with 10 common genera, Cmrroclisis, Creoleon, Cuetu, D~.clolco~r,Mc1c.t-o-oizenlurus, Mymleleorl, Nenzoleon, Neurok~orz,Palpurrs and Syng~nes(Hiil- zel 1986). The southern African fauna hears very l~ttleresemblance to that of the Nearctic and Ne- otropical Regions or the other great d~strlbutlonalcentre, the Australian Region It shares only two common genera w~ththe Americas, Mjrrneleorz and Hugenomyza and some dls- tat ~clat~on\hip\through the Acdnthacllsln~and Dendroleontln~ Auqtral~adnd the Afrotrop- ical Region have three genera in common, Distolron, H~igenornyiaand Mynnelcorz. It also shares the Acanthaclisini and Dendroleontini, but no common genera in thesc tribes. Apart from the species shared with sorne Indian Ocean islands and Madagascar (part of the Afrotropical Region), no southern African species is known to occur outside Africa. Southcrn Africa differs significantly from other zoogeographical regions in the cornposi- tion of its major myrmeleontid fauna1 component. In the western hemisphere, the main radiation has been in the Brachynernurini (now in the Myrmccaclurini) and Glenurini (now in Nemoleontini), both former tribes absent fro111 the Afrotropical Region. whilst in Aus- tralia the Dendroleontini have proliferated (New 1985). The Distoleontini (now also in the Nemoleontini) are the largest tribe in the Palacarctic Region, with about 50 % of all ant-lions belonging to this group (Hiilzel 1986). In the Afrotropical Region by contrast, the radiation has taken place in the Palparini and the epicentre of this emergence has been southern Africa. The Nenrnleontini are also well represented in the subregion but seem rather diverse as presently constituted, and may represent more than one lineage. It is noteworthy that no large suprageneric taxon of Myr- meleontidae with a particularly rich radiation is identifiable in the Oricntal Region. This may be be due to insufficient knowledge of that fauna, or possibly that the Region really lacks a dominant group.

EVO1,UTIONAKY TRENDS

On the basis of the above, it can be concluded that myrnieleontids have been evolving along different lincs in the various biogeographical regions and infbrmation on the antiq- uity of some groups can he gleaned. Zoogeographical evidence suggests that, based on present distribution patterns, the Dendroleontini, Acanthaclisini and Myrnieleontini, which are cosinopolitan, are ancient lineages and were present before the fragmentation of Gond- wanaland. This implies that the ubiquitous genus Myrzrlcon is very old, and that the pit- building habit was an early innovation which led to a succcsful and widespread radiation of this branch of the family. Of the aboveinentioned three tribes. the Dendroleontini have undergone their greatest ratli- ation in Australia (New 1985) which has been isolated for longer than the other zoogeographical regions. The arboreal habits of sorne taxa nlay also lend support to the suggestion that Dendroleontini represent a very ancient myrnmeleontid lineage. Other lines particularly the former tribes Brachynemurini and Glenurini in America and Palparini in Africa have emerged and radiated subsequent to the disintegration of Gond- wanaland. These groups arc dominant in their regions, but the fernier two tlo not occur in Africa and the latter is not represented in America, indicating that the radiations may be comparatively recent. One of the main factors, possibly the key, to the emergence of ant-lions as the largest and most widespread family of Neuroptera has been the exploitation of the sand environ- ment. This is a virtually limitless habitat and its colonization held many advantages for neuropterous insects. For example, protection from predators, caniouflage from prey, an aid to rapid subdual of prey by dragging it under the sand, thermoregulation through ver-- tical migration in the substrate, material for pupal construction and others. Also, the basic neuropreran feeding mechanism of the niandibuiariniaxillar piercing and sucking mouth- parts and external digestive process was an ideal preadaptation for predatory larvae living in sand (Mansell 1986). The protection afforded by the sandy habitat may also have contributed d~rectlyto the large size attained by Pulpc~resspecies. The larvae are not restricted by habitat size (eg. Southern African Myrmeleontidae tree-holes or rock ledges) or confined by small inconspicuous pits or limited in sizz when living exposed on vegetation or rocks. With these limitations removed, some palparines have developed into the largest Neuroptera under the protective mantle of sand. In so do- ing, they have also been able exploit a wider range of prey and prey sizes than other ncu- ropterans. The various desert and semi-desert biotopes of southern Africa have therefore provided ideal conditions for the radiation of Myrmeleontidae and, in particular, the great recent group, the tribe Palparini.

ACKNOWLEDGEMENTS

I am indebted to my esteemed friend and colleague, Prof. Dr. Horst Aapbck (University of Vienna). for his constructive criticism of the manuscript and his valuable suggestions for its irnproven~ent.

REFERENCES

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Bibliography of the Neuropterida Reference number (r#): 6995

Reference Citation: Mansell, M. W. 1990 [1990.??.??]. Biogeography and relationships of southern African Myrmeleontidae (Insecta: Neuroptera). Pp. 181-190 in Mansell, M. W.; Aspöck, H. (eds.). Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology (3-4 February 1988, Berg en Dal, Kruger National Park, South Africa). South African Department of Agricultural Development, Pretoria. 298 pp.

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