The Berothidae of Africa: a Review of Present Knowledge (Insecta: Neuroptera)

Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology. Berg en Dal, Kruger National Park, R.S.A., 1988. Mansell, M.W. & Aspdck, H. (Eds). Pretoria, R.S.A. 1990. F'p. 101-1 13.

The Berothidae of Africa: a review of present knowledge (Insecta: Neuroptera)

Ulrike ASPOCK Naturhistorisches Museum, Wien, Austria

ABSTRACT

Three' of the four berothid subfamilies are represented in Africa: Rhachiberothinae Tjeder, with Rhachiberorha Tjeder (two species) and Mucroberotha Tjeder (three species); Nosybinae MacLeod & Adams, with Nosybus Navis (seven species); Berothinae Handlirsch, with Nodalla Navis (ll species), Lekrugeria Navis (one species), Berlekrumyia U. Aspock & H. Aspock (one species), Isoscelipteron Costa (one species) and Podallea Navis (12 species). History of research on African Berothidae is concisely outlined. Occurrence of genera and species in the countries of Africa is recorded in tables. Chorology of genera is discussed and illustrated with distribution maps. Key words: Neuroptera, Berothidae, Africa, research history. distribution, zoogeography.

INTRODUCTION Africa seems to be of prime importance as a distribution centre of Berothidae. Three1 of the four subfamilies, Rhachiberothinae, Nosybinae and Berothinae occur in this region. About 40 species have hitherto been described, making Africa the continent with the richest known berothid fauna.

RESEARCH ON AFRICAN BEROTHIDAE History of research on African Berothidae is scarcely 100 years old. It began at the end of the nineteenth century with the description of Berotha eatoni in 1898, by Robert McLach- lan. Although the family had already been established by Handlirsch in 1908, subsequent species were described as Hemerobiidae as late as 1918. The chronological succession of new taxa (Tab. 1) reflects, to some extent, historical expeditions to Africa (see below). Two early monographical papers on Berothidae which include African species (Kriiger 1922; NavBs 1929) are only mentioned for the sake of completeness. The first outstanding and comprehensive study of African representatives of the family, was Bo Tjeder's monograph on the Berothidae of southern Africa (Tjeder 1959). A new, still effective, classifi- cation of the family was proposed a few years later by MacLeod & Adams (1967). A series of generic revisions of Podallea NavBs, 1936, Nosybus Navhs, 1910, Sphaeroberotha NavBs, 1930, and Lekrugeria NavBs, 1929, was started by us in the 1980s (U. Aspock & H. Aspock 1981, 1983, 1984, 1986), and is being continued. The present state of knowledge on a world-wide scale has recently been summarized (U. Aspock 1986).

'See addendum at the end of this paper. Table 1. Berothidae of Africa: chronological list of dcscribcd species.

I Original name 1898 Berotha eatoni McL...... Nodalla catoni (Mcl,.) 1910 Nosyhus nobilis NAV...... Berotha vasscana NAV ...... Podallca vasscana (NAV.) 1915 Berotha lcroiana ESB.-PET...... I'odallca lcroiana (1:SB.-PI:T.) Berotlia kocnigi 1:SB.-PET...... 1-ekrugcria kocnigi (ESl3.-P1:T.) 1918 Nosybus navasi BANKS...... 1920 Berotha geyri ESH.-PET...... Nodalla gcyrl (17SB.-PI T.) Berotha sal~aricaESB.-PliT...... Nodalla sal~arica(L SB -PI:T ) 1922 Nosybus togana KKUGLR...... 1926 Nodalla acgyptiaca NAV...... Nodalla sinaitica NAV...... 1929 Costachillea barbara NAV ...... ?syn. of Nodalla catoni (McL.) 1930 Costachillea werncri CSl3:PCT. .... Nodalla wcrncri (FSB.-PET.) Sphacroberotha dumonti NAV. .... ?syn. of Nodalla gcyri (I;SB.-PICT.) 1931 Bcrotl~asashilana NAV...... Podallea sashilana (NAV.) 1935 Berotha seyrigina NAV...... Podallca seyrigina (NAV.) 1936 Berotha squamulata NAV...... Podallell squalnulata (NAV.) Nodalla lincata NAV...... Nodalla striatella NAV...... Podallea seriata NAV...... nov. syn. of Podallca lcroiana (I-SB.-PIYI'.) 1952 Nosybus orthoptera FRASER ..... syn. of Nosyb~tsnavasi BANKS 1954 Acroberotha xiphophora TJID .... Podallea xiphophora (TSI:l>.) 1955 Acroberotha pauliani F'RASI'R .... Podallca patrlian~(I'RASI, R) 1959 Rhachibcrotha signifera TJCl). .... lit~achibcrothasrnithcrsi TJrD. .... Mucroberotl~afasciata TSllD...... Nosybus minutus TII:D...... Acrobcrotha tricirrata TJLD...... Podallca tricirrata (TJI:D.) Berotha cxarmata TJFD...... Podallca cxarmata (TJED.) 1968 Nodalla oranensis R0USSI:T ...... ?syn. of Nodalla catoni (McL.) Mucrobcrotha vcsicaria TJED...... Mucrobcrotha nigrcscens TJI:D. .... 1979 Bcrotha glaserella U.A. & H.A. & 1fZL...... 1981 Podallea tjcderi U.A. & H.A...... Podallca pcllita U.A. & t1.A...... 1983 Nosybus zernyi LJ..A. & H.A...... Nosybus venustus [).A. & H.A. .... Nosybus tjederi U.A. X H.A...... Nosybus hirsutipennis LJ.A. & I3.A. .. 1988 Hcrlckruniyia :tfricanclla U.A & H.A. . Podallea ~nansclliU.A. & H.A......

So far, 41 species (including a few synonymous and unclarified ones) of African provenance have been described. Some of these species were discovered more or less incidentally during the colonial history of Africa. Zoological expeditions to Africa were, however, also undertaken in early times. These include: the Sahara expedition led by Freiherr Geyr von Schweppenburg in 19131 19 14 (see Esben-Petersen 1920); the Austrian Kordofan expedition under the leadership of F. Werner in 1914 (see Esben-Petersen 1930); the French Ethiopian expedition 'Mission scientifique de I'Omo' in 193211933 (see Arambourg, Chap- puis & Jeanne! 1935; NavAs 1936), which are worthy of mention, not only for historical reasons but also for having had important implications for research on Berothidae. The Lund University expedition to southern Africa in 1950 (see Brinck 1955) was an outstanding event (and the reason for Bo Tjeder's above-mentioned monograph) in continuing Berothidue c?f Africa

Table ?. Records of herothid genera in countries of Africa.

V) 5 6 COUNTRY 0

I\LC; 1. R 1.4 ...... 4NGOL4 ...... 131 NIN ...... 130PHI~TFIATS\VhNA ...... 13O'TSiV ,AKA ...... BIIIIKINt1 l:..\SO ...... 13I:RI'NDI ...... C.hls:KOON ...... C41'1' V17RDl lSI.ANDS ...... CI'NTRAL !1f:RICAN RTP. .... CHAD ...... ('1Sh;L:l ...... COh30RO ISLAKIIS ...... CONC;O ...... I>JISOliTI ...... 1 (;\r'fT ...... l~Qt7.4TOKIAL.GL;INI.'A ...... KTHlOM.4 ...... G,\BON ...... GAMBIA ...... GHAKA ...... (; L'IN I'A ...... GIIINI:A BISSAU ...... IVORY COAST ...... KINYA ...... KWASDFBELC ...... LI~,SO'TII 0 ...... LIBERIA ...... LIBYA...... MAI~AC;ASCAI< ...... RIALAWI ...... MALl ...... MAURITANIA ...... MAI!RITIUS ...... MOROCCO ...... MOZAMBIQI.IL: ...... NAMIBIA ...... NIG1:K ...... NI(;I; Kl.4 ...... RWANDA ...... SAHARA ...... S;\O TO~I~AKD PRINCIPI: .... St7NI:GAL ...... Sl.TCHJ:LLI. S ...... S1 I:S

COUNTRY

TANZANIA ...... TOGO ...... TRANSKCI ...... TUNISIA ...... UGANDA ...... VI' NDA ...... ZAIRE ...... ZAMBIA ...... ZIMBABWL: ......

Swedish explorations of southern Africa, which had a long tradition dating back to Linnaearl times. This expedition stands as pars pro toto for present day collecting which, however, has now been largely taken over by local scientists. Our own field experiences in several parts of Palaearctic and Afrotropical Africa, have encouraged me to summarize what has already been done, and to indicate what still has to be done.

Fig. 1. Recorded distribution of Berothidae in Africa Berothidae of Africa

The projection of all traceable records of Berothidae onto a map of Africa (Fig. l) indi- cates an abundance that was never observed in collecting. On the other hand, recording of genera in the countries of Africa (see Tab. 2), certainly reflects the frequency of genera. but also collecting traditions in some of the countries and a total lack in others (This al- phabetical list of African countries corresponds with Der Fischer Welt Almanach 1988, Fischer Verlag, Frankfurt am Main, 1987).

BIOLOGY

The biology of African Berothidae (and the family in general, see New 1986) is almost unknown, most species having been collected at light. Eggs and early larval stages of a Podallea species and Mucroberotha vesicaria Tjeder have been studied by Minter (personal communication and Minter 1990). The whole life cycle, however, is only known for one Nearctic Lomamyia, which is a true termitophile (C.A. Tauber & M.J. Tauber 1968). The idea of termitophily is thus a guide for further investigations.

TAXONOMICAL ASPECTS AND CHOROLOGY OF AFRICAN BEROTHIDAE

As stated above, three1 of the four very distinctive berothid subfamilies are represented in Africa. Rhachiberothinae (two genera) and Nosybinae (monotypic) are extremely apo- morphic, each comprising a very homogeneous unit. The Berothinae (five genera) on the contrary, are surprisingly heterogeneous; sister group relationships of the subfamilies are, however, unresolved. Taxonomy of the species is in progress, and undiscovered taxa can still be expected. Numerous new species await description and several described taxa need to be clarified. They are, nevertheless, available for zoogeographical accounts at the generic level, so have been recorded in the corresponding distribution maps.

Subfamily Rhachiberothinae Tjeder, 1959

Rhachiberothinae are the only extant berothids with raptorial foreiegs. The male of Rhachiberotha Tjeder, 1959, is still unknown but the male of Mucroberotha Tjeder, 1959,

Table 3. Rhachiberothinae: records of species Rkhiderotha Tjeder and Mucroherothu Tjeder in countries of Africa.

Species 3 U N -- Rh. signifera TJ17D.

Ilh. srnittiersi TJIiD.

M. fiisciata TJED.

M. vcsicaria TJEY.

M. nlgrescens TJFD.

M. sp.

'see nddendunl at the end of this paper Fig. 2. Rhachiberothinae: recorded distribution oi Rhnchihrrotha Tjeder, and Muc.roberotha Tjcde~ shows a striking similarity to symphi-asine Mantispidae with respect to the genitalia and in having only four tarsomeres. These facts caused serious doubts concerning the affilia- tion of this subfamily (Tjeder 1968). We consider them as being true Berothidae, but the problem is complex indeed, and should not be limited to these characters only. Rhachiberotha is known by two species from Zimbabwe, Mucroberotha by three species from South Africa, S.W.A./Namibia and Zimbabwe and an undescribed species from An- gola (Tab. 3). Both genera are endemic and seem to be restricted to southern Africa (Fig. 2). A revision of this subfamily, including further new species (not yet included in the map), is planned for the near future in collaboration with M.W. Mansell.

Subfamily Nosybinae MacLeod & Adams, 1967.

Nosybus Navas, 1910, and consequently the monotypic Nosybinae, presently comprise seven species. The records of N. nohilis Navas from South Africa are new (M.W. Man- sell, personal communication). Nosybinae are endemic to Africa and restricted to the Afrotropical Region (Tab. 4 & Fig. 3) but significantly, are more frequent and more ex- tensively distributed than the Rhachiberothinae.

Subfamily Berothinae Handlirsch, 1908. This subfamily is represented on all continents, although restricted to the warnier climates, and comprises 15 genera. Berothinae occur throughout the African Continent being represented by five geographically restricted, heterogeneous genera.

Fig. 4. Berothinae: recorded distribution of Nodalla Navas. Table 5. Berothinae: records of species of Nodalla Navas in countries of Africa. Berothidae of Afiica

Fig. 5. Berothinae: recorded distribution of Berlekrumyirr U.A. & H.A. and African distribution of hkrugeriu Navas. and Isosceliptrron Costa. Table 6. Berothinae: records of species of Ltkrugeriu Navis, Berlekrumyia U.A. & H.A., and Isoscelipteron Costa in countries of Africa.

Nodalla Navris, 1926, of which Sphaeroberothu Navris, 1930, is now clearly a new syno- nym, has been somewhat problematic due to inaccessibility of types. Several species still require clarification, therefore the distribution map (Fig. 4) refers only to the genus. Ta- ble 5 includes all species described so far, ignoring synonymy. The discussion on syno- nyms which was initiated in our revision of Sphaeroberotha (U. Aspock & H. Aspock 1984), will soon be finalized in a forthcoming paper on Nodullu. Nodalla is the dominant genus of the family in Palaearctic Africa, several species certainly being restricted to the continent, whilst several probably also occur in the Middle East.

Berothidae of Africa

0 KHACHiBEROTHA AAA MUCROBEROTHA -... ivOSYBUS m- NOCALLA

,,o a--* LEKRUGERIA **a BEKL~EKRUMYIA - - - 0-*-ISOSCELIPTERON -- --- PODAiLEA

GENERA OF BEROTHIDRE 0 700 IUi'O I5C" ll.OU*rn ,- 1

.'~-- ~.-~ I

Fig. 7. Kccorded distribution of berothid gencra in Africa

Lekrugeria NavBs, 1929, with a total of three species, occurs disjunctly in Africa and Asia (U. Aspiick XL H. Aspijck 1986); the species are extremely rare. The only African species. L. kocnigi (Esben-Petersen, 191 S) occurs in a characteristic belt across the continent in the southern parts of the Sahcl zone (Fig. S), thus occurring in several countries (Tab. 6). Berleknlmyiu U. Aspock & H. Aspiick, 1988, is known only from Mozambique and Malawi (U. Aspock & H. Aspiick 1988) (Tab. 6 & Fig. 5). It is most probably endemic to Africa and restricted to the Afrotropical Region. Isoscelipteron Costa. 1863, is the genus with the widest known distribution among the Berothidae (U. Aspock 1986, 1987). It occurs extremely disjunctly in Europe, Asia and Australia. and only touches the northwestern edge of Palaearctic Africa with a single species (Tab. 6 & Fig. 5). Podalleu Navas, 1936, comprises 14 described species. Podalleas are the most frequent and typical berothids of Afrotropical Africa, and demonstrate the expansion of the Afrotrop- ical Region into the south of the Arabian Peninsula through one of its species. The genus is represented in the Madagascan subregion as well (Tab. 7 & Fig. 6). Table 7 and figure 6, also both contain unpublished new records of known species, as well as some species that await description. In summary, it can be stated that three of the eight known genera are certainly, and another probably, endemic to Afr~caand restricted to the Afrotropical Region: these are Rhachiherothu, Mucroheroth~~,No~ybus and Berlebumyia (Fig. 7). Nodullu and Podulleu divide Afr~caby occupying its Palaearctic and Afrotropical parts respectively, but share a common narrow central region of the continent. Isoscelipteron is represented only marginally as a relatively young gift from Europe to Africa. Lekrugeria is reduced to a supra-equatorial belt across the continent. A few berothid species are certainly eremial, the bulk of species is, however, arboreal in a broad sense. We are still far from an understanding of any causal zoogeography of African Berothidae, but may assume that the berothid fauna of Africa is a very old and autochthonous one.

ACKNOWLEDGEMENTS

I would like to express my cordial thanks to my husband, Horst, for his intensive scientif- ic co-operation and for many critical discussions. I am also very grateful to our friend and highly esteemed colleague, Dr. Mervyn Mansell, for reading the manuscript and for linguistic improvement.

REFERENCES

ARAMBOURG, C., CHAPPUIS, P.-A. & JEANNEL, R. 1935. Itineraire et liste des stations. In: Mission Scientifique de I'Omo, 11, jusc. I. Memoires du Musc'um Nutzonul d'Histoire Nuturelle, Paris 2: 1-22. ASPOCK, U. 1986. The Present State of Knowledge of the Family Berothidae (Neuropteroidea: Planipennia). In: Gepp, J., Aspiick, H. & Hiilzel, H. (Eds) Recent Re.srarch in Nc.uropterology. Proceedings of the Second Intc,rnurional Symposium on Neuropterology. Hamburg. F. R. G. . 1984. 87- 10 1 . Graz, Austria. ASPOCK, U. 1987. The Berothidae (Neuropteroidca: Planipennia) of the Middle East. In: Krupp, F., Schneider, W. & Kinzelbach, R. (Eds) Proceedings of the Symposium on the Fuunu und Zoogeogruphy ($the Middle East. Mainz, F.R. G., 1985. 160- 167. Beihejie zum Tiibinger Allas des Vorderen Orients, Reihe A, 28. Wiesbaden, F.R.G. ASPOCK, U. & ASPOCK, H. 1981. Das Genus Podallea Navas, 1936 (Neuropteroidea: Planipennia: Berothidae). Zeit.schriji der Arheitsgem~insch~lftinsclu 0sterreichi.scher Enrornologen 32: 81-96. ASPOCK, U. & ASPOCK, H. 1983. Das Genus Nosybus Navis, 1910 (Neuropteroidea: Planipennia, Berothidae). Zeitschriji der Arbeitsgemeinsc-hufi hterreichischer Entomologen 34: 9 1 - 105. ASPOCK, U. & ASPOCK, H. 1984. Zur Kenntniss des Genus Sphueroberothu Navas, 1930 (Neuropteroidea: Planipennia: Berothidae). Zei/.schrifr der Arbeitsgemeinschafi Osrerreichischer Entomologen 35: 65-83. ASPOCK, U. & ASPOCK, H. 1986. Das Genus Lekrugeriu Navas (Neuropteroidea: Planipennia: Berothidae: Bcrothinae). Zeitschriji der Arbeitsgemeinschafi hterreichischer Entomologen 37: 85-98. ASPOCK, U. & ASPOCK, H. 1988. Berlekrumyia africurrella n.g. et n.sp. und Podulleu munselli n.sp. - zwei ncue Berothiden-Spezies aus dem Siidosten Afrikas (Neuropteroidea: Planipennia: Berothidae). Zeitsc-hriji der Arbeitsgemeinschafr iirterreichischer Enromologen 39: 113-123. BRINCK, P. 1955. Swedish Exploration of South African Animal Life during 200 years. In: Hanstriim, B., Brinck, P. & Rudebeck, G. (Eds) South African Animal Lib 1: 11-61. Swedish Natural Science Research Council, Stockholm, Sweden. ESBEN-PETERSEN, P. 1920. Ncuropteren der Inneren Sahara. Archiv fir Nuturgeschichte 84: 143- 159. ESBEN-PETERSEN, P. 1930. XXVII. Neuropteren. In: Wisscnschaftliche Ergebnisse der mit Unterstiitzung der Akadcmie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommcncn zoologischen Expedition nach dem Anglo-~g~~tischenSudan (Kordofan) 1914. Denkschrijien der Aklemie der Wissc~n.schufienin Wien. Mathematisch-nuturwiis.sennschafi1icheKlusw 102: 201-206. Wicn. KRUGER, L. 1922. Berothidae. Beitrage zu einer Monographie der Neuropteren-Familie der Berothiden. Stettiner Entomologische Zeirung 83: 49-88. MacLEOD, E.G. & ADAMS, P.A. 1967. A review of the taxonomy and morphology of the Berothidae, with the description of a new subfamily from Chile (Neuroptera). Psyche 74: 237-265. MINTER, L.R. 1990. A comparison of the eggs and first instar larvae of klucroberotha vesicuriu Tjeder, with those of other species in the families Berothidae and Mantispidae. In: Mansell, M.W. & Aspock, H. (Eds) Advances in Neuropterc~logy.Proceedings (fthc, Third International Symposium on Neuropterol- ogy. Berg en Dal, Kruger Nutionul Purk, R. S. A., 1988. Pp. 115-129 Pretoria, R.S.A. 1990. NAVAS, R.P.L. 1929. Monografia de la Familia de los Berotidos (lnsectos Neurcipteros). Memorius de lu Academia de Ciencicis Exuctu.~,Fisico-Quimiras y Narurales de Zaragozu 2: 1- 107. NAVAS, R.P.L. 1936. Neuroptera, Embioptera, Plecoptera, Ephemeroptera et Trichoptera. In: Mission Scientifique de I 'Omo, 111, fasc. 19. MPmoires du Museum National d 'Histoire Nuturelle, Puris 4: 101-128. Berothidae of Africa

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ADDENDUM

During the course of our post-symposium excursion in February 1988, a representative of the subfamily Cyrenoberothinae MacLeod & Adams was unexpectedly and surprisingly discovered on Ameib Ranch, Erongo Mountains in S.W.A./Namibia. This subfamily had hitherto only been known from a single species (Cyrenoberotha penai MacLeod & Adams) from South America (Chile). The discovery of the subfamily in southern Africa demonstrates the presumable Gondwanian character of this taxon; the African representative being a new genus and species was described as Manselliberotha neuropterologorum and dealt with in detail by U. Aspock & H. Aspock (1988) in Zeitschrift der Arbeitsgemein- schaft Osterreichischer Entomologen 40: 1-13. All four subfamilies of Berothidae are now known to occur in this subregion.

Address of author: Dr Ulrike Aspock Naturhistorisches Museum Wien Burgring 7 A-1014 Wien Austria Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 1430

Reference Citation: Aspöck, U. 1990 [1990.??.??]. The Berothidae of Africa: a review of present knowledge (Insecta: Neuroptera). Pp. 101-113 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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