Introduction
The superorder Neuropterida comprises about 6500 extant valid species worldwide, distributed across three orders, Neuroptera (“lacewings”) Megaloptera (“alderfies”) and Raphidioptera (“snakeflies”), with sixteen, two and two families respectively.
South Africa harbours a rich and diverse lacewing and alderfly fauna, with 415 valid species currently known from the country. This number continues to increase through documentation of undescribed taxa, further exploration, and the revelation of cryptic species through morphological and molecular analyses. While the orders Neuroptera and Megaloptera are well represented in South Africa, Raphidioptera are absent from the Afrotropical Region. All families of Neuroptera, except Nevrorthidae, Nymphidae, and Ithonidae are represented in South Africa and the Megaloptera fauna includes both Corydalidae and Sialidae.
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Since April 1994, South Africa has been divided into nine provinces (Figure 1). These vary widely in size, from the Northern Cape, which covers nearly one-third of the country's land area, to Gauteng, which includes only 1.5% (Table 1).
Prior to 1994, South Africa had a long and complex history of political boundaries, which reflect the occupation and settlement of the country. The Dutch colonized southwestern South Africa in the vicinity of the Cape of Good Hope (now part of Western Cape Province) in 1652 under the Dutch East India Company, and the area later came under British control at the turn of the eighteenth century. The British named the area the Cape Colony, but it was known under a variety of names in the major European languages of that era, as reflected on the labels and in the publications of many early European collectors and authors. These include: Kaap de Goede Hoop (Dutch); Cape, Cape Colony, Cape of Good Hope (English); Cap, le Cap, Cap de Bonne Esperance, Cap d. Bonn Esp. (French); Capitae bonnae spei, Cap. b. sp. (Latin); and Kap, Kap der Guten Hoffnung, südlichen Afrika (German).
By the time the British formally annexed the Cape Colony in 1814, much of the area had been occupied by farmers of Dutch descent, known as the Boers (farmers) or Afrikaners, who had evolved their own language, Afrikaans, from Dutch. By 1835 the British had extended their territory eastwards as far as the Great Kei River and northwards to the Orange River. They named a number of districts, including British Kaffraria, the area between the Kei and Keiskamma rivers. Specimen labels of several early collectors and species authors bear the epithet “Caffraria”, a locality that became rather loosely interpreted to encompass most of present-day Eastern Cape Province. It was in this area that the British first encountered the Xhosa, a branch of the Nguni people who had originally migrated southwards over several centuries.
In 1835 the Afrikaner farmers (Boers) started a migration northwards to escape the repressive British rule of the Cape Colony, a migration known as “Die Groot Trek” (The Great Migration). They moved across the Orange and later the Vaal Rivers and established two separate Afrikaner Republics, the Orange Free State and the Zuid-Afrikaansche Republiek (later Transvaal). The former comprised what is now known as Free State province, while the latter, proclaimed in 1839, now includes the current provinces of Gauteng, Mpumalanga, North West, and Limpopo.
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The British had established a trading post at Port Natal, now Durban, in 1824. In the same year they signed a treaty with the Zulus, another branch of the Nguni people, ceding them Port Natal and the immediate surrounding area, which became Natal, and now KwaZulu-Natal Province. The interior of Natal had been occupied since the 16th century by the Nguni branch of Bantu-speaking peoples, who later merged into the Zulu nation under successive chiefs, most notably Shaka and Dingane. Part of the Boer leadership migrated eastwards into this territory in about 1837 and established the Republic of Natal, with its capital at Pietermaritzburg and its northern border at the Tugela River. The British annexed this Boer republic in 1843. In the meantime, numerous battles and skirmishes ensued between the Boers, the British, and the Zulus, resulting in one of the bloodiest periods in South African history. The Zulus largely retreated to the area north of the Tugela into the area known as Zululand, where they still retain much of the territory. At that time, South Africa comprised four provinces: the Cape Province and Natal under British rule and the Orange Free State and Transvaal (Zuid-Afrikaansche Republiek) under the Boers. The British and the Boers fought two major wars until the defeat of the Boers during the Anglo-Boer War (1899-1902). The four provinces then merged to form the Union of South Africa in 1910, which remained under British administration until 1961 when it formally became the independent Republic of South Africa. The four-province division prevailed until 1994 when the current government under the African National Congress came to power after independence from the Nationalist Party government that had ruled since 1948. The existing four provinces were then subdivided into the nine current divisions: Cape Province was split into the Western Cape, Northern Cape, and Eastern Cape provinces; Natal was renamed KwaZulu-Natal; Transvaal was split into Gauteng, Mpumalanga, North West, and Limpopo provinces; and Free State Province remained unchanged.
Prior to the Nguni (Bantu-speaking peoples) and European immigrations, South Africa had been populated by the KhoiSan peoples, two groups with similar origins and language. The San people were hunter-gathers who left the only documented history of original South African inhabitants through thousands of rock paintings and engravings throughout southern Africa. The Khoikhoi people were largely pastoralists. The San and Khoikhoi roamed a pristine southern Africa with abundant game but left no structural records of their presence before they were displaced to remote areas of the western and northern parts of South Africa by the advancing Nguni migrations from the north and European settlers from the south.
Biomes of South Africa
South Africa is one of the most biologically diverse countries in the world (https://www.sanbi.org), after Indonesia and Brazil, due to its varied topography and climate, which have engendered the formation and evolution of the biomes that support biodiversity. South Africa has 2,798 kilometres of coastline, with the cold Atlantic Ocean to the west and the warmer Indian Ocean to the east. These two oceans influence the climate, which, together with geology and resultant topography, has combined to produce a suite of distinct biomes that range from Desert in the west, through the Fynbos, Succulent Karoo, Nama-Karoo, Grassland, Savanna, Albany Thicket, Indian Ocean Coastal belt to Forest biomes in the east. Mucina and Rutherford (2006; Figure 2) mapped these nine major biomes on the basis of vegetation, but each has also engendered the evolution of a distinctive lacewing fauna, especially in the families Nemopteridae and Myrmeleontidae, which have radiated significantly resulting in many endemic taxa. South Africa is home to two thirds of the global species of Nemopteridae, while 41 of the 121 world Palparinae species are endemic to South Africa. This diversity, and the high levels of endemism, has led to growing concern about the conservation status and preservation of many populations of lacewings that are now threatened by burgeoning agriculture and urban development, and the ever-present threat of habitat destruction and insecticide drift. Many taxa, especially the unique Cape fauna of Nemopteridae, are only known from single populations or unique types and are gravely threatened in the fragile habitats in which they evolved. The major objective of the Southern African Lacewing Project (Mansell 2002) is consequently to explore and document the fauna, to establish the conservation status of its constituent taxa and to inform the protection and conservation of this unique South African heritage.
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Each of the provinces of South Africa includes two or more of the country’s nine major biomes, which have strongly influenced the evolution and diversity of the local Neuroptera fauna. The number of species currently known from each province is also, to some extent, a reflection of collecting effort, with the best surveyed areas being those in close proximity to one or more major museums and/or universities.
Distribution of South African Neuropterida by Province
Of the present provinces of South Africa, Limpopo Province is the most speciose with 182 species (43.4% of the known South African fauna; Table 2), even though it encompasses only two of the country’s major biomes (Savanna and sparse Grassland). Its richness is enhanced by several geographical and climatic characteristics, as well as being relatively well surveyed for lacewings. The physical features include the Soutpansberg and Lebombo mountain ranges, and the Blouberg inselberg, all of which are heavily forested and are biodiversity hotspots. There is also reduced rainfall to the north in the Limpopo River valley where intrusions of deep Kalahari sand provide a corridor for species extending into Limpopo Province from the arid west.
Limpopo is followed by the Western Cape Province with 156 species (37.6% of the total country fauna). It includes four of the major biomes: Fynbos, Succulent Karoo, Nama Karoo, and small patches of Forest. This area has been a major evolutionary centre for plants and animals and especially the lacewing family Nemopteridae and, to a lesser extent, Myrmeleontidae that have undergone major radiations in this region, resulting in the majority of South Africa’s endemic taxa. This province, together with the Eastern Cape harbours the Fynbos biome, regarded as one of the six Floral Kingdoms of the world in what is known as the Cape Floral Kingdom. The Nemopteridae in particular, whose adults are exclusively pollenophagous have undoubtedly co-evolved with the Cape Flora. The Cape Fold Mountains that extend into the Eastern Cape also harbour a rich complement of Megaloptera that are all endemic to the Western and Eastern Cape provinces, with one species occurring in KwaZulu-Natal and Limpopo.
Mpumalanga Province includes only two biomes, Grassland and Savanna, and contains 147 species (35.4% of the country total). This relatively low number is mainly due the large proportion of the province being historically covered by highland grassland, the “Highveld”, which is not particularly conducive to lacewing diversity. This is, however, offset in the east by the lower altitude subtropical “Lowveld,” comprising the Savanna biome, that includes the southern portion of the Kruger National Park which has been well surveyed for lacewings.
KwaZulu-Natal with 146 species (35.1% of the fauna) is unique in that it includes the Indian Ocean Coastal Belt biome, along with Grassland and Savanna. The diversity of this province is consequently influenced by faunal elements, including lacewings, that extend their ranges southward along the coast and into the province from East and Central Africa. Consequently, KwaZulu-Natal has very few endemics. One of the taxa that epitomizes this distribution is one of South Africa’s largest antlions, Palpares inclemens (Walker), whose range extends from Durban in the south, northward through Mozambique to Tanzania, Kenya and beyond.
The North West Province has 136 species (32.8% of the fauna) in two biomes, Grassland and Savanna. It lies between the Northern Cape and Limpopo provinces and consequently shares many taxa with them.
The Northern Cape Province, which occupies about 30.5% of South Africa’s land area and encompasses five biomes, has 133 species (32.0% of the fauna). The biomes include Desert, Succulent Karoo, Nama Karoo, Fynbos, and Savanna. The province also harbours a number of species endemic to South Africa, and the relatively low number of taxa is certainly due to the size of the area and inadequate collecting effort. The mostly-arid landscape is, however, richly endowed with Myrmeleontidae and Nemopteridae.
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The Eastern Cape Province with 102 species (24.6% of the fauna) occupies South Africa’s second-largest land area, and includes seven biomes: Fynbos, Grassland, Nama Karoo, Savanna, Indian Ocean Coastal Belt, Forest, and the unique Albany Thicket. Despite its size and richness, the low number of species is certainly due to large parts of the area not being adequately surveyed, particularly the coastal and montane forests. There are several species that are endemic to the province, including two species of Proctarrelabis (Ascalaphidae) and an undescribed species of Maula that is endemic to the narrow coastal belt.
Gauteng Province -- the smallest South African province but the one that contains the country’s industrial, mining, and economic heartland, has 96 species (23.1% of the total fauna). It is probably the best explored of all provinces, mainly because of its small size and the presence of several major museums and universities. It encompasses two biomes, Grassland and Savanna, and is bisected by the Magaliesberg mountain range, which offers various habitats and has a relatively rich diversity of lacewings, but no endemic species.
Free State Province, with only 53 species (12.8% of the lacewing fauna of South Africa) is also the most depauperate of the nine provinces with regard to family representation. Although it includes three biomes, Grassland, Nama Karoo, and Savanna, the Grassland biome, which is not conducive to lacewing diversity, is by far the most dominant vegetation type in this province.
A summary of family representation by province is provided in Table 3. From this, it can be seen that only five families, Ascalaphidae, Chrysopidae, Coniopterygidae, Hemerobiidae, and Myrmeleontidae are known from all nine provinces. Limpopo and the Western Cape each harbour 12 of the 15 families, while Gauteng, KwaZulu-Natal, Mpumalanga, North West, and Eastern Cape each have 10, followed by Northern Cape (8) and Free State (5).
Taxonomic History of South African Lacewings and Alderflies
The documented history of South African lacewings is intricately linked with that of the Afrotropical Region as many taxa that were described from further north in Africa also occur in South Africa. This historical review focuses on taxa that were originally described from South Africa. Historically, 325 species names, described by 40 authors, have been based on specimens collected from South African localities. Of these names, 267 are currently regarded as valid, although several of these will require future synonymization.
Hemerobius speciosus Linnaeus, 1758 (now Palpares speciosus) was the first neuropteran recorded from South Africa, followed during the 18th century by Mantis pusilla Pallas, 1772 (Sagittalata pusilla), Ascalaphus capensis Fabricius, 1781 (Nephoneura capensis), Myrmeleon capense Thunberg, 1784 (Proctarrelabis capensis), Myrmeleon luteus Thunberg, 1784 (Pamexis luteus), and Mantis fausta Thunberg, 1784 (Mantispa fausta). The holotype of H. speciosus is still in good condition in the Linnaean Society vault in London.
The first half of the 19th century saw contributions from eight authors: Olivier (1811; 2 species described, 2 still valid; Crambomorphus sinuatus, Pamexis bifasciatus); Leach (1815; 1, 1; Nemopterella africana); Klug (1836; 3, 2; Sicyoptera dilatata, Laurhervasia setacea); Westwood (1836; 2, 2; Nemia angulata, Nemia costalis); Burmeister (1837, 1839; 5, 2; Palpares caffer, Creoleon luteipennis); Erichson (1839; 4, 3; Afromantispa tenella, Mantispa fuscipennis, Sagittalata dorsalis); Guérin-Méneville (1844; 1, 1; Pseudoclimaciella erichsoni), and Rambur (1842; 7, 6; Italochrysa nevrodes, Apochrysa leptalea, Helcopteryx rhodiogramma, Myrmeleon ochronevrus, Myrmeleon lanceolatus, Banyutus pulverulentus).
Of the pre-1850 works that contributed to the early knowledge of the South African fauna that of Rambur (1842) is particularly significant. Not only did he describe several valid species in three families (Ascalaphidae, Chrysopidae, and Myrmeleontidae), an additional species (Syngenes longicornis) that was described from an unknown locality has subsequently proven to be of South African provenance and, perhaps, most significantly, he described the charismatic genus Palpares, within which most of the large, strikingly-patterned, antlions of southern Africa were originally placed.
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Seven authors contributed to knowledge of the fauna during the second half of the 19th century, with John Obadiah Westwood, the Oxford academic and first Hope Professor of Entomology, being the only one of the early entomologists to continue publishing during this period – Pseudoclimaciella tropica (Westwood, 1852) and Nemopterella remifera (Westwood, 1874). Apart from Schneider (1851) (Pseudomallada chloris), the remainder of the century saw contributions by five authors: Walker (1853, 1860; 9, 4); Hagen (1866, 1886; 2, 1), Gerstaecker (1863, 1885; 4, 3), McLachlan (1867, 1869, 1870, 1873; 8, 6), and Kolbe (1900; 1, 1), although most of these authors also described taxa from other African countries, which are now known from South Africa.
The first half of the 20th century was dominated by the infamous describer Longinos Navás whose publications on Afrotropical Neuropterida spanned over 30 years from 1908 to 1939. During this time, he described at least 660 species from the Afrotropics, of which only 351 are currently valid, and many more await formal synonymy. Navás described 45 species, in 31 different papers, specifically from South Africa, but only 25 remain valid, and at least another eight will require future synonymization (Navás 1908, 1909, 1911, 1912a,b,c,d,e,f, 1913a,b,c,d,e, 1914a,b,c,d,e, 1921a,b,c, 1922, 1923a,b, 1925a,b, 1927a,b, 1929, 1934). It is unlikely that the taxonomic disorder caused by this author will ever be fully resolved, as most descriptions are too poor to interpret, and many type specimens are lost, destroyed, or based on unique females that are impossible to identify. By contrast, several valuable contributions were also made during this time by Enderlein (1914; 1, 1): van der Weele (1909; 6, 6); Esben-Petersen (1912, 1916, 1917, 1920, 1922, 1924, 1925; 18, 16); Banks (1909, 1911; 4, 3); and Kimmins (1929, 1930, 1933, 1935, 1937, 1948, 1956; 13, 12). It was also the era of the first South African taxonomist, Louis Péringuey, Director of the South African Museum, Cape Town, who made a significant contribution in two papers (1910, 1911), describing 20 local species, 15 of which remain valid. He was followed by Keppel Barnard (1931, 1940), his successor at the museum, who described three species of Corydalidae from the Western Cape Province.
The second half of the twentieth century was the era of the great Swedish neuropterist, Bo Karl Herman Tjeder, who made the finest contributions yet to knowledge of South African Neuroptera (1955, 1957, 1959, 1960, 1961, 1966, 1967, 1968, 1969, 1992). In a series of landmark publications in South African Animal Life, Tjeder (1957, 1960, 1961, 1966, 1967) significantly elevated the scope and rigour of revisionary studies for South African Neuroptera, establishing the precedent for neuropteran monography that has rarely been exceed anywhere in the world before or since. In all, Tjeder described 92 species from South Africa, 83 of which remain valid. He also described 34 genera, 26 of which are still recognized. Kimmins (1956) continued to publish during the 1950s, and Handschin (1959) added one additional species. Meinander (1972, 1975, 1983, 1998) and Monserrat (1994) added 16, and four species of Coniopterygidae respectively. Oswald (1994) contributed a species of Psychopsidae, and U. Aspöck & H. Aspöck (1988, 1997) documented two new species (Berothidae and Rhachiberothidae), with U. Aspöck & Mansell (1994) describing a further two rhachiberothids. Tjeder (1992) and Tjeder & Hansson (1992) added 13 new species of Ascalaphidae from South Africa, while Hölzel (1987, 1989, 1990, 1993) added six new species (five still valid) of Chrysopidae as part of his revisionary studies of that family. This period also marked the beginning of the active involvement of South African neuropterists in the local fauna when Mansell (1977, 1980, 1981, 1987, 1990, 1992, 1996) commenced his publications on Nemopteridae and Myrmeleontidae. Picker (1984, 1987) described two Nemopteridae and Minter (1986) recorded the first species of Dilaridae from South Africa.
So far, the 21st century has yielded 15 new taxa: two Chrysopidae by Hölzel & Duelli (2001, 2003), one species of Psychopsidae (Bakkes 2017), six additional Myrmeleontidae by Mansell (2004, 2013, 2018) and Mansell & Ball (2016); while Price et al. (2012), Liu & Price (2013) and Liu & Hayashi (2013) have added seven species of Megaloptera, which has greatly enhanced knowledge of this order in South Africa.
Besides these known taxa, there are approximately 106 taxa in South Africa that still require formal descriptions. These comprise 84 Myrmeleontidae, 15 Nemopteridae and 7 Ascalaphidae, which are currently the priority of the Southern African Lacewing project. There are also several undescribed taxa in other families, which bring the total of species that possibly occur in South Africa to about 550–580 species. Approximately 20 currently named species will also be synonymized during revisions, which should, however, not affect the above estimate too adversely.
Collectors and Collections
There was no specialist neuropterist in South Africa until the early 1970s and all historical material in local and overseas collections was opportunistically collected by other entomologists or explorers in the course of general collecting. Despite this, many specimens made their way to specialists and museums in Europe, most notably the specimen of Palpares speciosus described by Carl Linnaeus in 1758. He was a known correspondent of Ryk Tulbagh, Governor of the Kaap de Goede Hoop colony (now part of Western Cape Province) from 1751 to 1771 under the Dutch East India Company. Tulbagh is known to have sent specimens to Linnaeus (Giliomee 2013), and this is doubtless how the specimen ended up in the Linnaean collection, now in London. Carl Thunberg, a Swedish botanist and student of Linnaeus collected plants and insects in South Africa between 1772 and 1775 and travelled extensively during his three-year stay. In the process, he discovered and described three of the earliest known South African Neuroptera. Carl F. Drège, an apothecary in the Cape from about 1826, and an early insect collector and trader, sold lacewings to collectors in Europe, including the specimens described by Klug (1834). A significant number of lacewings were collected by Hans J. C. E. Brauns, a medical doctor in Willowmore between 1895 and 1929. His lacewing material is deposited in the Transvaal Museum, Pretoria (Ditsong Museum) (TMSA) and several European museums. Nesoleon braunsi (Banks) is named after him. Two local museums, TMSA and the South African Museum, Cape Town (Iziko Museum) (SAMC) added many lacewings to their collections from about 1883 onwards, as part of the by-catch of general collecting by museum staff or by light-trapping for Lepidoptera. These collectors included L. Péringuey, K. H. Barnard, V. B. Whitehead, S. van Noort, and anonymous “S.A. Museum Staff” (SAMC), and the well known Lepidopterists, A. J. T. Janse, G. van Son, and L. Vari (TMSA). Numerous donations were also made by other collectors including H. W. Bell-Marley, W. E. Jones, G. A. Alston, H. G. Wood, R. M. Lightfoot, R. F. Lawrence, J. S. Brown, C. W. Thorne (SAMC) and H. G. Breijer, J. H. Potgieter, C. J. Swierstra, A. Walton, M. J. Scoble and R. Toms (TMSA).
The 1951-1952 Lund University Expedition collected Neuroptera that provided the basis and impetus for Bo Tjeder’s fundamental studies of South African lacewings, together with the collections of the TMSA, SAMC and other South African institutions.
Subsequent to Tjeder’s studies there was a burgeoning interest in South African and Afrotropical lacewings, with the initiation of the Southern African Lacewing Survey by the current author, Mervyn W. Mansell, while a post-graduate student at Rhodes University in Grahamstown. Specimens from these ongoing surveys now form the basis of the lacewing collections of the South African National Collection, Plant Protection Research Institute, Pretoria (SANC). These holdings currently comprise about 50,000 pinned specimens housed in 600 drawers. These were the first surveys focussed explicitly on the Neuropterida of southern Africa and were compiled during numerous field trips to all countries of southern Africa and by many individual contributors. The SANC field trips in South Africa and Namibia from 1979-2004, yielded the bulk of the collections, while several simultaneous expeditions with American (L. A. Stange, R. B. Miller, J. D. Oswald, and J. B. Johnson), Austrian (H. Aspöck, U. Aspöck and H. Hölzel), German (P. Ohm), and Swiss (P. Duelli) colleagues have provided additional valuable study material and publications. The extensive field trip through South Africa and Namibia following the Third International Symposium on Neuropterology in 1988 included most of these colleagues and many others, and yielded much material documented by Mansell & Aspöck (1990).
Numerous colleagues and collectors have contributed significantly to the lacewing surveys over the years, including: J. B. Ball, H. S. Staude, D. M. Kroon, R. G. Oberprieler, C. H. Scholtz, E. Grobbelaar, C. D. Eardley, H. de Klerk, R. A. Adam, P. J. Ashton, D. K. Bakkes, C. L. Bellamy, P. Bayliss, H. D. Brown, M de Jager, N. J. S. Duke, E. Grei, M. E. Lee, L. R. Minter, C. G. E. Moolman, S., O., & W. Neser, R. D. Stephen, R. P. Urban, and especially A. K. Brinkman and A. P. Marais. Extensive Neuroptera surveys have also been undertaken in several National Parks: Kruger National Park by H., A., and L. E. O. Braack and M. W. Mansell; and Kalahari Gemsbok and Karoo National Parks by Mansell and colleagues.
Several collectors, J. B. Ball, and A. K. Brinkman (Cape Town); A. I. Curle, and H. C. Ficq (Johannesburg); and S. F. & G. A. Henning (Johannesburg) curate significant private collections of lacewings that add further to the South African discourse.
Finally, modern technology, by way of electronic relational databases have facilitated the compilation, coordination, and querying of data pertaining to lacewing specimens collected over more than two centuries and have now made accessible data that have previously been unavailable in numerous collections.
Methods and Resources
Studies of South African Neuropterida are based on field-collected specimens, data from the literature, and historical data associated with museum specimens from most of the major collections in southern Africa and Europe. All data are recorded, collated, and analysed in a relational database developed in Microsoft Access (Mansell & Kenyon 2002). This provides the basis for the Catalogue of Afrotropical Neuropterida, some contents of which are provided here in the NidaZA module of the Lacewing Digital Library. These data are also provided to the Animal Demography Unit, University of Cape Town (ADU) for the Lacewing Map project (Mansell 2014), and to the South African National Biodiversity institute (SANBI) (www.sanbi.org.za).
Material is collected by means of non-destructive umbrella light traps, by hand-net, and through rearing of larvae. Specimens are all pinned, labelled, and accessioned onto the database with a unique accession number for each lot (all specimens of the same species collected at the same locality and date). More recently, the middle and hind left legs of freshly-collected specimens are removed and stored in absolute ethyl alcohol in a deep-freeze refrigerator for future molecular analysis. These molecular samples are allocated the same accession number as the pinned voucher specimen. When published, the molecular sequences are provided to GenBank.