Advances in the systematics and ecology of African Corinnidae spiders (Arachnida: Araneae), with emphasis on the Castianeirinae

English: The Corinnidae is one of 76 families of spiders (Arachnida: Araneae) presently recognised in the Afrotropical Region. By the end of the last century their taxonomy and systematics had been very poorly studied and no modern revisions existed on the group. At that time, 110 species in 22 genera were known from the region, making it a family with moderate species richness. The description of the new genus Hortipes Bosselaers & Ledoux, 1998 in the family Liocranidae signalled the start of modern systematics studies in that family, and following the transfer of Hortipes to the Corinnidae, of that family by default too. Since that time, 20 taxonomic papers have been published on the Afrotropical Corinnidae and 10 new genera (all endemic to the region) and 164 new species have been described, of which three species form part of the current study (Chapters 7 and 9). Several genera have also been transferred to or from the Corinnidae in those papers. Presently there are 35 genera and more than 270 species known from the region, with the Corinnidae now ranking eighth in species richness in the region. Most of the revisionary work so far has focused on the subfamilies Trachelinae and Phrurolithinae, while the Corinninae sensu lato and Castianeirinae have largely remained neglected. The broad aim of the current study was to focus on the systematics of the latter group, treat the taxonomy of each of the currently known genera, at least in part, and provide a basis for future work on the subfamily. As such, many of the smaller genera in the subfamily were revised in the Afrotropical Region and two new genera were described. The genus Apochinomma Pavesi, 1881, the only described genus of accurate antmimicking castianeirines from the region, is revised and separated into two species groups based on genitalic and abdominal morphology. The type species, A. formicaeforme Pavesi, 1881, is redescribed and three new species are described in the A. formicaeforme species group: A. malkini sp. nov., A. parva sp. nov. and A. tuberculata sp. nov.. Two new species, A. decepta sp. nov. and A. elongata sp. nov., are described in the A. decepta species group, although an additional species only known from juveniles can also be placed in the latter group. Members of the A. formicaeforme species mimic Polyrhachis ants and are mainly arboreal, while members of A. decepta species group are ground- or grass-dwelling and probably mimic ponerine ants. The genus Cambalida Simon, 1909 is revised and three species are transferred from Castianeira Keyserling, 1879 to Cambalida: C. deminuta (Simon, 1909) comb. nov., C. fulvipes (Simon, 1896) comb. nov. and C. loricifera (Simon, 1885) comb. nov.. An additional species is transferred from Brachyphaea Simon, 1895 to Cambalida: C. fagei (Caporiacco, 1939) comb. nov.. All of these species are redescribed, as is Cambalida coriacea Simon, 1909. Two species, Castianeira depygata Strand, 1916 syn. nov. and C. mestrali Lessert, 1921 syn. nov., are considered junior synonyms of C. fulvipes. The type material of the type species of the genus, C. insulana Simon, 1909 from Annobon Island, is lost, and only immature specimens have been subsequently collected from a nearby island. The species is regarded as a nomen dubium until fresh adult material can be collected. A replacement name, Cambalida simoni nom. nov., is proposed for Cambalida fulvipes Simon, 1909, the latter being a secondary junior homonym of Cambalida fulvipes (Simon, 1896) comb. nov.. The type material of C. simoni is also lost and it too is considered a nomen dubium. Five new species are described: C. compressa sp. nov., C. dippenaarae sp. nov., C. griswoldi sp. nov., C. lineata sp. nov. and C. unica sp. nov.. Castianeira Keyserling, 1879 is the largest genus in the Corinnidae with 131 described species, of which 22 are presently known from the Afrotropical Region. There is a very rich undescribed fauna known from the region, and the variable morphology of its component species would suggest it is polyphyletic and should be divided into several genera. For example, six species are misplaced and have been transferred to or synonymised with species in Cambalida or the new genus Copuetta gen. nov.. In the present study, five species are redescribed and illustrated for the first time based on the type material: C. delicatula Simon, 1909, C. formosula Simon, 1909, C. majungae Simon, 1896, C. phaeochroa Simon, 1909 and C. thomensis Simon, 1909. The female holotype of C. bicolor (Simon, 1890) lacks an abdomen and the species is considered a nomen dubium. The types of several Afrotropical species could not be traced as yet and the species should be redescribed, if possible, based on recently collected material from near their type localities. The ground-dwelling genus Copa Simon, 1885 is one of four genera in the Afrotropical Region that have cryptic colouration that bears a resemblance to that of wolf spiders (Lycosidae), hereafter referred to as cryptic lycosiform colouration. The type species of the genus, C. flavoplumosa Simon, 1885, is redescribed and proposed as a senior synonym of C. benina Strand, 1916 syn. nov. and C. benina nigra Lessert, 1933 syn. nov.. This is possibly the most widespread corinnid in the Afrotropical Region albeit that is has not yet been recorded from any of the islands. A new species, C. kei sp. nov., is described from South Africa. Copa agelenina Simon, 1910, originally described from a subadult female from southern Botswana, is considered a nomen dubium. Although the Madagascan fauna was not included in this revision, nearly 30 new species have been distinguished from museum collections, and once that fauna is revised it will provide an exceptional example of island radiation. In a revision of the Afrotropical species of the ant-mimicking genus Corinnomma Karsch, 1880, Apochinomma semiglabrum Simon, 1896 is redescribed from both sexes, and based on these descriptions it is transferred to Corinnomma as C. semiglabrum (Simon, 1896) comb. nov.. A new species, C. lawrencei sp. nov., is described from Mozambique, Tanzania and South Africa. The taxonomic status of C. olivaceum Simon, 1896 is discussed and the first illustrations of the female genitalic structures are presented. Since no fresh material of this species is available and the female holotype is badly faded, it is not thoroughly redescribed. An English translation of Simon’s (1896) Latin description of C. olivaceum is provided with the intention of more accurately describing the colouration of this species. The arboreal cryptic lycosiform castianeirine genus Echinax Deeleman-Reinhold, 2001, previously known only from South-East Asia, is recorded from the Afrotropical Region for the first time. Copa longespina Simon, 1909 is redescribed and the species is transferred to Echinax as E. longespina (Simon, 1909) comb. nov.. Six new species are described from both sexes: E. clara sp. nov., E. hesperis sp. nov., E. natalensis sp. nov., E. scharffi sp. nov., E. similis sp. nov. and E. spatulata sp. nov.. The genus Graptartia Simon, 1896, presently known only from Africa, is revised. The type species, G. granulosa Simon, 1896, is redescribed and the first genitalic sketches of the species are provided. Two new species, G. mutillica sp. nov. and G. tropicalis sp. nov., are described. Unique amongst African castianeirines, all species of Graptartia are mimics of wingless female velvet ants (Mutillidae). Although the genus Merenius Simon, 1909 is not revised, a single common species, Merenius alberti Lessert, 1923, is redescribed. The species was previously known only from South Africa, and is recorded for the first time from Mozambique, Swaziland and Zimbabwe. While most populations of M. alberti comprise the typical black morph of the species, a red morph is described for the first time here. As part of a field study to identify the potential models of the two colour morphs of M. alberti, spiders were collected by hand and ants by pitfall trapping in the Ndumo Game Reserve in northern KwaZulu-Natal, South Africa. The ants assemblages sampled at 20 sites in the reserve seem to indicate that the black morph is a generalised mimic of black ground-dwelling ants, most likely Camponotus cinctellus (Gerstäcker, 1859), Streblognathus peetersi Robertson, 2002 and Polyrhachis gagates F. Smith, 1858, while the red morph is a mimic of Anoplolepis custodiens (F. Smith, 1858) ants. Lastly, the genus Messapus Simon, 1898, presently placed in the Castianeirinae, is reviewed. The type species, M. martini Simon, 1898, clearly represents two different species, one a corinnine (female lectotype) and the other a castianeirine (male paralectotype). The female is redescribed and the true male described for the first time. Based on the redescription, Messapus is transferred to the Corinninae. Corinna natalis Pocock, 1898 is misplaced and is transferred to Messapus as M. natalis (Pocock, 1898) comb. nov., while Messapus secundus Strand, 1907 is misplaced and transferred to Merenius as M. secundus (Strand, 1907) comb. nov.. A new cryptic lycosiform genus, Copuetta gen. nov., with the type species C. maputa sp. nov., is established to accommodate the castianeirine male paralectotype of M. martini, and its matching female is described for the first time. Castianeira kibonotensis Lessert, 1921 syn. nov. is considered a junior synonym of Copa lacustris Strand, 1916 and the species is redescribed and transferred to Copuetta gen. nov. as C. lacustris (Strand, 1916) comb. nov.. An additional eleven new species of Copuetta gen. nov. are described: C. comorica sp. nov., C. erecta sp. nov., C. kakamega sp. nov., C. kwamgumi sp. nov., C. lesnei sp. nov., C. litipo sp. nov., C. lotzi sp. nov., C. magna sp. nov., C. naja sp. nov., C. uzungwa sp. nov. and C. wagneri sp. nov.. A second new cryptic lycosiform genus, Wasaka gen. nov., is described for four new species from tropical Africa: W. imitatrix sp. nov., W. montana sp. nov., W. occulta sp. nov. (type species) and W. ventralis sp. nov.. A phylogenetic analysis of the subfamily Castianeirinae from the Afrotropical Region was carried out. Forty-one ingroup taxa (Castianeirinae) were included, of which 39 were Afrotropical, one Australasian and one Brazilian. Outgroup taxa included three species of Trachelinae, one Phrurolithinae, two Corinninae and two Corinnidae incertae sedis, with Drassodes sesquidentatus Purcell, 1908 used to root the trees. Only species of Castianeirinae treated in this thesis and known from both sexes were included in the analysis. Analyses performed in Winclada, TNT and PAST all produced similar but very unsatisfactory results, with the outgroups grouping together with part of the Castianeirinae. Consequently, a second analysis was conducted with the exclusion of most of the outgroup taxa (except D. sesquidentatus and Corinninae). These results improved the resolution of the results considerably. but still did not resolve the placement of the Medmassa–Messapus clade within Castianeirinae; these genera can be considered to belong to Corinninae and should hypothetically have been placed outside the Castianeirinae clade. A single analysis produced in PAST produced the most parsimonious tree, with Medmassa–Messapus placed outside the Castianeirinae and each of the Afrotropical castianeirine genera as monophyletic. The results are inadequate to support any systematic changes in the Corinnidae, but future analyses need to include a more diverse range of castianeirine genera from outside the Afrotropical Region to better understand the relationships of the Afrotropical fauna. In the final chapter, the role of Castianeirinae as components of arthropod mimicry complexes is described for three species of ants, Anoplolepis custodiens (F. Smith, 1858), Polyrhachis gagates F. Smith, 1858 and Camponotus fulvopilosus (De Geer, 1778). There are respectively two out of 10, four out of six, and zero out of five species of Castianeirinae forming part of the arthropod complexes associated with these ants. All of these castianeirines are inaccurate (weak/ generalised) mimics of their models except for Apochinomma formicaeforme, which is an accurate (good/specialised) mimic of P. gagates. Colour polymorphism is also described for the first time in four species of Afrotropical Castianeirinae, i.e. Corinnomma semiglabrum, Merenius alberti, Castianeira cf. venustula (Pavesi, 1895) and Copa flavoplumosa. Three of these species are inaccurate mimics of ants, while C. flavoplumosa is a species with a widespread variant with cryptic lycosiform colouration and a nigrito form restricted mainly to tropical forests. High Castianeirinae biodiversity and endemism corresponds to most of the main Biodiversity Hotspots and Centres of Endemism (CE) in the Afrotropical Region: Maputaland-Pondoland-Albany CE (five endemics), Madagascar and Indian Ocean Islands CE (>30 endemics), East African Afromontane Forests CE (four endemics), East African Coastal Forests CE (five endemics), Guinean Forests of West Africa CE (seven endemics) and the Horn of Africa CE (one endemic). No endemic castianeirines have been recorded in the Succulent Karoo and Cape Floristic Region CE’s in southern Africa, although this corinnid fauna of these two CE’s is largely dominated by Trachelinae, most of which are endemics.