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

[摘要] English: The Corinnidae is one of 76 families of spiders (Arachnida: Araneae) presentlyrecognised in the Afrotropical Region. By the end of the last century their taxonomy andsystematics had been very poorly studied and no modern revisions existed on the group. Atthat time, 110 species in 22 genera were known from the region, making it a family withmoderate species richness. The description of the new genus Hortipes Bosselaers & Ledoux,1998 in the family Liocranidae signalled the start of modern systematics studies in thatfamily, 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 AfrotropicalCorinnidae and 10 new genera (all endemic to the region) and 164 new species have beendescribed, of which three species form part of the current study (Chapters 7 and 9). Severalgenera have also been transferred to or from the Corinnidae in those papers. Presently thereare 35 genera and more than 270 species known from the region, with the Corinnidae nowranking eighth in species richness in the region. Most of the revisionary work so far hasfocused on the subfamilies Trachelinae and Phrurolithinae, while the Corinninae sensu latoand Castianeirinae have largely remained neglected. The broad aim of the current study wasto focus on the systematics of the latter group, treat the taxonomy of each of the currentlyknown 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 twonew genera were described.The genus Apochinomma Pavesi, 1881, the only described genus of accurate antmimickingcastianeirines from the region, is revised and separated into two species groupsbased on genitalic and abdominal morphology. The type species, A. formicaeforme Pavesi,1881, is redescribed and three new species are described in the A. formicaeforme speciesgroup: 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 lattergroup. Members of the A. formicaeforme species mimic Polyrhachis ants and are mainlyarboreal, while members of A. decepta species group are ground- or grass-dwelling andprobably mimic ponerine ants.The genus Cambalida Simon, 1909 is revised and three species are transferred fromCastianeira Keyserling, 1879 to Cambalida: C. deminuta (Simon, 1909) comb. nov., C. fulvipes (Simon, 1896) comb. nov. and C. loricifera (Simon, 1885) comb. nov.. Anadditional species is transferred from Brachyphaea Simon, 1895 to Cambalida: C. fagei(Caporiacco, 1939) comb. nov.. All of these species are redescribed, as is Cambalidacoriacea 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 typematerial of the type species of the genus, C. insulana Simon, 1909 from Annobon Island, islost, 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. Areplacement 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 nomendubium. 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 131described species, of which 22 are presently known from the Afrotropical Region. There is avery rich undescribed fauna known from the region, and the variable morphology of itscomponent species would suggest it is polyphyletic and should be divided into severalgenera. For example, six species are misplaced and have been transferred to or synonymisedwith species in Cambalida or the new genus Copuetta gen. nov.. In the present study, fivespecies 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 ofseveral 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 theAfrotropical Region that have cryptic colouration that bears a resemblance to that of wolfspiders (Lycosidae), hereafter referred to as cryptic lycosiform colouration. The type speciesof the genus, C. flavoplumosa Simon, 1885, is redescribed and proposed as a senior synonymof C. benina Strand, 1916 syn. nov. and C. benina nigra Lessert, 1933 syn. nov.. This ispossibly the most widespread corinnid in the Afrotropical Region albeit that is has not yetbeen recorded from any of the islands. A new species, C. kei sp. nov., is described fromSouth Africa. Copa agelenina Simon, 1910, originally described from a subadult female fromsouthern Botswana, is considered a nomen dubium. Although the Madagascan fauna was not included in this revision, nearly 30 new species have been distinguished from museumcollections, and once that fauna is revised it will provide an exceptional example of islandradiation.In a revision of the Afrotropical species of the ant-mimicking genus CorinnommaKarsch, 1880, Apochinomma semiglabrum Simon, 1896 is redescribed from both sexes, andbased 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, Tanzaniaand South Africa. The taxonomic status of C. olivaceum Simon, 1896 is discussed and thefirst illustrations of the female genitalic structures are presented. Since no fresh material ofthis species is available and the female holotype is badly faded, it is not thoroughlyredescribed. An English translation of Simon's (1896) Latin description of C. olivaceum isprovided 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 Regionfor the first time. Copa longespina Simon, 1909 is redescribed and the species is transferredto Echinax as E. longespina (Simon, 1909) comb. nov.. Six new species are described fromboth 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. Thetype species, G. granulosa Simon, 1896, is redescribed and the first genitalic sketches of thespecies are provided. Two new species, G. mutillica sp. nov. and G. tropicalis sp. nov., aredescribed. Unique amongst African castianeirines, all species of Graptartia are mimics ofwingless 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 fromSouth 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 redmorph is described for the first time here. As part of a field study to identify the potentialmodels of the two colour morphs of M. alberti, spiders were collected by hand and ants bypitfall trapping in the Ndumo Game Reserve in northern KwaZulu-Natal, South Africa. Theants assemblages sampled at 20 sites in the reserve seem to indicate that the black morph is ageneralised 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, isreviewed. The type species, M. martini Simon, 1898, clearly represents two different species,one a corinnine (female lectotype) and the other a castianeirine (male paralectotype). Thefemale is redescribed and the true male described for the first time. Based on theredescription, Messapus is transferred to the Corinninae. Corinna natalis Pocock, 1898 ismisplaced and is transferred to Messapus as M. natalis (Pocock, 1898) comb. nov., whileMessapus 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 thetype species C. maputa sp. nov., is established to accommodate the castianeirine maleparalectotype 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 Copalacustris Strand, 1916 and the species is redescribed and transferred to Copuetta gen. nov. asC. 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 crypticlycosiform 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 Regionwas carried out. Forty-one ingroup taxa (Castianeirinae) were included, of which 39 wereAfrotropical, one Australasian and one Brazilian. Outgroup taxa included three species ofTrachelinae, one Phrurolithinae, two Corinninae and two Corinnidae incertae sedis, withDrassodes sesquidentatus Purcell, 1908 used to root the trees. Only species of Castianeirinaetreated in this thesis and known from both sexes were included in the analysis. Analysesperformed in Winclada, TNT and PAST all produced similar but very unsatisfactory results,with the outgroups grouping together with part of the Castianeirinae. Consequently, a secondanalysis was conducted with the exclusion of most of the outgroup taxa (except D.sesquidentatus and Corinninae). These results improved the resolution of the resultsconsiderably. but still did not resolve the placement of the Medmassa–Messapus clade withinCastianeirinae; these genera can be considered to belong to Corinninae and shouldhypothetically have been placed outside the Castianeirinae clade. A single analysis producedin PAST produced the most parsimonious tree, with Medmassa–Messapus placed outside theCastianeirinae and each of the Afrotropical castianeirine genera as monophyletic. The results are inadequate to support any systematic changes in the Corinnidae, but future analyses needto include a more diverse range of castianeirine genera from outside the Afrotropical Regionto better understand the relationships of the Afrotropical fauna.In the final chapter, the role of Castianeirinae as components of arthropod mimicrycomplexes is described for three species of ants, Anoplolepis custodiens (F. Smith, 1858),Polyrhachis gagates F. Smith, 1858 and Camponotus fulvopilosus (De Geer, 1778). There arerespectively two out of 10, four out of six, and zero out of five species of Castianeirinaeforming part of the arthropod complexes associated with these ants. All of thesecastianeirines are inaccurate (weak/ generalised) mimics of their models except forApochinomma formicaeforme, which is an accurate (good/specialised) mimic of P. gagates.Colour polymorphism is also described for the first time in four species of AfrotropicalCastianeirinae, 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 lycosiformcolouration and a nigrito form restricted mainly to tropical forests.High Castianeirinae biodiversity and endemism corresponds to most of the mainBiodiversity Hotspots and Centres of Endemism (CE) in the Afrotropical Region:Maputaland-Pondoland-Albany CE (five endemics), Madagascar and Indian Ocean IslandsCE (>30 endemics), East African Afromontane Forests CE (four endemics), East AfricanCoastal 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 inthe Succulent Karoo and Cape Floristic Region CE's in southern Africa, although thiscorinnid fauna of these two CE's is largely dominated by Trachelinae, most of which areendemics.

[发布机构] University of the Free State