Charles R. Haddad

Phylogeny of entelegyne spiders: Affinities of the family Penestomidae (NEW RANK), generic phylogeny of Eresidae, and asymmetric rates of change in spinning organ evolution (Araneae, Araneoidea, Entelegynae)

Penestomine spiders were first described from females only and placed in the family Eresidae. Discovery of the male decades later brought surprises, especially in the morphology of the male pedipalp, which features (among other things) a retrolateral tibial apophysis (RTA). The presence of an RTA is synapomorphic for a large clade of spiders exclusive of Eresidae. A molecular data matrix based on four loci was constructed to test two alternative hypotheses: (1) penestomines are eresids and the RTA is convergent, or (2) penestomines belong within the RTA clade. Taxon sampling concentrated on the Eresidae and the RTA clade, especially outside of the Dionycha and Lycosoidea. Evolution of the cribellum, conventionally characterized as a primitive araneomorph spinning organ lost multiple times, is explored. Parsimony optimization indicates repeated appearances of the cribellum. Exploration of asymmetric rates of loss and gain in both a likelihood framework and using a Sankoff matrix under parsimony reveals that cribellum homology is supported when losses are two times more likely than gains. We suggest that when complicated characters appear (under parsimony optimization) to evolve multiple times, investigators should consider alternative reconstructions featuring a relatively high rate of loss. Evolution of other morphological characters is also investigated. The results imply revised circumscription of some RTA-clade families, including Agelenidae, Amaurobiidae, Cybaeidae, Dictynidae and Hahniidae. Some nomenclatural changes are formally proposed here; others await further investigation. The family Penestomidae (NEW RANK) is established. Tamgrinia, not Neoramia, is the cribellate sister clade of the ecribellate Agelenidae. Tamgrinia and the subfamily Coelotinae are transferred from the family Amaurobiidae to the family Agelenidae. Zanomys and its relatives are not coelotines but belong to a clade tentatively identified as Macrobuninae.

Jumping Spiders (Araneae: Salticidae) of the Ndumo Game Reserve, Maputaland, South Africa

ABSTRACT Salticids from the Ndumo Game Reserve (KwaZulu-Natal, South Africa) are described. In total 72 species of 38 genera were found, of which one genus Aenigma gen. n. and 14 species are described as new: Aenigma incognita sp. n., Bianor eximius sp. n., Evarcha mirabilis sp. n., E. striolata sp. n., Habrocestum africanum sp. n., Icius nigricaudus sp. n., Massagris natalensis sp. n., Phlegra arborea sp. n., Ph. certa sp. n., Pignus pongola sp. n., Pseudicius venustulus sp. n., Rhene pinguis sp. n., Thyenula fidelis sp. n., and Th. magna sp. n. Three specific names are synonymised: Cyrba armata Wesołowska, 2006 with C. lineata Wanless, 1984; Pellenes pulcher Wesołowska, 1999 (junior homonym of P. pulcher Logunov, 1995) with P. tharinae Wesołowska, 2006; and Thyene strandi Caporiacco, 1939 with Th. natalii Peckham & Peckham, 1903. A new combination, Evarcha annae (ex Habrocestum annae Peckham & Peckham, 1903) is proposed. Twenty species are recorded from South Africa for the first time: Asemonea Stella Wanless, 1980; Cyrba boveyi Lessert, 1933; Evarcha elegans Wesołowska & Russell-Smith, 2000; E. ignea Wesołowska & Cumming, 2008; E. prosimilis Wesołowska & Cumming, 2008; Goleba puella (Simon, 1885); Heliophanus fascinatus Wesołowska, 1986; H. pauper Wesołowska, 1986; Hispo georgius (Peckham & Peckham, 1892); Menemerus minshullae Wesołowska, 1999; Modunda staintoni (O.P.-Cambridge, 1872); Myrmarachne lulengana Roewer, 1965; Nigorella plebeja (L. Koch, 1875); Pellenes epularis (O.P.-Cambridge, 1872); P. bulawayoensis Wesołowska, 1999; P. tharinae Wesołowska, 2006; Pseudicius alter Wesołowska, 1999; Schenkelia modesta Lessert, 1927; Thyene bucculenta (Gerstaecker, 1873); and Th. semiargentea (Simon, 1884). The rich diversity of Salticidae collected in this study, and the proportion of new species relative to the total jumping spider fauna (19%), highlight the need to maintain conservation efforts within Maputaland protected areas with management plans that also take invertebrate taxa into consideration.

Discovery of a monophagous true predator, a specialist termite-eating spider (Araneae: Ammoxenidae)

True predators are characterised by capturing a number of prey items during their lifetime and by being generalists. Some true predators are facultative specialists, but very few species are stenophagous specialists that catch only a few closely related prey types. A monophagous true predator that would exploit a single prey species has not been discovered yet. Representatives of the spider family Ammoxenidae have been reported to have evolved to only catch termites. Here we tested the hypothesis that Ammoxenus amphalodes is a monophagous termite-eater capturing only Hodotermes mossambicus. We studied the trophic niche of A. amphalodes by means of molecular analysis of the gut contents using Next Generation Sequencing. We investigated their willingness to accept alternative prey and observed their specific predatory behaviour and prey capture efficiency. We found all of the 1.4 million sequences were H. mossambicus. In the laboratory A. amphalodes did not accept any other prey, including other termite species. The spiders attacked the lateral side of the thorax of termites and immobilised them within 1 min. The paralysis efficiency was independent of predator:prey size ratio. The results strongly indicate that A. amphalodes is a monophagous prey specialist, specifically adapted to feed on H. mossambicus.

THE INFLUENCE OF MOUND STRUCTURE ON THE DIVERSITY OF SPIDERS (ARANEAE) INHABITING THE ABANDONED MOUNDS OF THE SNOUTED HARVESTER TERMITE TRINERVITERMES TRINERVOIDES

Abstract The dynamics of spiders present in abandoned Trinervitermes trinervoides (Sjöstedt) termite mounds were studied over a period of one year, from March 1999 to January 2000, with five mounds excavated on a bimonthly basis. All spiders present in the mound were collected by hand and preserved in 70% ethanol. A total of 771 spiders represented by 21 families and 82 species were collected from the 30 mounds during the course of the study. The most abundant were the Gnaphosidae, which represented 37.87% of all spiders collected, followed by the Salticidae (12.97%), Pholcidae (10.51%) and Oonopidae (9.60%). These were the only families that represented more than 5% of the spider fauna. The most abundant species were Zelotes fuligineus (Purcell 1907) (Gnaphosidae) (11.69%), Smeringopus sambesicus Kraus 1957 (Pholcidae) (10.51%), Heliophanus sp. (Salticidae) (9,86%) and a Gamasomorphinae sp. (Oonopidae) (9.21%). A correlation was found between spider abundance and mound height, surface perforation of the mound and season of collection. Spider numbers were highest in mounds with a high surface degradation, while a tendency existed for an increase in numbers with increased mound height. Web-building spiders (Pholcidae and Theridiidae) were largely limited to mounds with a cavity in the structure.

Three New Genera of Tracheline Sac Spiders from Southern Africa (Araneae: Corinnidae)

ABSTRACT Three new genera of sac spiders of the subfamily Trachelinae (Corinnidae) are described from southern Africa. Fuchiba gen. n. is represented by F. aquilonia sp. n. (type species), F. capensis sp. n., F. montana sp. n., F. similis sp. n., F. tortilis sp. n. and F. venteri sp. n. All species are endemic to South Africa except for F. aquilonia sp. n., which is also recorded from Mozambique and Botswana, and F. montana sp. n., which is known from Lesotho. Fuchibotulus gen. n. is represented by F. bicornis sp. n. from South Africa (type species) and F. kigelia sp. n. from South Africa and southern Mozambique, Poachelas gen. n. is represented by three new species, and two species groups are recognised based on leg morphology. The striatus species group is represented by P. striatus sp. n. (type species, South Africa) and P. solitarius sp. n. (Zimbabwe). The montanus species group, represented by P. montanus sp. n., is endemic to South Africa. The three genera show different microhabitat preferences. Representatives of Fuchiba gen. n. and Fuchibotulus gen. n. were collected primarily by pitfall trapping or by hand from leaf litter in various biomes, while representatives of Poachelas gen. n. were collected primarily from the base of grass tussocks, or occasionally by sweep-netting in grassland and savannah. Identification keys to new species are provided for each genus.

The faunistic diversity of spiders (Arachnida: Araneae) of the Savanna Biome in South Africa

Invertebrates include more than 80% of all animals, yet they are severely under-represented in studies of southern African diversity. Site biodiversity estimates that ignore invertebrates, not only omit the greatest part of what they are attempting to measure, but also neglect major contributors to essential ecosystem processes. All available information on spider species distribution in the South African Savanna Biome was compiled. This is the largest biome in the country, occupying over one third of the surface area. A total of 23 739 records from 1260 localities were recorded in the South African Savanna Biome until the end of 2010. This included 1230 species represented by 381 genera and 62 families. The last decade has seen an exponential growth in the knowledge of the group in South Africa, but there certainly are several more species that have to be discovered, and the distribution patterns of those listed are partly unknown. Information is summarised for all quarter-degree squares of the biome and reveals considerable inequalities in knowledge. At a large scale the eastern region is much better surveyed than the western parts, but at finer scales throughout the region, several areas have little information. The Salticidae is the most diverse family (162 spp.) and also has the most endemic savanna species (42 spp.). An endemicity index indicates that 366 species are endemic to the biome, with 322 species that are near endemics, i.e., also occurring in an adjacent biome. An abundance index (1–3) was also calculated for each species based on numbers sampled. A rarity index for each species gives a preliminary indication of their conservation importance. Patterns of guild composition are summarised and species known to play a role as predators in agro-ecosystems found within the biome are also discussed.

Trophic strategy of ant-eating Mexcala elegans (Araneae: Salticidae): looking for evidence of evolution of prey-specialization

Abstract We investigated the trophic strategy of Mexcala elegans Peckham & Peckham 1903, an ant-eating salticid spider from South Africa, in order to gain baseline information concerning the evolution of prey specialization. We studied its natural prey, prey acceptance, and choice using a variety of prey species. In its natural habitat, the spider captured only ants, mainly its mimetic model Camponotus cinctellus, indicating that the species is a stenophagous ant-eater. However, in the laboratory, M. elegans captured 12 different invertebrate taxa with efficiency similar to the capture of ants, suggesting that it is euryphagous. For the capture of ants but not for other prey, it used a specialized prey-capture behavior. In prey-choice experiments, the spiders did not prefer ants to flies. We found no evidence for neural and behavioral constraints related to identification and handling of prey. Our results suggest that M. elegans is a euryphagous specialist using a specialized ant-eating capture strategy in which prey specialization has evolved as a byproduct of risk aversion (“enemy-free space” hypothesis).

South African Spider Diversity: African Perspectives on the Conservation of a Mega-Diverse Group

Any field of endeavour requires retrospection after a period of substantial activity. This process provides a measure of what has been achieved and identifies future directions. Studies of spider diversity in South Africa have gone through an intense growth phase over the past ten years and reached a stage in its development where reflections on patterns and processes observed could provide meaningful input into the identification of further work. This chapter establishes the background and framework for such a discussion on the path to a more holistic conservation planning that includes invertebrates. Invertebrate conservation and diversity pose a significant challenge to planners and managers (Engelbrecht, 2010), and in spite of the central role that insects and arachnids play in terrestrial biodiversity, they still remain peripheral to decision-making processes. The reality is that, for Africa in particular, there are very few conservation areas that have both the resources and expertise to include invertebrates as part of their monitoring and management initiatives (however, see South African River Health Programme1). The advent of adaptive management, with a strong emphasis on experimental implementation of alternative management options (Johnson, 1999), has informed much of recent thinking and has cast a dim light on classical inventory studies that generate species lists. However, records of the numbers of species and their distribution provides a fundamental starting point for the conservation of biodiversity (Pullin, 2002). This view also ignores the contribution that basic inventories and alpha taxonomy make to the initial development of a field. This chapter will show that South African spider systematics and ecology are in an exploratory phase, and that traditional approaches to mapping diversity has enabled spider ecology in the country to generate species lists that are often resolved up to species level. Very few other studies on mega-diverse invertebrate groups in Africa can match this taxonomic resolution (see e.g. Formicinae). This descriptive phase will provide the

A revision of the tracheline sac spider genus Cetonana Strand, 1929 in the Afrotropical Region, with descriptions of two new genera (Araneae : Corinnidae)

ABSTRACT The dark sac spiders of the genus Cetonana Strand, 1929 (Corinnidae: Trachelinae) of the Afrotropical Region are revised. Following a detailed comparison of the somatic and genitalic morphology of the Afrotropical C. martini (Simon, 1897) with the European type species, C. laticeps (Canestrini, 1868), Afroceto gen. n. is established to include the transfer of two Afrotropical species previously described in Cetonana, namely A. martini (Simon, 1897) comb. n. (type species) and A. coenosa (Simon, 1897) comb. n. Three species are proposed as junior synonyms of A. martini, namely C. curvipes (Tucker, 1920) syn. n., C. tridentata (Lessert, 1923) syn. n. and C. simoni (Lawrence, 1942) syn. n. The type of C. aculifera (Strand, 1916) from Madagascar is presumed destroyed and this species is considered nomen dubium. Additionally, 12 new Afroceto gen. n. species are described from southern Africa, namely A. arca sp. n., A. bulla sp. n., A. bisulca sp. n., A. capensis sp. n., A. corcula sp. n., A. croeseri sp. n., A. flabella sp. n., A. gracilis sp. n., A. plana sp. n., A. porrecta sp. n., A. rotunda sp. n. and A. spicula sp. n. Afroceto gen. n. species display variable ecological preferences, occurring in tree canopies, lower foliage strata, bark, and on the ground in contrasting habitats (forests, savannah, grassland, karoo and fynbos) in southern Africa, with single records from Malawi and Tanzania. Patelloceto gen. n., closely resembling Afroceto gen. n. but distinguished by the reduced leg spination and distinctive genitalic morphology, is also described, with three new species, P. secutor sp. n. (type species), P. denticulata sp. n. and P. media sp. n. from southern, central and east Africa. Patelloceto gen. n. species are primarily arboreal, occurring in tree canopies, lower foliage strata or on bark in forests and savannahs.

A revision of the purse-web spider genus Calommata Lucas, 1837 (Araneae, Atypidae) in the Afrotropical Region

Abstract The purse-web spider genus Calommata Lucas, 1837 is revised in the Afrotropical Region. Following examination of the female type material, Calommata transvaalica Hewitt, 1916 is removed from synonymy with Calommata simoni Pocock, 1903 and revalidated. The females of both species are redescribed and their males described for the first time. While Calommata simoni is very widespread across tropical Africa, Calommata transvaalica is endemic to northern South Africa. Four new species are described, all known only from males: Calommata megae sp. n. (Zimbabwe), Calommata meridionalis sp. n. (South Africa), Calommata namibica sp. n. (Namibia) and Calommata tibialis sp. n. (Ivory Coast and Togo). Notes are presented on the biology of each species.