Araneae: Salticidae)

FOUR JUMPING SPIDER GENERA OF THE COCALODES-GROVV ARE MONOPHYLETIC WITH GENERA OF THE SPARTAEINAE (ARANEAE: SALTICIDAE) A.C RODRIGO1 & R.R. JACKSON2 department of Zoology, School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New Zealand & 2Department of Zoology, University of Canterbury, Private Bag, Christchurch, New Zealand. (Received 6 January 1992; revised and accepted 30 March 1992) ABSTRACT Rodrigo, A.G. & Jackson, R.R. (1992). Four jumping spider genera of the Cocalodes-group are monophyletic with genera of the Spartaeinae (Araneae: Salticidae). New Zealand Natural Sciences 19: 61-67. Wanless (1984a) erected a new sub-family, the Spartaeinae, to contain all members of the incorrectly named Boethinae except the genera Allococalodes, Cocalodes, Holcolaetis, and Sonoita. These were excluded from the sub-family because they possessed a median tegular apophysis on the male palp, a feature which Wanless believed to be synapomorphy of the group, and one which separated the group from members of the Spartaeinae. Our phylogenetic analysis of the extant members of the Spartaeinae, and the "Cocalodes-group" indicate that the latter are monophyletic with other genera of the sub-family. An amended definition of the sub-family is provided. KEYWORDS: Spartaeinae - "Cocalodes-group" - phylogeny - monophyly - quasi-polymorphic coding. INTRODUCTION however, that these genera are closely affiliated with members of the Spartaeinae and another Wanless (1984a) erected the jumping spider allied sub-family, the Lyssomaninae (Wanless sub-family Spartaeinae to accomodate members 1984a). of the incorrectly named Boethinae. He also These three groups are considered to be removed the genera Cocalodes Pocock, Sonoita primitive salticids (Blest & Carter 1987), and our Peckham & Peckham, and Holcolaetis Simon interest in them stems from a desire to test the from the sub-family on the grounds that males of hypothesis first proposed by Jackson & Blest these genera possess a median apophysis on the (1982) that jumping spiders evolved from web- palpal tegulum, a feature not possessed by other building ancestors. As a consequence, we recon genera of the Spartaeinae. Wanless (1984a) sug structed the phylogeny of the Spartaeinae using gested that another sub-family should be erected morphological characters, and we included the to contain these aberrant genera and a closely related genus, Allococalodes Wanless, and he in genera Cocalodes, Allococalodes, Holcolaetis, and formally referred to them collectively as the "Co- Sonoita in the analysis. Our results indicate that calodes-group". He was later to reassess this the "Cocalodes-gpoxxip" forms a monophyletic claim on the grounds that Holcolaetis and Sonoita clade within the Spartaeinae and, in accordance share little in common with Cocalodes and Alloco with current phylogenetic systematic practice, calodes, apart from the presence of a tegular should be included as members of the sub-family. median apophysis, the homology of which he Here, we describe the phylogenetic analysis, as doubted (Wanless 1985). There is little doubt, well as the implications of its results. 62 New Zealand Natural Sciences 19 (1992) METHODS AND MATERIALS or more groups, with at least two groups having more than one taxon as a member, were used. TAXA STUDIED Characters which satisfy this criterion are phylo Descriptions ofthe taxa listed in Table 1 were genetically informative (sensu Hennig 1965). obtained from published literature (cited in Table Characters which are monomorphic for all taxa 1). A list of characters were obtained from these offer no information on within-group relation descriptions and these are described below. ships. Characters for which there is only one partition with more than one taxon are similarly MORPHOLOGICAL CHARACTERS uninformative, for it implies that all other taxa The following morphological characters were possess unique character states (ie. autapomor- used to construct a taxon-character dataset. The phies). reader is directed to the excellent papers by Wanless (1984a, 1984b) for more complete de 1. Posterior median eyes (PMEs): 1-small and scriptions ofthe characters. The method of maxi vestigial; O-large (Wanless 1984a, p. 137). mum parsimony (Kluge & Farris 1969, Wiley Only some extant and extinct (ie. amber fossil) 1981) was used to reconstruct the phylogeny ofthe members of the Spartaeinae, Lyssomaninae group. As is appropriate for this method, only and Cocalodes-group possess large and func characters whose states partition the taxa into two tional PMEs. These are vestigial in other advanced salticids. Wanless (1984a) notes that with the exception of Taraxella, all salti Table 1. List of genera studied. Asemonea is a member ofthe cids can be placed easily into one or the other Lyssomaninae, and was included as an outgroup. Allococal- category. odes, Cocalodes, Holcolaetis, and Sonoita have been infor 2. PME in relation to anterior median/anterior mally referred to by Wanless as the "Cocalodes-group". lateral eyes (AMEs/ALEs) and posterior lat eral eyes (PLEs): 1-closer to AME/ALEs GENUS REFERENCE than to PLEs; 0-closer to PLE (Wanless 1980, p. 217). The closeness of the PMEs to the ALLOCOCALODES Wanless Wanless 1982 anterior eyes is most marked in the lyssoman ASEMONEA Wanless 1980 ine genera Asemonea O.P.-Cambridge, Go- BRETTUS Thorell Wanless 1979 leba Wanless, Macopaeus Simon, and Pan COCALODES Pocock Wanless 1982 disus Simon, where the tubercles on which the COCALUS Koch Wanless 1981a PMEs are situated abut those which hold the CYRBA Simon Wanless 1984b ALEs. The PMEs of some spartaeine genera GELOTIA Thorell Wanless 1984a are also found close to the ALEs, although not Wijesinghe, 1991b to the extent found in the lyssomanine genera. HOLCOLAETIS Simon Wanless 1985 3. Femoral organ: 1-present; 0-absent (Wanless MELEON Wanless Wanless 1984a 1984a p. 139). This structure takes the form of MINTONIA Wanless Wanless 1984a a perforated region situated either on a tu NEOBRETTUS Wanless Wanless 1984a bercle or in a gully on the underside of the PHAECIUS Simon Wanless 1981b femora of Legs I in the males of some genera. Wijesinghe 1991a An amorphous exudate is often associated PORTIA Karsch Wanless 1978 with the organ, and it may have a pheromone Wanless 1984a secreting function. SONOITA Peckham & Peckham Wanless 1985 4. Minute tarsal and metatarsal setae: 1-present; SPARTAEUS Thorell Wanless 1984a 0-absent (Wanless 1984a, p. 169; 1984b, p. Wanless 1987 450). Numerous minute setae may be found TARAXELLA Wanless Wanless 1984a on the underside of tarsi and metatarsi of Wanless, 1987 some genera. VEISSELLA Wanless Wanless 1984a 5. Retrolateral tibial apophysis (RTA): 1-mem- YA GINUMANIS Wanless Wanless 1984a braneous base present; 0-membraneous base A.G. Rodrigo & R.R. Jackson: Phylogeny of jumping spiders of Cocalodes-group 63 absent (Wanless 1984a, p. 139). The RTA is presence of structures (30-40 um in length) found on the male palpal tibia. The presence which resemble mussel shell valves. These of a membraneous base to the RTA allows it fields are situated on the dorsal surface ofthe to have some mobility. This feature is only abdomen, and are associated with pores of present in some genera. some form or other, suggesting a secretory 6. Ml apophysis of the distal haematodocha: 0- function. They are difficult to find in all but absent, 1-minute lobe, 2-large and petal-like, fresh specimens, and have only been found in 3-long and filamentous, 4-membraneous some species. We have been conservative in patch (Wanless 1984a, p. 140). The distal our estimation of their presence, and have haematodocha of the male palps of some only recorded them as present in those genera genera are made up of two apophyses which with species which are known to possess these are labelled by Wanless (1984a) as Ml and fields. All other genera have been treated as M2. The reader is referred to Wanless not possessing these fields. (1984a) for more complete descriptions of 12. Cymbial flange: 0-absent; 1-present (Wanless these structures, but it is noteworthy that these 1978, p. 84). A flange-like protuberance is apophyses are absent from the Cocalodes- found on the pro- or retrolateral dorsal sur group. Wanless (1984a) therefore suggested face at the base of the cymbium of some that the presence of Ml and M2 may be a genera. synapomorphy (ie. a shared-derived charac 13-16. Number of pro-marginal teeth on chelic ter) of the Spartaeinae. erae (Wanless 1984a, p. 138). Salticids, like 7. Median tegular apophysis: 1-present; 0-ab many other spiders, possess dentate armature sent (Wanless 1982, p. 264; Wanless 1984a, p. on their chelicerae. The numbers of these 141). This apophysis takes the form of a bifid cheliceral teeth have often been used as di prong arising from the palpal tegulum, and is agnostic characters. However, because ofthe only found in members of the "Cocalodes- variability in numbers of these teeth between group", as well as some lyssomanine genera. species within genera, such characters have 8. Tegular furrow: 0-lacking; l-short and shal recently been rejected as having systematic low; 2-deep (Wanless 1984a, p. 140). Part of value. Nonetheless, while the numbers of the tegulum, usually found on its dorsal side. cheliceral teeth may be variable, this variabil It is often obscured by other structures of the ity, as measured by its range, may itself be palp, and information on this feature is not phylogenetically informative. always available. Wanless (1984a)

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