Zootaxa 2021: 1–22 (2009) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition) New cocalodine jumping spiders from Papua New Guinea (Araneae: Salticidae: Cocalodinae) WAYNE P. MADDISON Departments of Zoology and Botany and Beaty Biodiversity Museum, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia, V6T 1Z4, Canada. E-mail: [email protected] Abstract Six new species and three new genera of cocalodine jumping spiders are described. Restricted to New Guinea and nearby areas, the Cocalodinae are basal salticids, outside the major salticid clade Salticoida. The new genera are Yamangalea (type species Y. frewana, new species), Tabuina (type species T. varirata, new species) and Cucudeta (type species C. zabkai, new species). In addition to these type species, described are the new species Tabuina rufa, Tabuina baiteta, Cucudeta uzet, Cucudeta gahavisuka, and Allococalodes madidus. The first description of females of the genus Allococalodes is provided. Natural history observations and photographs of living specimens are provided for all five genera of cocalodines. Key words: Araneae, Salticidae, Cocalodinae, jumping spider Introduction Among the salticid spiders endemic to New Guinea and nearby islands are the unusual genera Cocalodes Pocock and Allococalodes Wanless, notable for having a median apophysis on the male palp, widespread in spiders (Coddington 1990) but rare in salticids. Having a median apophysis indicates these genera are outside both of the two major clades of salticids, the Salticoida (Maddison & Hedin 2003) and the Spartaeinae (Wanless 1984). This puts these two genera among the sparse basal lineages of the family, and raises the possibility that they are an isolated, relictual group with only two body forms (Cocalodes, an elongate foliage dweller, and Allococalodes, more robust but with only two specimens previously described). However, in a recent expedition to Papua New Guinea, several new lineages of apparently-related salticids were found, revealing that Cocalodes and Allococalodes are part of a radiation of basal salticids in Australasia much more diverse than previously recognized. Described here are three new genera, Yamangalea (one species), Tabuina (three species), and Cucudeta (three species). In addition, a new species of Allococalodes is described, including the first known females of the genus. Material and methods Figure 1 shows the localities at which the specimens here described were sampled. All specimens except those of Tabuina baiteta are deposited in the Spencer Entomological Museum of the University of British Columbia (UBC-SEM). Tabuina baiteta is deposited in the Royal Belgian Institute of Natural Sciences (RBINS). Photographs of living specimens were taken with a Pentax Optio 33WR digital camera with a small lens glued to it for macro capability. Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Accepted by C. Muster: 19 Jan. 2009; published: 27 Feb. 2009 1 Terminology is standard for Araneae. All measurements are given in millimeters. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the "small eyes"). FIGURE 1. Papua New Guinea, with collecting localities for cocalodines described here. All localities except the Baiteta Forest were sampled during the 2008 Conservation International expedition. The Baiteta Forest site was sampled by Olivier Missa in 1995 by canopy fogging. Listed are all known cocalodines at each site. Taxonomy Subfamily Cocalodinae Simon A median apophysis on the male palpus is a striking rarity in salticid spiders. When present, it is a sclerite just clockwise from the base of the embolus (in a left palp, ventral view), surrounded by hematodocha, and at least partially surrounded by the tegulum (Wanless 1982, 1985; Maddison 2006; Figs 26, 57, 73). The sperm duct typically approaches it before turning counter-clockwise into the embolus. Wanless (1982) assumed the median apophysis was derived within salticids and thus was evidence that Cocalodes, Allococalodes, and the African Holcolaetis Simon and Sonoita Peckham and Peckham form a monophyletic group. However, the median apophysis is probably ancestral for salticids, based on its widespread presence throughout araneomorph spiders (Coddington 1990), in particular among families related to salticids (e.g. Bosselaers & Jocqué 2002; Ramírez 2003; Silva 2003; Benjamin et al. 2008), and based on its distribution among basal salticids in recent molecular phylogenies (Maddison & Needham 2006, Maddison et al. 2007). If so, then the presence of a median apophysis does not indicate to what clade a salticid belongs, but rather to what clades it 2 · Zootaxa 2021 © 2009 Magnolia Press MADDISON doesn't belong — a salticid with a median apophysis is a member of neither the Salticoida nor the Spartaeinae, both of which have lost it. [The sclerite reported as a "median apophysis" from the salticoid Tarne Simon by Szûts and Rollard (2007) is almost certainly not homologous to that in basal salticids: Tarne's sclerite is counterclockwise from the base of the embolus in the left palp ventral view, and is not cradled by the tegulum.] Wanless (1985) later preferred the hypothesis that Holcolaetis and Sonoita formed a clade with the spartaeines to the exclusion of Cocalodes. The placement of Cocalodes is therefore unresolved. Wunderlich (2004) synonymized the Spartaeinae with the Cocalodinae, choosing Simon's family-group name Cocalodeae because of priority over Wanless's name Spartaeinae. I reject this synonymy primarily because Cocalodes can be excluded from the Spartaeinae in Wanless's strict sense (1984) because it lacks two spartaeine synapomorphies, a tegular furrow and loss of the median apophysis. Even if these cocalodines and the spartaeines were found to be sister groups, we could still retain them as separate, as I do here. Wunderlich's broad sense of Cocalodinae, which includes the Baltic Amber salticids, is united as far as we know only by plesiomorphic character states (presence of conductor and median apophysis, many retromarginal cheliceral teeth, large posterior median eyes). Here I take a much more restricted concept of the Cocalodinae, to include only the five genera discussed below. The Baltic Amber salticids, other than the hisponines, are therefore Salticidae incertae sedis. I will not here attempt to resolve the placement of cocalodines in the phylogeny of the Salticidae, as I have found no morphological synapomorphies that link it with particular subgroups of salticids. Some of the cocalodines I describe here resemble spartaeines closely in habitus, but this observation is insufficiently precise to provide evidence for relationship. However, the three new genera described here are provisionally placed within the Cocalodinae along with Cocalodes and Allococalodes. This group has one proposed synapomorphy, the internal sclerotized spheres of the epigynum first reported by Wanless in Cocalodes. These are, however, also present in at least a few salticoids (see e.g., Galiano 1963, 1970: Tullgrenella morenensis (Tullgren) and Chira gounellei (Simon)). These spheres are visible clearly in the new species Yamangalea frewana, Tabuina varirata, Tabuina baiteta, Tabuina rufa, and Cucudeta zabkai (Figs 30, 43, 50, 61, 78; see arrow in Fig. 43), but were not seen in C. uzet and C. gahavisuka (Figs 82, 86) and are ambiguous in Allococalodes madidus (Fig. 19). Additional evidence for the monophyly of the cocalodines comes from preliminary data from the 28S gene, by which Cocalodes, Allococalodes and the three genera described here are resolved as a monophyletic group (Maddison & Zhang unpublished). Monophyly and limits of genera Justification of the new genera described here is not straightforward. I am reluctant to erect many new genera given the surplus that the Salticidae already has, but some new genera are needed, because body forms and genitalia of cocalodines are at least as diverse as the entire subfamily Spartaeinae. On the other hand, potential synapomorphies among these disparate forms are not obvious. Proposed synapomorphies are as follows: Cocalodes has a unique form of the male palp conductor, and has a more elongate body than any other cocalodine. Allococalodes species have no known synapomorphies, but are united with Cocalodes by the elongate chelicerae and the intercheliceral horn of males. Cucudeta species share the distinctive spination of the first metatarsus, a medial displacement of the posterior median eyes, the copulatory duct entering anteriorly into the spermathecae, and the leaf litter habitat. Tabuina varirata and T. baiteta share median apophyses with a similar small terminal hook, and sclerotized conductors (shared with Cocalodes, but possibly independently derived given the relationship of Cocalodes with Allococalodes). These characters leave the placement of Yamangalea frewana and Tabuina rufa unresolved. Underlying the choice to place them as I have is the preliminary unpublished molecular data cited above. It suggests three clades: (1) Cucudeta, (2) Tabuina varirata