Arachnida: Opiliones: Laniatores)

New familial assignment for two harvestmen species of the infraorder Grassatores (Arachnida: Opiliones: Laniatores)

ABEL PÉREZ-GONZÁLEZ Grupo de Sistemática e Biologia Evolutiva (GSE), Núcleo em Ecologia e Desenvolvimento Sócio-Ambiental de Macaé (NUPEM), Universidade Federal do Rio de Janeiro (UFRJ), CP 119331, CEP 27910-970, Macaé, RJ, Brazil. E-mail: [email protected]

Incorporating masculine genitalic characters into Opiliones taxonomy has produced important revisions in the systematics of this group of arachnids. Currently, the inclusion of penis morphology in the description of any taxon of Phalangida (harvestmen with penis: Eupnoi + Dyspnoi + Laniatores, as used in Pinto-da-Rocha et al. 2007) has become an almost “mandatory” standard (e.g. Acosta et al. 2007), and opilionologists have been working to establish the masculine genital pattern for each family (e.g., Martens 1986; subchapters in Pinto-da-Rocha & Giribet 2007). Still, in the infraorder Grassatores the diversity in penis morphology is enormous and much structure and functionality remains poorly understood. Unfortunately, for many of the described Grassatores, the genitalia are entirely unknown, and this constitutes an important impediment to reliable familial assignation (e.g., in Kury 2003, 41 genera were considered as incertae sedis). This problem is quite relevant to “phalangodid-like” genera, considering their rather homogeneous external appearance but highly diverse genitalia (Martens 1988). One of the most illustrative examples is the subfamily Tricommatinae Roewer, 1912, that has been originally described under Phalangodidae, but which has a male genitalia groundplan matching the Gonyleptoidea, a very distant superfamily (Giribet et al. 2010). Consequently it was raised to a separate family, closer to the Gonyleptidae than to the Phalangodidae (Kury 1992), and finally regarded as a member of Gonyleptidae (Kury 2003). While revising the members of Samoidae and the Brazilian Grassatores incertae sedis in the Naturmuseum Senckenberg, Sektion Arachnologie (SMF) in Frankfurt am Main, Germany, I had the possibility to study two harvestmen species whose male genitalia were previously unknown and whose systematic placement remained obscure even after the comprehensive systematic overview offered in Pinto-da-Rocha & Giribet (2007). The male genitalia of both species exhibit characters to support their new familial assignments. The penial morphology nomenclature follows the recent taxonomical works: In Zalmoxidae the stragulum refers to a glans rigid sclerite articulated to the truncus like a jackknife (Kury & Pérez-González 2007; Macias-Ordoñez et al. 2010). The truncus lamina ventralis is divided into two tagmata: the distal rutrum which is hammer or spade shaped, usually bearing two pairs of paramedian setae, and the basal pergula which is a girdle bearing two to four pairs of erect setae (Kury & Pérez-González 2007). In Podoctidae the definition of follis follow Macias-Ordoñez et al. (2010), synonymies: “lamellar sac” in Martens (1986) and Kury & Machado (2009); “inflatable sac” in Kury (2007); for further details of Podoctidae male genitalia see the same bibliography. The method of male genitalia preparation and illustration follows Acosta et al. (2007).

Taxonomic account

Zalmoxidae Sørensen, 1886

Pirassunungoleptes H.E.M.Soares, 1966

Pirassunungoleptes analis (Roewer, 1949) new combination, new familial assignment (Figs 1 a–e)

Phalangodinus analis Roewer, 1949: 14, pl. 2, figs 9 a–f [Phalangodidae: Phalangodinae]; Kury 2003: 27. [Grassatores incertae sedis]

24 Accepted by A. Schönhofer: 4 Dec. 2010; published: 4 Feb. 2011 FIGURE 1. Phalangodinus analis Roewer, 1949, male holotype; (a) dorsal habitus; (b) leg IV, prolateral view; (c–e) distal part of penis: (c) dorsal view, (d) lateral view, (e) ventral view. Abbreviations: A1–4, Mesotergal areas 1 to 4; Fe, femur; Mt, metatarsus; Pa, patella; Pe, pergula; Ru, rutrum; S1–4, sulci 1 to 4; Ti, tibia. Stragulum in dark grey; capsula interna in medium lighter grey. Scale bars: a, 1mm; b, 0.5mm; c–e, 0.1 mm.

NEW FAMILIAL ASSIGNMENT FOR TWO GRASSATORES Zootaxa 2757 © 2011 Magnolia Press · 25 Types: SMF 9907447 (ex RII 7447/193), male holotype and two female paratypes (examined). Type locality: Brazil: Pernambuco State, Goiana (“Goyanna” in Roewer 1949). Remarks: Originally described as Phalangodidae: Phalangodinae by Roewer (1949) and later considered as Grassatores incertae sedis by Kury (2003). Justification of the new familial assignment: The male genitalia (Figs 1 c–e) show clearly the ventral plate of the pars distalis divided into pergula and rutrum, which is a characteristic of Zalmoxidae and Fissiphalliidae (Tourinho & Pérez-González 2006; Pinto-da-Rocha 2007) and the stragulum is quite different from the enlarged fingerlike stragulum of Fissiphalliidae (Martens 1986, 1988; Pinto-da-Rocha 2004, 2007; Tourinho & Pérez-González 2006). The short rutrum and the wide and short stragulum with a deep median cleft (dividing the stragulum in two branches) diagnose the species as a member of Zamoxidae. Some features of the external morphology of this species are also typical of Zalmoxidae (Kury & Pérez-González 2007): the pyriform body, the well marked bulla, the sulcus II being slightly “V” shaped and the enlarged tibia IV armored with spines (Figs 1 a–b and also see figs 9 a–f in Roewer 1949). Justification of the new combination: Phalangodinus surinamensis Roewer, 1912 (the type species of Phalangodinus Roewer, 1912) exhibits remarkabe morphological differences with Phalangodinus analis Roewer, 1949. These include a very wide and larger ocularium, enlarged chelicerae, parallel mesotergal sulci, and the anal operculum unarmed (Roewer 1949, personal observation). These differences point to both species belonging to different genera. I was not able to decide if Phalangodinus also belongs to Zalmoxidae because the male genitalia of P. surinamensis is still unknown. I therefore retained this taxon as Grassatores incertae sedis. A reliable allocation of P. analis into one of the present zalmoxid genera is not possible ahead of a taxonomical revision of this family whose taxonomy is affected by an enormous number of genera erected under a highly typological approach. Phalangodinus analis is therefore tentatively placed in the genus Pirassunungoleptes H.E.M. Soares, 1966 (type species: Pirassunungoleptes calcaratus H.E.M. Soares, 1966, from São Paulo State, Brazil), to which it shows the closest affiliation in terms of the following characters: minute size (less than 2 mm); pyriform habitus, ocularium small, near the frontal margin of the carapace (Fig. 1 a); anal operculum with a row of pointed aphophyses, and femur and tibia IV armed with a remarkable ventral acute apophyses (Fig. 1 b). This placement has to be verified within a more comprehensive revision of this and related genera.

Podoctidae Roewer, 1912

Waigeucola Roewer, 1949, new familial assignment

Waigeucola palpalis Roewer, 1949 (Figs 2 a–e)

Waigeucola palpalis Roewer, 1949: 59, figs 115 a–h. [Phalangodidae: Samoinae]

Types: SMF 9908683 (ex RII/8683/6), one male holotype and one male paratype (examined). Type locality: Indonesia: Waigeo Island (“Insel Waigeu” in the original label). Justification of the new familial assignment: Waigeucola Roewer, 1949, is a monotypic genus originally described in Phalangodidae: Samoinae (Roewer 1949). The type species is Waigeucola palpalis by original designation. This species is hereby considered as a member of Podoctidae because it possesses the male genitalia typical of the family (Figs 1 c–e) with the ventral plate deeply cleft and a well developed follis. Also, the tubercular bridge in the carapace (also called “ocular bridge” in Kury & Machado 2009) exhibited by this species (Fig. 1 a–b) is considered a possible synapomorphy of Podoctidae (Kury 2007). No evidence was found for placing this species in another existing podoctid genus, therefore it is transferred to this family with its original valid genus.

Acknowledgments

Thanks are given to Peter Jäger and Julia Altmann (SMF) for their kind attention during my visit to the Naturmuseum Senckenberg Sektion Arachnologie (SMF) in Frankfurt a. Main; this trip was supported by an Ernst Mayr Travel Grant in Animal Systematics (awarded by the Museum of Comparative Zoology at Harvard University). Peter Jäger kindly sent pictures of the types, to allow improvement of the plates. Cahyo Rahmadi kindly help with the Indonesian locality. Thanks to Adriano B. Kury for providing bibliography and helpful discussions about Opiliones systematics. I am

26 · Zootaxa 2757 © 2011 Magnolia Press PÉREZ-GONZÁLEZ indebted to Jerry Cates and Casey Richart for their kind revision of English language of the manuscript. The manuscript was greatly improved thanks to the critical review of Luis E. Acosta, Adriano Kury and Axel Schönhofer.

FIGURE 2. Waigeucola palpalis Roewer, 1949, male holotype; (a) detail of lateral carapace; (b) lateral habitus; (c–e) distal part of penis: (c) dorsal view, (d) lateral view, (e) ventral view. Abbreviations: Fo, follis; TB, tubercular bridge; VP, ventral plate. Scale bars: c–e, 0.1 mm.

References

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