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A Phylogenetic Analysis of the Subfamilies of Anyphaenidae (Arachnida, Araneae) MARTIN J

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A Phylogenetic Analysis of the Subfamilies of Anyphaenidae (Arachnida, Araneae) MARTIN J A phylogenetic analysis of the subfamilies of Anyphaenidae (Arachnida, Araneae) MARTIN J. RAMIREZ Ramirez, M. J.: A phylogenetic analysis of the subfamilies of Anyphaenidae (Arachnida, Ent. scand . Araneae). Ent. scand. 26: 361-384. Copenhagen, Denmark. December 1995. ISSN 0013- 8711. Anyphaenidae are diagnosed by the pattern of tracheal system, the forwardly advanced tra- cheal spiracle, and the lamelliform claw tuft setae. Cladistic analyses of the relationships of the anyphaenid genera are made from a selection of 9 exemplar taxa scored for 20 characters. In result, three subfamilies are proposed. Malenellinae subfam. n. is erected for Malenella nana gen. & sp. n. from southern Chile. This subfamily is considered as the sister group of all other anyphaenids. Anyphaeninae and Amaurobioidinae fonn a monophyletic group unit- ed by the orientation of the claw tuft setae and the grooved cymbial tip. Anyphaeninae are diagnosed by three characters associated with the advancement of the spiracle toward the epi- gastric furrow. Amaurobioidinae are diagnosed by the ingression of the median hematodocha in the male tegulum, and a characteristic secondary conductor. The relationships among the genera of Amaurobioidinae are discussed. The complex tracheal system of anyphaenids develops, during ontogeny, from a simple four-branched system. The genera Aysenia (from Clubionidae), Philisca (from Miturgidae) and Sanogasta (from Corinnidae) are transferred to Anyphaenidae: Amaurobioidinae. M. J. Ramirez, Department of Biology, FCEyN, Pabellon II Ciudad Universitaria, 1428 Bue- nos Aires, Argentina. Current address: Museo Argentino de Ciencias Naturales, Av. Angel Gallardo 470, 1405 Buenos Aires, Argentina. Introduction needed for a revision of the amaurobioidines, and continued reference or correction of an unpub- This paper had its origin after I started the revi- lished work is inconvenient, I decided to continue sion of the anyphaenid genera bearing a ventral Kochalka's effort here. membranose area in the male tegular base. Plat- nick (1977), who first noted this character, sug- gested that this group (including the Gondwanan Materials and methods Amaurobioides O. P.-Cambridge, and most of the species assigned to the Neotropical genera Gay- Tracheae were examined after digestion of non- enna Nicolet, Oxysoma Nicolet and Tomopisthes cuticular tissues with KOH 10-20% solution at Simon), probably deserved subfamily status. 100°C in a double boiler. Photographs of the tra- Kochalka, in an unpublished thesis (1980), agreed cheal system were made in 75% ethanol. The with him and used for the group the name Amau- preparation of spinnerets for SEM study followed robioidinae, an available family-rank name after Coddington (1989), except that these were air Hickman's (1949) Amaurobioididae. The name dried. Dissections for muscle observations were was still used by Lehtinen (1967) in Miturgidae, made on alcohol-fixed specimens; tissues and where he provisionally placed Amaurobioides. organs were removed with fine needles, alternat- Kochalka (1980) presented a partial generic revi- ing with ca. 5 seconds of sonication. Postem- sion of the amaurobioidines, and proposed char- bryonic stages are numbered according to Galiano acters for two monophyletic groups of genera, one (1991). All measurements are given in millime- related to Amaurobioides, the other to Gayenna. ters. See 'Acknowledgments' for acronyms of As a phylogenetic framework of the family was institutions. @EntGmalagica scandinavica (Grp 9) 362 RamIrez,M. J. ENT. SCANO. VOL. 26:4 (1995) The abbreviations of morphological terms used Bertkau (1878) proposed the family Anyphaeni- in the text and figures are: dae because of the complex tracheal system of Anyphaena Sundevall. At that time, and through IstB = first bifurcation of median tracheae. the following decades, some large families were 3rdOV =third dorso-ventral abdominal muscle. 3rdPO =third posterior oblique abdominal muscle. recognized on what we today consider as sym- 3rdVL =third ventral longitudinal abdominal muscle. plesiomorphic criteria. These dump-families, as 4thAO =fourth anterior oblique abdominal muscle. some authors have termed them, were found on 4th VL = fourth ventral longitudinal abdominal muscle. closer examination to be polyphyletic groups Ac = aciniform gland spigot. ALE = anterior lateral eye. whose components could not be placed in well ALS = anterior lateral spinneret. defined families. Good examples of dump-fami- AME =anterior median eye. lies are the amaurobiids, agelenids and clubion- ASC =amaurobioidine secondary conductor. C =conductor. ids. The latter family contained most of the two- CO = copulatory duct. clawed spiders with unmodified eyes and spinne- Col =colulus. rets. Considering that Simon (1897) included in CO =copulatory opening. Clubionidae groups as disparate as ctenids and OP = "dictynoid" pore. E = embolus. heteropodids, it is understandable that the differ- EB = embolar base. ences in the tracheal system were overlooked, and FO = fertilization duct. that the Anyphaenidae were abandoned as a sep- OrC = groove of the conductor. arate family. OrASC = groove of the amaurobioidine secondary con- ductor. The placement of anyphaenids in a separate LTr = lateral tracheae. family seemed natural after the dissociation of MA = median apophysis. major groups of clubionids into several families. mAmp = minor ampullate gland spigot. The most comprehensive works on anyphaenids MAmp =major ampullate gland spigot. MH = median hematodocha. (Platnick 1974; Platnick & Lau 1975) devoted MTr =median tracheae. much space to justify the distinction of anyphae- Nu = nubbin of major ampullate gland spigot. nids from clubionids. These authors also provided Pi =piriform gland spigot. synapomorphies in support of the monophyly of PLE =posterior lateral eye. PLS =posterior lateral spinneret. anyphaenids, primarily from the tracheal system PlSp = pulmonar spiracle. and the claw-tuft hairs. APmA =amaurobioidine paramedian apophysis. PME =posterior median eye. Inteifamilial relationships. - Outgroup relation- PMS = posterior median spinneret. RTA =retrolateral tibial apophysis. ships of the anyphaenids are far from understood. Sp = spermatheca. Basically, two alternative hypotheses have been St = subtegulum. proposed, each supported by different and con- T =tegulum. flicting character systems. These assign anyphae- TrSp = tracheal spiracle. nids to a dictynoid and the dionychan lineage, respectively. Forster (1970) included the anyphaenids in his Background Dictynoidea, defined by branched median tra- The family Anyphaenidae has traditionally cheae. He did not expalin, however, why other included two-clawed spiders with the tracheal two-clawed spider families with branched median spiracle conspicuously separated from the spinne- tracheae, such as salticids, thomisids or philodro- rets. As nearly all spiders known to possess these mids, were not included in the Dictynoidea. characters were included in Anyphaenidae, the Kochalka (1980) accepted Forster's view, and naturalness of the group was not questioned. The proposed a sister group relationship between only exception involved the genus Amaurobioides Notomatachia Forster (Desinae) and Anyphaeni- for which Hickman (1949) proposed a separate dae, based on the median tracheae supplying family, later synonymized with Anyphaenidae tracheoles to the prosoma, the protruded chelice- (Platnick 1974). In contrast, the rank and superfa- rae, the elongate body and the tip of the male pal- milial placement of anyphaenids have remained pal cymbium being relatively short. All these matters of controversy. characters occur in some anyphaenids, but the After studying the tracheal system of spiders, association with Notomatachia is untenable. Male ENT. SCAND. VOL. 26:4 (1995) Phylogeny of anyphaenid spiders 363 Figs 1-5. Tracheal system: (1) Malenella nana sr. n.; (2) Teudis cordobensis Mello-Leitao; (3) Tomopisthes varius Simon; (4) Tomopisthes varius, male (same scale as Fig. 3); (5) Arachosia sp., female. IstB = first bifurcation of median tracheae; LTr =lateral tracheae; MTr = median tracheae; PlSp = pulmonar spiracle; TrSp = tracheal spiracle. and female genitalia are almost identical in nids, but autapomorphies of some genera (e.g., Notomatachia and other desine genera such as Osoriella Mello-Leitao and Acanthoceto Mello- Matachia Dalmas, Goyenia Forster and PanDa Leitao). Forster (all figured by Forster 1970), and the char- Bennet (1992) proposed the secondary pores of acters invoked by Kochalka do not appear to be the spermathecae as a further synapomorphy for part of the groundplan of all these genera, but dictynoids, but they were not found in dictynines. rather autapomorphies of Notomatachia. Further- The discovery of this kind of pores in anyphae- more, the protruded chelicerae and elongate body nids (Fig. 34; Ramirez, in prep.) might reinforce form are not groundplan characters of anyphae- the hypothesis of a dictynoid lineage. Neverthe- 364 Ramirez, M. 1. ENT. SCAND. VOL. 26:4 (1995) Figs 6-12. Male and female tracheal systems: (6) Aysha prospera Keyserling, female; (7) male; (8) Teudis procerus Keyserling, female; (9) male; (10) Tasata parcepunctata Simon, female; (11) male; (12) Tomopistes immanis Simon, female, walls of two adjacent tracheal tubules near the division at pedicel; arrow indicates spiral thread. 1stB = first bifurcation of median tracheae; LTr = lateral tracheae. less, the distribution of this striking character is Notwithstanding, some dictynoid families bear
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