MYSMENA SIMON, 1894

(FIGS 28–44, 51, 52, 65A, 129C, G, 130A–C, E, F, 133A–F, H, I, 134A–C, 142D–L, 143A–C, G–O, 144O, 147A, C: CLADE C144)

Mysmena Simon, 1894: 588 . Bishop & Crosby 1926: 177 (synonymized Microdipoena). Levi, 1956: 3 (synonymized with Calodipoena, Tamasesia, and Microdipoena). Forster, 1959: 303–307; 1977: 129. Gertsch, 1960b: 13. Kraus, 1967: 388. Loksa, 1973: 283. Saaristo, 1978: 125 (rejected synonymy with Microdipoena). Hickman, 1979: 74. Brignoli, 1980: 729 (rejected synonymy with Calodipoena, Tamasesia, and Microdipoena). Wunderlich, 1980b: 267; 1986: 218. Davies, 1985: 91 (transfer to Calodipoena by Brignoli, 1983 rejected). Snazell, 1986: 62. Trotta, 2005: 170. Ono, in Ono, Chang & Tso, 2007: 73. Lopardo & Dupérré, in Lopardo et al., 2008: 37.

Calodipoena Gertsch & Davis, 1936: 8 (type species by original designation C. incredula Gertsch & Davis, 1936, type material in AMNH, not examined). Brignoli, 1983: 376 (transfer from Mysmena because of alleged relationships with C. incredula Gertsch & Davis, 1936). New synonymy.

Itapua Baert, 1984 b: 604 (type species by original designation and monotypy I. tembei Baert 1984, type material in MHNG, examined). New synonymy.

Calomyspoena Baert & Maelfait, 1983: 104 (type species by original designation and monotypy C. santacruzi Baert & Maelfait, 1983, type material in IRSN, examined). New synonymy.

Tamasesia Marples, 1955: 476 (type species by original designation T. rotunda Marples, 1955, type material in BMNH and MNHN, examined); Levi 1956: 3 (transfer from Tamasesiidae to Theridiidae, synonymized with Mysmena); Brignoli, 1980: 730 (transfer to Mysmenidae, rejected synonymy with Mysmena). New synonymy.

Kekenboschiella Baert, 1982: 303 (type species by original designation K. marijkeae Baert, 1982, type material in IRSN, examined). Baert 1984a: 230. New synonymy.

Type species

Mysmena leucoplagiata (Simon, 1879) by original designation, type material in MNHN, examined (see below, and also Kraus, 1967).

Synonymy justification

Relationships among and within mysmenine clades are highly unstable and poorly supported (see Results). Morphologically, these results are not surprising. In particular, homoplasy is widespread among the Mysmena representatives, and no notable synapomorphy characterizes this genus. The inclusion in the analysis of several undescribed mysmenid species with distinct and diverse morphology (especially genitalic morphology) might obscure relationships, producing an even more unstable pattern. In this case, although distinct undescribed species might possibly represent new genera (see comments below), to date the available data do not support such hypotheses. As circumscribed here, Mysmena includes a polyphyletic Calodipoena, the monotypic genera Itapua and Calomyspoena, and a (strictly) monophyletic Tamasesia, Kekenboschiella, and Mysmena s.s., although the latter three genera are supported by just one (or none in the case of Mysmena) homoplastic character change, and no molecular transformations. In the absence of concise and unique diagnostic features for any of the aforementioned genera, we have re-circumscribed the genus Mysmena to avoid proliferation of monotypic genera and non-monophyletic taxa, losing phylogenetic information.

Familial placement, composition, and re-circumscription

Mysmena was transferred to Symphytognathidae from Theridiidae by Forster (1959), and to Mysmenidae from Symphytognathidae by Forster & Platnick (1977). In the proposed phylogenetic hypothesis, Mysmena comprises a large clade distally within the Mysmeninae lineage (Fig. 160; for synonymies and new combinations, refer to Fig. 161B). The re-circumscribed Mysmena comprises a total of 42 described species: 23 from Mysmena, ten from Calodipoena, four from Kekenboschiella, three from Tamasesia, and one from each of the two monotypic genera Calomyspoena and Itapua (Platnick, 2014). Mysmena is here represented by 11 described plus 18 undescribed species (12 of the latter undescribed were species scored only for molecular data): M. leucoplagiata, M. leichhardti, M. tasmaniae, M. mootae comb. nov. (from Calodipoena), M. incredula comb. nov., M. santacruzi comb. nov., M. tembei comb. nov., M. acuminata comb. nov. (from Tamasesia), M. rotunda comb. nov., M. awari comb. nov. (from Kekenboschiella), M. marijkeae comb. nov., Mysmena- MYSM-011-ARG, Mysmena- MYSM-014- THAI, Mysmena- MYSM-(015 018)-MAD, MYSM-(005 038–042)-ARG, MYSM-(007 010)- MEX, MYSM-(013 035–037)-THAI, and MYSM-028-MAD.

Monophyly and diagnosis

Ambiguously optimized synapomorphies for Mysmena, shared by most of the taxa, include the spermatic duct switchback SB I distally bending at a right angle [Figs 133D–F, 134A, B, E; straight in Mysmena- MYSM-015-MAD, Mysmena (= Tamasesia) rotunda and MYSM-005-ARG, Fig. 133A, B, H, I], and the presence of a long ventral scapus (Figs 29C, 31G, 37C, 42C, 129C, G, 130B) and weakly sclerotized fertilization ducts, with a distinguishable wall (Figs 42D, 49A, 51D, 129C, G, 130A; independently membranous, translucent in Itapua tembei, Mysmena leucoplagiata, and MYSM-034- MAD). Mysmena monophyly is also supported by 265 molecular synapomorphies.

Because of its previous placement within Theridiidae, and its recurrent synonymies with Calodipoena, Tamasesia, and/or Microdipoena, the previous diagnoses of Mysmena include features that are currently considered synapomorphic for the subfamily Mysmeninae or even Mysmenidae (see e.g. Simon, 1894; Levi, 1956; Forster, 1959; Gertsch, 1960a; Kraus, 1967). Also, because of the mislabelling of the vial containing the type specimen of Mysmena leucoplagiata, which also included specimens of Mysmenella jobi, the diagnosis of Mysmena has been rather confusing and inaccurate (vial examined; see also Kraus, 1967). For example, the type species Mysmena leucoplagiata has been correctly redescribed by Kraus (1967) and Wunderlich (1980b), whereas Mysmenella jobi was mistakenly redescribed as Mysmena leucoplagiata by Levi (1956) and Loksa (1973). Furthermore, diagnostic features of the here-synonymized genera are also largely broad for the family or at least Mysmeninae (Gertsch & Davis, 1936; Marples, 1955; Baert, 1982, 1984a), except for a few diagnostic features of Calomyspoena (Baert & Maelfait, 1983) .