Mysmena, SIMON, 1894

Lopardo, Lara & Hormiga, Gustavo, 2015, Out of the twilight zone: phylogeny and evolutionary morphology of the orb-weaving spider family Mysmenidae, with a focus on spinneret spigot morphology in symphytognathoids (Araneae, Araneoidea), Zoological Journal of the Linnean Society 173 (3), pp. 527-786 : 784-785

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) .

Figures

Figure 28. MYSM-005-ARG (Mysmena, Mysmenidae) from Misiones, Argentina; male. A, prosoma, lateral view; B–G, left palp. B, dorsal view; C, prolateral view; D, prolateral–ventral view; E, retrolateral–ventral view; F, detail of embolus tip, ventral view; G, same, retrolateral view. See Appendix 3 for the list of abbreviations.

Figure 29. MYSM-005-ARG (Mysmena, Mysmenidae) from Misiones, Argentina. A–C, E, F, female; D, male. A, digested abdomen, detail of left spermatheca; B, same, detail of posterior tracheal system; C, epigynal area and scapus; D, left femur I, retrolateral view; E, left tibia IV, prolateral view; F, left metatarsus IV, prolateral view. See Appendix 3 for the list of abbreviations.

Figure 30. MYSM-007-MEX (Mysmena, Mysmenidae) from Chiapas, Mexico; male left palp. A, retroventral view; B, ventral view; C, prolateral view; D, dorsal view; E, retrolateral view; F, detail of tip of palp, ventral–prolateral view. See Appendix 3 for the list of abbreviations.

Figure 31. MYSM-007-MEX (Mysmena, Mysmenidae) from Chiapas, Mexico; A–D, male left palp; E–G, female. A, detail of proximal palp, prolateral basal expansion and paracymbium, retroventral view; B, detail of dorsal–retrolateral tip of cymbium; C, same, prolateral view; D, same, retrolateral view; E, cheliceral bases, frontal view; F, abdomen, lateral view; G, abdomen, lateral view, detail of scapus and spinnerets. See Appendix 3 for the list of abbreviations.

Figure 32. Mysmena (= Calodipoena) incredula (Mysmenidae); male palp. A–C, E–H, left palp; D, right palp, subtly expanded, inverted. A, D retrolateral view; B, ventral–prolateral view; C, detail of proximal palp, retrolateral view; E, detail of tibia and prolateral basal expansion, ventral view; F, same, close up; G, detail of tip of palp, ventral view; H, same, retrolateral view. See Appendix 3 for the list of abbreviations.

Figure 33. Mysmena (= Calodipoena) incredula (Mysmenidae); abdomen. A, B, female digested abdomen; C–H, male spinnerets. A, tracheal system and vulva; B, same, detail of vulva; C, spinning field, ventral–posterior view; D, posterior median spinnerets; E, right anterior lateral spinnerets (ALS); F, left ALS; G, right posterior lateral spinnerets; H, same, detail. See Appendix 3 for the list of abbreviations.

Figure 34. Mysmena (= Calodipoena) incredula (Mysmenidae); legs. A, B, female right leg I; C–F, male left legs. A, femoral spot, retrolateral view; B, metatarsal trichobothrium, dorsal view; C, metatarsus I, prolateral view; D, tarsus I, prolateral view; E, femur IV, prolateral view; F, claws IV, retrolateral view.

Figure 35. Mysmena (= Calodipoena) incredula (Mysmenidae): A, E, F, male; B–D, female; A, B, ocular area, frontal view; C, prosoma, ventral view; D, same, detail of anterior sternum and labium; E, abdomen, pedicel area and epigastric furrow; F, epiandrous spigots.

Figure 36. Mysmena-MYSM-015-MAD (Mysmena, Mysmenidae) from Antananarivo, Madagascar; male left palp; A, retrolateral view; B, distal view; C, prolateral view; D, ventral view; E, dorsal view. See Appendix 3 for the list of abbreviations.

Figure 37. Mysmena-MYSM-015-MAD (Mysmena, Mysmenidae) from Antananarivo, Madagascar; female abdomen; A, epigynal area and scapus, ventral view; B, left posterior spinnerets (PS); C, same as A, posterolateral view; D, digested abdomen, detail of vulva; E, right PS. See Appendix 3 for the list of abbreviations.

Figure 38. Mysmena-MYSM-015-MAD (Mysmena, Mysmenidae) from Antananarivo, Madagascar; prosoma: A, B, male; C–H, female; A–C, lateral view; D, carapace, frontal view; E, chelicerae and palps, frontal view; F, mouthparts, ventral view; G, right palp, detail of palpal tip; H, distal promargin of right chelicera.

Figure 39. Mysmena-MYSM-015-MAD (Mysmena, Mysmenidae) from Antananarivo, Madagascar; abdomen and left legs: A, C, male; B, D–J, female. A, abdomen, ventral view; B, booklung cover; C, leg I, tarsal organ, retrolateral view; D, femur I, retrolateral view; E, metatarsus I, retrolateral view; F, femur IV, retrolateral view; G, tibia IV, retrolateral view; H, tarsus I, retrolateral view; I, tarsus IV, retrolateral view; J, leg IV, metatarsus–tarsus junction, dorsal view.

Figure 40. Mysmena-MYSM-015-MAD (Mysmena, Mysmenidae) from Antananarivo, Madagascar; male left palp, details. A, ventral-distal view; B, prolateral view; C, same, close up; D, ventral view; E, retrolateral view; F, dorsal view. See Appendix 3 for the list of abbreviations.

Figure 41. Mysmena leichhardti (Mysmenidae) from Queensland, Australia; male left palp. A, prolateral–ventral view; B, dorsal view; C, retrolateral–ventral view; D, prolateral view; E, retrolateral view. See Appendix 3 for the list of abbreviations.

Figure 42. Mysmena leichhardti (Mysmenidae) from Queensland, Australia: A, B, E, G, H, male; C, D, F, female. A, prosoma, lateral view; B, same, lateral–frontal view; C, epigynal area and scapus, ventral view; D, digested abdomen, detail of vulva, tracheae removed; E, mouthparts, distal view; right femur I, prolateral–dorsal view; G, left femur I, ventral view; H, left metatarsus I, prolateral view. See Appendix 3 for the list of abbreviations.

Figure 43. Mysmena leichhardti (Mysmenidae) from Queensland, Australia; male left palp, details. A, B, embolus– conductor interaction; C, distal palp, distal–prolateral view; D, dorsal cymbial tip; E, ventral–prolateral cymbial tip. See Appendix 3 for the list of abbreviations.

Figure 44. Mysmena-MYSM-018-MAD (Mysmena, Mysmenidae) from Mahajanga, Madagascar: A, female epigynal area and scapus, ventral–posterior view; B–D, male left palp; B, prolateral view; C, detail of embolus and pars pendula, retrolateral– ventral view; D, cymbial distal tip. See Appendix 3 for the list of abbreviations.

Figure 49. MYSM-023-MAD (Mysmeninae, Mysmenidae) from Antananarivo, Madagascar: A, D, E, female; B, C, F–H, male. A, digested abdomen, detail of vulva; B, left palp, prolateral view, bulb displaced; C, same, detail of cymbial tip, retrolateral–dorsal view; D, abdomen, detail of epigynal area and respiratory spiracles, posterior view; E, same, ventral view; F, colulus and spinnerets, ventral view; G, posterior median spinnerets; H, prosoma, detail of sternum, labium and maxillae, ventral view. See Appendix 3 for the list of abbreviations.

Figure 51. Mysmena tasmaniae (Mysmenidae). A–C male left palp; A, dorsal view; B, retrolateral–dorsal view; C, expanded palp, distal–ventral view. D, female digested abdomen, detail of vulva, tracheae removed. See Appendix 3 for the list of abbreviations.

Figure 52. A, MYSM-034-MAD (Mysmena, Mysmenidae), female; B–D, MYSM-029-MAD (Mysmeninae, Mysmenidae), female; E, F, MYSM-028-MAD (Mysmena, Mysmenidae), female. A, prosoma, lateral view; B, right anterior lateral spinneret; C, right posterior lateral spinneret; D, right leg I, claws, retrolateral view; E, chelicerae and palps, frontal view; F, epigynal area, ventral view. See Appendix 3 for the list of abbreviations.

Figure 65. A, Mysmena (= Tamasesia) rotunda (Mysmenidae), female syntype, left posterior lateral spinneret. B–H, Trogloneta cantareira (Mysmenidae). B, female left leg I; C–F, male left leg I; G, H, female left leg IV. B, femur, patella and tibia; C, metatarsus, dorsal view; D, tarsus, prolateral view; E, proximal metatarsal cuticular pattern, dorsal view; F, claws, prolateral view; G, tarsus, retrolateral view; H, metatarsus, retrolateral view. See Appendix 3 for the list of abbreviations.

Figure 129. Mysmenidae female genitalia, cleared: A, Microdipoena guttata, ventral view; B, Microdipoena nyungwe, ventral view; C, MYSM-007-MEX (Mysmena), from Chiapas, Mexico, ventral view, arrow to diverticle of copulatory duct; D, Microdipoena elsae, ventral copulatory duct; E, Microdipoena (= Mysmenella) samoensis, syntype, ventral view; F, Microdipoena (= Mysmenella) jobi, paratype, ventral view; G, Mysmena-MYSM-015-MAD (Mysmena), from Antananarivo, Madagascar, dorsal view; H, MYSM-029-MAD (Mysmeninae), from Antsiranana, Madagascar, ventral view. See Appendix 3 for the list of abbreviations.

Figure 130. Mysmenidae female genitalia, cleared. A, Mysmena (= Calodipoena) tarautensis, paratype, dorsal view, arrow to broken scapus; B, Mysmena (= Calodipoena) incredula, dorsal view; C, Mysmena leichhardti, from Queensland, Australia, ventral view; D, MYSM-023-MAD (Mysmeninae), from Antananarivo, Madagascar, ventral view; E, Mysmena tasmaniae, ventral view; F, MYSM-005-ARG (Mysmena), from Misiones, Argentina, ventral view; G, MYSM-019-MAD (Mysmeninae), from Toliara, Madagascar, ventral view. See Appendix 3 for the list of abbreviations.

Figure 133. Mysmenidae male left palp (unless otherwise stated), cleared. A, MYSM-005-ARG (Mysmena), from Misiones, Argentina, dorsal view, right palp, inverted, arrow to cymbial groove (CyG); B, same, ventral view; C, MYSM-007-MEX (Mysmena), from Chiapas, Mexico, retrolateral view, arrows to CyG; D, Mysmena (= Kekenboschiella) awari, paratype, prolateral view; E, same, dorsal–retrolateral view; F, Mysmena (= Kekenboschiella) marijkeae, holotype, retrolateral view, right palp, inverted; G, Brasilionata arborense, holotype, retrolateral view, right palp, inverted; H, Mysmena (= Tamasesia) rotunda, type, prolateral view; I, same, retrolateral view, schematic drawing. See Appendix 3 for the list of abbreviations.

Figure 134. Mysmenidae male left palp (unless otherwise stated), cleared: A, Mysmena leucoplagiata, type, prolateral view, right palp, inverted; B, Mysmena leichhardti, from Queensland, Australia, prolateral–ventral view; C, Mysmena tasmaniae, prolateral–ventral view, right palp, inverted; D, Mysmeniola spinifera, holotype, prolateral view, right palp, inverted; E, MYSM-019-MAD (Mysmeninae), from Toliara, Madagascar, prolateral view; F, MYSM-023-MAD (Mysmeninae), from Antananarivo, Madagascar, retrolateral view, right palp, inverted; G, MYSM-020-MAD (Mysmeninae), from Toamasina, Madagascar, dorsal view, right palp, inverted; H, same, ventral view. See Appendix 3 for the list of abbreviations.

Figure 142. Composite images of Mysmenidae species: Mysmeninae and Trogloneta. A, B, Microdipoena nyungwe; A, male, lateral view; B, female, ventral view. C, Trogloneta granulum, female, ventral view. D, Mysmena (= Calodipoena) incredula, female, lateral view, inverted. E, Mysmena (= Calodipoena) mootae, female holotype, dorsal view. F, Mysmena (= Calomyspoena) santacruzi, female paratype, dorsal view. G–I, Mysmena (= Itapua) tembei; G, male holotype, lateral view, inverted; H, male holotype, frontal view; I, female paratype, frontal view. J–L, Mysmena (= Kekenboschiella) awari, male paratype; J, lateral view; K, dorsal view; L, ventral view. M, N, Mysmeniola spinifera, male holotype; M, lateral view; N, ventral view. O, Brasilionata arborense, male holotype, dorsal view, abdomen detached. Scale bars: 0.5 mm.

Figure 143. Composite images of Mysmeninae species. A–C, Mysmena tasmaniae; A, female, lateral view; B, female, ventral view; C, male, ventral view. D–F, MYSM-019-MAD (Mysmeninae); D, female, lateral view; E, female, ventral view; F, male, lateral view. G–I, MYSM-005-ARG (Mysmena), female; G, lateral view; H, dorsal view; I, ventral view. J–L, MYSM-007-MEX (Mysmena); J, female, lateral view; K, female, dorsal view; L, male, ventral view. M, Mysmena- MYSM-015-MAD (Mysmena), female, lateral view. N, Mysmena leichhardti, female, lateral view. O, MYSM-028-MAD (Mysmena), female prosoma, ventral view. Scale bars: 0.5 mm.

Figure 144. Composite images of Mysmenidae species and outgroups. A, MYSM-023-MAD (Mysmeninae, Mysmenidae), male, lateral view. B, Phricotelus stelliger (Araneoidea), female, type, lateral view. C, Iardinis mussardi (Symphytognathidae), male holotype, dorsal view. D–F, Leucauge venusta (Tetragnathidae); D, male, lateral view; E, female, lateral view; F, female, dorsal view. G–J, Linyphia triangularis (Linyphiidae); G, male, lateral view; H, male, ventral view; I, female, dorsal view; J, female, lateral view. K, Elanapis aisen (Anapidae), male, ventral view; L, Mysmena (= Tamasesia) rotunda, female, dorsal view. Scale bars: A–C, K, 0.5 mm; D–J, 1 mm.

Figure 147. Webs of Mysmenidae: A, Mysmena tasmaniae, B, MYSM-005-ARG (Mysmena), from Misiones, Argentina, female with egg sac; C, Mysmenidae from Chiapas, Mexico, detail of centre of web, external threads removed to expose the hub; D, Maymena sp. from Misiones, Argentina; E, same, detail of centre of web.

Figure 160. Phylogenetic hypothesis rendered by the complete morphological and molecular data set analysed under equal-weights parsimony using the dynamic homology criterion. Cladogram taken from Lopardo et al. (2011: fig. 12). Numbers below each node indicate node numbers. Note that the micropholcommatine Taphiassa punctata and the theridiosomatid Coddingtonia euryopoides were referred to as Parapua punctata and TSMD-002-THAI (respectively) in Lopardo et al. (2011). Family codes used for unidentified species are as follows: ANAP, Anapidae; MYSM, Mysmenidae; SYMP, Symphytognathidae; TSMD, Theridiosomatidae.

Figure 161. Hypothesis of relationships within Mysmenidae, taken from the results of the combined analysis from Lopardo et al. (2011) (see Fig. 160). New taxonomic (i.e. generic) status as proposed in our study is assigned to each representative; new combinations are emphasized in bold. Unambiguous morphological character optimizations are shown for every branch in the tree. Numbers below each node indicate node numbers. Empty and filled boxes represent homoplasious and non-homoplasious transformations, respectively. Major clades representing taxonomic decisions discussed in the text are highlighted in grey boxes.