Maymena, GERTSCH, 1960

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 : 778

MAYMENA GERTSCH, 1960

( FIGS 10–16 , 128B, D, E , 131I, J , 140M–O , 141A–I , 147D,E : CLADE C168)

Maymena Gertsch, 1960a: 30–38 . Gertsch, 1971: 94–95. Brignoli, 1974: 224. Baert, 1990: 17. Platnick, in Eberhard, Platnick & Schuh, 1993: 11. Griswold et al., 1998: 63.

Type species

Maymena mayana (Chamberlin & Ivie, 1938) by original designation, type material in MCZ, not examined.

Familial placement and composition

Transferred to Mysmenidae from Symphytognathidae by Forster & Platnick (1977). Maymena is closely related to Mysmenopsinae in our working phylogenetic hypothesis ( Fig. 161B ). Currently, Maymena comprises 13 described species (Platnick, 2014), and it is represented here by three described plus three undescribed species (the latter species scored only for molecular data): M. mayana , M. ambita , M. rica , Maymena -MYSM-003- ARG, Maymena -MYSM-004- MEX, and Maymena - MYSM-016-ARG).

Monophyly

Morphological synapomorphies of Maymena include: posterior lateral spinnerets with an anterior flat spatulate modified seta ( Fig. 11H ); aciniform gland spigots of the posterior spinnerets of two different shapes ( Fig. 13D ); seta on major ampullate field with one row of long ‘branches’ ( Figs 11E , 13C , 16B ); leg spination (i.e. macrosetae occurring on tibiae, femora, and metatarsi; Figs 140M , 141C ); and males with cylindrical palpal tibia ( Fig. 10A ). Ambiguously optimized synapomorphies for Maymena include: males with a femoral spot at least on femur I; metatarsal clasping spine particularly proximal ( Fig. 16G ); embolus rim deeply grooved ( Fig. 10H ); the primary cymbial conductor apically bent over the ventral side ( Fig. 10D, G ); epiandrous fusules dispersed in a row ( Figs 12B , 16A ); and macrosetae on female palpal tarsus ( Figs 13A , 15A ). Maymena is one of the few clades in this data set that is strongly supported and stable, and its monophyly is supported by most data partitions and parameter combinations. The genus is also supported by 125 molecular synapomorphies.

Diagnosis

Maymena shares with Mysmeninae the anterior flat spatulate modified seta on PLS and the aciniform gland spigots of two different shapes; with Trogloneta the presence of femoral spot on males; and with Mysmenopsinae the dispersed male epiandrous fusules, the macrosetae on female palpal tarsus, the leg spination, and the absence of tegular conductor (see also Gertsch, 1960a). Maymena differs from all other mysmenid genera by the combination of the aforementioned features, and by the presence of one row of long ‘branches’ on the major ampullate field seta, males with cylindrical palpal tibia, metatarsal clasping spine particularly proximal, seemingly interacting with the particularly distal tibial clasping spine in most species, and the male palpal morphology, with primary cymbial conductor apically bent over the ventral side. In addition, the respiratory arrangement distinguishes Maymena from other mysmenids. It consists of anterior booklungs and two long lateral tracheal tubes and two shorter median apodemes arising from a narrow posterior single spiracle (L. Lopardo, P. Michalik & G. Hormiga, unpubl. data; also Gertsch, 1960a).

Figures

Figure 10. Maymena ambita (Mysmenidae), male left palp: A, dorsal view; B, prolateral–dorsal view; C, prolateral view; D, prolateral–ventral view; E, ventral view; F, retrolateral–ventral view; H, retrolateral view, detail of tip of embolus and housing cymbial conductors; G, same, retrolateral–dorsal view; I, same, retrolateral–dorsal view; J, detail of squared area from I; K, detail of tibia from C. See Appendix 3 for the list of abbreviations.

Figure 11. Maymena ambita (Mysmenidae), female: A, abdomen, ventral view; B, same, detail of epigynum; C, same, posterior view; D, digested abdomen, vulva; E, left anterior lateral spinneret; F, posterior median spinnerets; G, H, posterior lateral spinnerets (PLS); G, right PLS; H, left PLS. See Appendix 3 for the list of abbreviations.

Figure 12. Maymena mayana (Mysmenidae): A, B, male; C–H, female. A, left palp, dorsal–retrolateral view; B, abdomen, detail of epiandrous spigots; C, digested abdomen, detail of spermathecae; D, epigynum, ventral view; E, same, detail of copulatory openings; F, G, right leg I; F, claws; G, tarsus–metatarsus junction; H, left leg IV, claws. See Appendix 3 for the list of abbreviations.

Figure 13. Maymena mayana (Mysmenidae): A–C, E, F, H–J, female; D, G, male. A, prosoma, lateral view; B, same, dorsal view; C, left anterior lateral spinnerets; D, posterior median spinnerets; E–G, right posterior lateral spinnerets; E, detail of modified spatulate seta and aggregate spigots; H, detail of female palpal tibia; I, mouthparts, distal right chelicera; J, same, detail of promarginal teeth. See Appendix 3 for the list of abbreviations.

Figure 14. Maymena rica (Mysmenidae): A, B, D, male left palp; C, E, female. A, dorsal view; B, same, prolateral view, detail of tip of embolus and housing cymbial conductors; C, epigynum, ventral view; D, same as B, dorsal view; E, digested abdomen, detail of vulva. See Appendix 3 for the list of abbreviations.

Figure 15. Maymena rica (Mysmenidae), prosoma: A, C, F, G, female; B, D, E, H, I, male. A, B, lateral view; C, dorsal view; D, carapace, frontal view; E, prosoma, posterior view; F, ocular area, detail from panel C; G–I, mouthparts, detail of cheliceral fang and teeth; G, distal promargin of left chelicera; H, distal promargin of right chelicera; I, distal right chelicera.

Figure 16. Maymena rica (Mysmenidae), abdomen and legs: A, B, G, H, male; C–F, female. A, epiandrous spigots; B, left anterior lateral spinnerets; C, abdomen, ventral view. D, G, H, left leg I; E, F, left leg IV. D, femur, ventral view; E, tibia, dorsal view; F, claws, retrolateral view; G, tibia–metatarsus junction, prolaterodorsal view; H, tarsus, prolaterodorsal view. See Appendix 3 for the list of abbreviations.

Figure 128. Mysmenidae female genitalia, cleared: A, Isela okuncana, dorsal view; B, Maymena mayana, dorsal view; C, Kilifina-MYSM-002-KENYA (Isela sp.), from Kwale, Kenya, dorsal view; D, Maymena ambita, ventral view; E, Maymena rica, dorsal view; F, Trogloneta granulum, ventral view; G, Mysmenopsis dipluramigo, dorsal view. See Appendix 3 for the list of abbreviations.

Figure 131. Mysmenidae male left palp (unless otherwise stated), cleared: A, Mysmenopsis penai, retrolateral view, right palp, inverted; B, Mysmenopsis dipluramigo, retrolateral view; C, Mysmenopsis cidrelicola, presumably paralectotype, retrolateral view, right palp, inverted; D, Kilifina-MYSM-002-KENYA (Isela sp.), from Kwale, Kenya, ventral view, right palp, inverted; E, Trogloneta granulum, ventral view; F, Trogloneta cantareira, ventral view; G, Kilifina-MYSM-002- KENYA (Isela sp.), from Kwale, Kenya, dorsal view, right palp, inverted; H, Isela okuncana, ventral view; I, Maymena ambita, ventral view; J, Maymena mayana, ventral view. See Appendix 3 for the list of abbreviations.

Figure 140. Composite images of Mysmenidae species: Mysmenopsinae and Maymena. A–C, Isela okuncana; A, female, lateral view; B, female, ventral view; C, male, ventral view. D–F, Kilifina-MYSM-002-KENYA (Isela sp.), from Kwale, Kenya; D, female, lateral view; E, male, lateral view; F, male, ventral view. G, H, Mysmenopsis dipluramigo; G, female, lateral view; H, male, lateral view. I, Mysmenopsis palpalis, female, ventral view. J, K, Mysmenopsis cidrelicola, male paralectotype; J, lateral view; K, ventral view. L, Mysmenopsis penai, female, dorsal view. M–O, Maymena ambita; M, male, lateral view; N, female, lateral view; O, female, dorsal view. Scale bars: A–I, L–O, 0.5 mm; J, K, 1 mm.

Figure 141. Composite images of Mysmenidae species: Maymena and Mysmeninae. A, B, Maymena rica, female allotype; A, lateral view; B, dorsal view. C–F, Maymena mayana; C, female, frontal view; D, male, lateral view; E, male, dorsal view; F, male, ventral view. G–I, Maymena species, female, abdomen ventral; G, Maymena ambita; H, Maymena mayana; I, Maymena rica, female allotype. J–L, Microdipoena (= Anjouanella) comorensis; J, female paratype, lateral view; K, male holotype, lateral view; L, male holotype, ventral view. M, N, Microdipoena elsae; M, male holotype, lateral view; N, female allotype, lateral view. O, Microdipoena (= Mysmenella) illectrix, male, ventral view. Scale bars: A, B, G, I–O, 0.5 mm; C–F, H, 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 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.