Microdipoena, BANKS 1895

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 : 783-784

MICRODIPOENA BANKS 1895   Edit

( FIGS 17–27 , 129A, B, D–F , 132 , 141J–O , 142A,B : CLADE C125)

Mysmena Simon, 1895b: 149   Edit . Bishop & Crosby, 1926: 177. Levi, 1956: 8. Forster, 1959: 306. Kraus, 1967: 392. Gruia, 1977: 162. Shinkai, 1977: 326. Roberts, 1978: 932. Wunderlich, 1980b: 267; 1986: 222. Kasal, 1982: 75. Heimer & Nentwig, 1991: 306.

Microdipoena Banks, 1895: 85   Edit . Saaristo, 1978: 124–125 (rejected synonymy to Mysmena   Edit by Bishop & Crosby, 1926: 177). Brignoli, 1980: 731 (rejected synonymy to Mysmena   Edit by Bishop & Crosby, 1926: 177). Baert, 1984b: 608; 1985: 51; 1989: 29.

Anjouanella Baert, 1986: 265   Edit (type species by monotypy A. comorensis Baert, 1986   Edit , type material in MRAC, examined). New synonymy.

Mysmenella Brignoli, 1980: 731   Edit (transfer from Mysmena   Edit , type Mysmena illectrix Simon, 1895b   Edit , type material in MNHN, examined). Baert, 1984a: 240 (transfer from Mysmena   Edit ); 1989: 32. Namkung & Lee, 1987: 46. Coddington, 1990: 19. Thaler & Noflatscher, 1990: 174. Namkung, 2002: 146; 2003: 148. Wunderlich, 2004: 1073 (considered a junior synonym of Mysmena Simon, 1894   Edit ). Yin et al., 2004: 80. Lee et al., 2004: 100. Trotta, 2005: 170. Ono, 2007: 170. New synonymy.

Type species

Microdipoena guttata Banks, 1895   Edit by original designation, type material in MCZ, examined.

Familial placement, composition, and re-circumscription

Our working phylogenetic hypothesis places Microdipoena   Edit sister to Brasilionata   Edit within the mysmenine clade C128, which also comprises Mysmeniola   Edit and MYSM- 019- MAD. Microdipoena   Edit comprises four described species (Platnick, 2014) and under the current re-circumscription, 11 other described species are transferred here (a total of 15 described species). Microdipoena   Edit is here represented by seven described plus two undescribed species ( Fig. 161B ; the latter two species are scored only for molecular characters): M. guttata   Edit , M. elsae   Edit , M. nyungwe   Edit , M. samoensis   Edit comb. nov. (from Mysmenella   Edit ), M. jobi   Edit comb. nov. (from Mysmenella   Edit ), M. illectrix   Edit comb. nov., M. comorensis   Edit comb. nov., Microdipoena   Edit -AToL-DR, and MYSM- 030- MAD.

Monophyly, diagnosis, and synonymy justification The following combination of morphological synapomorphies is unique and therefore diagnostic for Microdipoena   Edit (and are shared among all Microdipoena   Edit representatives, unless noted): abdomen with a whitish ventral ring around the spinnerets ( Fig. 142A ; except Anjouanella   Edit , with all ventral abdominal area lighter, Fig. 141J–L ); males with two prolateral apical clasping spines on tibia I ( Figs 26C , 27I , 141K, L, O ), thick embolus with an apical switch in the coiling direction ( Figs 18C, D, F , 27C , 132B, D, E ; also in Brasilionata   Edit ), and with either a distal apophysis ( Fig. 18F ) or a distal irregular membrane ( Fig. 27A– C ; except in Anjouanella   Edit , without modifications), spermatic duct switchback SB I parallel, with the portions of the spermatic duct before and after the switch SB I run close with each other and with one pair of extra switches (SB III and IV, Fig. 132B–E ); and small paracymbium ( Figs 17C , 22G , 27B ). As most Microdipoena   Edit representatives in this data set were scored only for morphology, no molecular synapomorphies optimize at the node of this genus; however, its distal clade (clade C172), which includes the only sequenced species of this genus, is supported by 92 molecular synapomorphies. Previous diagnoses for Microdipoena   Edit s.s., Mysmenella   Edit , and Anjouanella   Edit are in agreement with the current diagnosis of the enlarged Microdipoena   Edit (see e.g. Banks, 1895; Brignoli, 1980; Baert, 1986).

BRASILIONATA WUNDERLICH, 1995   Edit

FIGS 133G , 142O

Brasilionata Wunderlich, 1995: 545   Edit .

Type species

Brasilionata arborense Wunderlich, 1995   Edit by original designation and monotypy, holotype in AMNH, examined.

Familial placement and composition

Brasilionata   Edit is a member of the mysmeninae   Edit clade C128 (also comprising Mysmeniola   Edit , Microdipoena   Edit , and MYSM- 019- MAD), and is sister to Microdipoena   Edit ( Fig. 161B ). Brasilionata   Edit is here represented by its type and only species B. arborense   Edit .

Monophyly and diagnosis

This Brazilian monotypic genus is only known by the male holotype specimen, and it is diagnosed by the following combination of autapomorphies: male palpal tibial rim scoop-shaped, cymbial fold with row of setae similar to surrounding setae (i.e. not minute), embolus with an apical switch in the coiling direction as in Microdipoena   Edit ( Fig. 133G ), uniform abdominal dorsal colour pattern and anterior median eyes separate ( Fig. 142O ). There appears to be a subtle depression between the anterior median eyes, which could also be synapomorphic for the genus ( Fig. 142O ; see Wunderlich, 1995: fig. 10), although it has not been explicitly proposed as such in previous studies. Brasilionata   Edit has been previously diagnosed by somehow vague characters (Wunderlich, 1995), which when examined within a revisionary context can be assigned to any other mysmenid genus or are simply symplesiomorphies: lack of femoral spot on male, no trichobothrium on metatarsus IV, eight eyes equal in size, one metatarsal prolateral clasping spine, male palpal femur, patella and tibia without structures, cymbium long and distally slender, with a cymbial process, and embolus with apophysis. We could not find an embolic apophysis and instead we report a coiling switch of the distal part of the embolus.

MYSMENIOLA THALER, 1995   Edit

FIGS 134D , 142M, N

Mysmeniola Thaler, 1995: 429   Edit .

Type species

Mysmeniola spinifera Thaler, 1995   Edit by original designation and monotypy, holotype in MHNG, examined.

Familial placement and composition

Our working phylogenetic hypothesis places Mysmeniola   Edit within the mysmenine clade C128 ( Brasilionata   Edit , Mysmeniola   Edit , Microdipoena   Edit , and MYSM- 019- MAD), as sister to the clade including Brasilionata   Edit and Microdipoena   Edit (clade C126; Fig. 161B ). Mysmeniola   Edit is here represented by its type and only species M. spinifera   Edit .

Monophyly and diagnosis

This singular Venezuelan monotypic genus, only known by males (see male palp drawing on Fig. 134D ), is here diagnosed by the following combination of autapomorphies: six eyes (anterior median eyes absent; Fig. 142M ); prolateral cymbium with a prolateral basal expansion surrounding the entire basal bulb and with minute setae at tip, male palpal tibia with prolateral apical process and irregular rim setal conformation, median trichobothrium on metatarsus I, tarsus I with prolateral row of setae distributed along tarsus, and minute but distinct dorsal–posterior abdominal hump ( Fig. 142M, N ). The presence of a cluster of strong setae at the base of the clypeus seems to be autapomorphic for the genus (see Fig. 142M ; also Thaler, 1995: figs 1, 2). Previously proposed diagnostic features for Mysmeniola   Edit are in agreement with the diagnostic features suggested here.

Figures

Figure 17. Microdipoena elsae (Mysmenidae), male right palp, inverted images: A, prolateral view; B, ventral view; C, retrolateral view; D, retroventral view. See Appendix 3 for the list of abbreviations.

Figure 18. Microdipoena guttata (Mysmenidae): A–F, male left palp; G, female. A, ventral–proximal view; B, retrolateral– distal view; C, dorsal–retrolateral view; D, expanded bulb, prolateral view; E, prolateral view; F, expanded bulb, detail of tip of embolus, retrolateral–distal view; G, digested abdomen, detail of vulva. See Appendix 3 for the list of abbreviations.

Figure 19. Microdipoena guttata (Mysmenidae). A, B, D–F, male; C, female. A, prosoma, frontal view; B, C, same, lateral view; D, same, ventral view, detail of labium–sternum junction; E, mouthparts, detail of tip of left chelicera; F, posterior median spinnerets. See Appendix 3 for the list of abbreviations.

Figure 20. Microdipoena guttata (Mysmenidae), abdomen: A, female; B–G, male. A, ventral view; B, detail of booklung cover; C, detail of abdominal cuticular pattern; D, pedicel area and booklung covers; E, detail of area above pedicel; F, posterior–ventral view, detail of epigastric furrow, epiandrous spigots and spiracular openings; G, same, ventral view.

Figure 21. Microdipoena guttata (Mysmenidae), legs: A, D, F–I, male; B, C, E, female. A, left leg I, detail of femoral spot; B, right leg I, metatarsus–tarsus junction; C, right femur II, dorsal view; D, left patella I, dorsal view; E, right femur II, prolateral view; F, left tibia I, prolateral view; G, right metatarsus III, dorsal view; H, right femur II, ventral view; I, right leg III, metatarsal trichobothrium.

Figure 22. Microdipoena nyungwe (Mysmenidae): A, B, E, female; C, D, F, G, male left palp. A, digested abdomen, respiratory system and vulva; B, same, detail of vulva; C, distal–retrolateral view, detail of tip of embolus and primary cymbial conductor; D, ventral–proximal view; E, tip of palp and tarsal organ; F, dorsal view; G, retrolateral view. See Appendix 3 for the list of abbreviations.

Figure 23. Microdipoena nyungwe (Mysmenidae), spinnerets: A, B, female; C–H, male. A, C, anterior lateral spinnerets (ALS); B, E, left posterior lateral spinnerets; D, posterior median spinnerets; F, ALS, detail of major ampullate field; G, H, ALS, detail of intersegmental lobe. See Appendix 3 for the list of abbreviations.

Figure 24. Microdipoena nyungwe (Mysmenidae), abdomen: A–D, female; E–G, male. A, epigastric furrow, epigynal area and spiracular openings, posterior–ventral view; B, same, detail of epigynal area; C, pedicel area and booklung covers; D, epiandrous spigots, detail from figure F; E, posterior respiratory spiracle and colulus, ventral view; F, pedicel area, booklung covers and epiandrous spigots.

Figure 25. Microdipoena nyungwe (Mysmenidae), prosoma: A, male; B–F, female. A, lateral view; B, ventral view; C, ocular area, frontal view; D, cheliceral bases, frontal view; E, distal chelicerae; F, labrum.

Figure 26. Microdipoena nyungwe (Mysmenidae), left legs: A, C, D, G, male; B, E, F, female. A, tarsus I, prolateral view; B, claws I, retrolateral view; C, leg I, tibia–metatarsal junction, prolateral view; D, metatarsus I, dorsal view; E, leg I, metatarsus–tarsus junction, dorsal view; F, tarsus IV, prolateral view; G, tibia IV, prolateral view.

Figure 27. Microdipoena (= Mysmenella) samoensis (Mysmenidae), syntypes: A–C, F, H, I, male; D, E, G, female. A, left palp, prolateral–ventral view; B, same, retrolateral–ventral view; C, same, detail of tip of embolus and interaction with primary cymbial conductor, prolateral view; D, digested abdomen, vulva, and part of respiratory system; E, same, detail of vulva; F, G, prosoma, lateral view; H, mouthparts, ventral view, detail of retromargin of chelicerae and labium; I, left tibia I, dorsal 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 132. Mysmenidae male left palp, cleared: A, Microdipoena elsae, prolateral view, arrow to spine of basal prolateral cymbial expansion; B, Microdipoena guttata, prolateral view, arrow to spine of basal prolateral cymbial expansion; C, Microdipoena (= Mysmenella) illectrix, type, prolateral view; D, Microdipoena (= Mysmenella) samoensis, syntype, prolateral view; E, same, retrolateral view; F, Microdipoena (= Mysmenella) jobi, holotype, retrolateral 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 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 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 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.