MYSMENINAE PETRUNKEVITCH, 1928

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 : 782

MYSMENINAE PETRUNKEVITCH, 1928 (CLADE C133)

Composition

This morphologically distinct subfamily is circumscribed here to comprise the following genera: Anjouanella , Brasilionata , Calodipoena , Calomyspoena , Itapua, Kekenboschiella , Microdipoena , Mysmena , Mysmenella , Mysmeniola , and Tamasesia [ Fig. 160 , see below and refer to Fig. 161B for synonymies, see below for comments on recently erected Chinese genera not included in the analyses; see main text for the removal and suggested new familial status of Crassignatha , Iardinis , Leviola , and Phricotelus ; see Miller et al. (2009) for the removal of Crassignatha based on morphology; and see Rix & Harvey (2010) for the transfer of Taphiassa to Micropholcommatinae and its synonymy with Parapua ]. In our phylogenetic analyses, Mysmeninae included the following taxa: Microdipoena , Mysmeniola , Brasilionata , Mysmena (see below for total number of species and generic composition), and the following undescribed species (regarded here as Mysmeninae incertae sedis): MYSM-001-MAD, MYSM-006-MAD, MYSM-008-ARG, MYSM-009-MAD, MYSM-012-MAD, MYSM-019-MAD, MYSM-020-MAD, MYSM-021-MAD, MYSM-023-MAD, MYSM-026- MAD, MYSM-027-MAD, MYSM-029-MAD, MYSM-031-MAD, MYSM-032-MAD, and MYSM-033-MAD.

Monophyly

Relationships within Mysmeninae are unstable, except for a few taxa (see below). The following synapomorphies support Mysmeninae as we have circumscribed it: respiratory system with anterior tracheae restricted to opisthosoma ( Fig. 37A ; extending into prosoma in MYSM-005-ARG, MYSM-007- MEX, and Microdipoena s.s.); advanced and wide posterior spiracular opening located midway between the spinnerets and epigastric groove ( Fig. 24A, E ), connected to branched posterior lateral tracheae extending into prosoma ( Fig. 22A, B ); aciniform gland spigots of posterior spinnerets with two different outlines ( Figs 19F , 33D , 37 ,B E); posterior lateral spinnerets with an anterior flat spatulate modified seta [ Figs 23B, E , 33G, H , 52C ; secondarily absent in Mysmena (= Tamasesia ) rotunda ], and slim cylindrical spigots ( Figs 23B , 37B, E ); palpal tibial trichobothria of females lacking ( Figs 38E , 52E ); female copulatory openings within the epigastric furrow ( Fig. 24A ; secondarily external in MYSM-023-MAD), membranous atrium ( Figs 129A, E, G , 130B ), and irregular membranous copulatory ducts ( Figs 18G , 27D , 129A–C, E, H , 130A– G ; distally sclerotized independently in two clades); males with coiled embolus ( Figs 27A , 47A, B , 132D, E , 134G ) and secondary (external) cymbial conductor ( Figs 31C , 40A , 43C ; secondarily absent in clade C128: Microdipoena , Brasilionata , Mysmeniola , and MYSM-019-MAD; Fig. 22F ). Other synapomorphies include: abdomen with a whitish ventral ring surrounding the spinnerets ( Figs 142J, L , 143A, B, G, I, L ), trichobothria on tibia III and IV between two and three tibia diameters in length ( Figs 26G , 29E , 39G ; short trichobothria occurs independently within the group), and strongly serrated distal promarginal curved seta ( Figs 19E , 38H , 42E , 48B ). Ambiguously optimized synapomorphies for this clade include: a posterior tracheal arrangement consisting of lateral tracheae surrounding the minute median apodemes ( Fig. 29B ); fingerprint cuticular pattern on the piriform field ( Figs 23A, C , 33E ); flagelliform spigots absent in males ( Fig. 23E ; secondarily present in Mysmena leichhardti and MYSM-005-ARG); orb webs with a proliferation of out-of-plane radii both above and below the orb plane ( Fig. 147A, B ); males with prolateral row of slim setae occupying only distal half of tarsus I ( Figs 16H , 26A , 34D , 45I , 50F ; all along tarsus in Mysmeniola spinifera ); cymbial prolateral basal expansion ( Figs 27A , 30B, C , 36C , 47B ); embolus with pars pendula ( Figs 32H , 36B , 132C–F , 133C , secondarily absent in MYSM-005- ARG); palpal tibial rim setae longer and arranged distally in a row or two [ Figs 18A , 32E , 36D , 42B , 45C ; secondarily irregular in Mysmeniola spinifera and Microdipoena (= Mysmenella ) jobi ]; and females with weakly modified epigynal area (i.e. epigynum absent; Figs 14C , 37A , 52F ; modified copulatory area in MYSM-023-MAD, Fig. 49D, E ). A total of 163 molecular synapomorphies support this subfamily.

Diagnosis

Mysmeninae differs from all other mysmenid genera and subfamilies by the following unique combination of features: the characteristic architecture of their orb webs with a proliferation of out-of-plane radii both above and below the orb plane; anterior tracheae restricted to opisthosoma (extending into prosoma in MYSM-005-ARG, MYSM-007- MEX, and Microdipoena s.s.), and an advanced and wide posterior spiracular opening located midway between the spinnerets and epigastric groove, connected to branched posterior lateral tracheae extending into prosoma and surrounding the minute median apodemes; anterior lateral spinnerets with fingerprint cuticular pattern on the piriform field, posterior spinnerets with aciniform gland spigots of two different outlines, and posterior lateral spinnerets with an anterior flat spatulate modified seta [as in Maymena , secondarily absent in Mysmena (= Tamasesia ) rotunda ], slim cylindrical spigots, and flagelliform spigots absent in males (secondarily present in males of Mysmena leichhardti and MYSM-005-ARG); females without epigynum and with the copulatory openings within the epigastric furrow (secondarily external in MYSM-023- MAD), a membranous internal atrium and irregular membranous copulatory ducts (distally sclerotized independently in two clades); males with secondary (external) cymbial conductor (secondarily absent in clade C128: Microdipoena , Brasilionata , Mysmeniola , and MYSM-019-MAD), cymbial prolateral basal expansion, coiled embolus with pars pendula (secondarily absent in MYSM-005-ARG), palpal tibial rim setae long and arranged distally in a row or two [secondarily irregular in Mysmeniola spinifera and Microdipoena (= Mysmenella ) jobi ], a prolateral row of slim setae occupying only distal half of tarsus I (all along tarsus in Mysmeniola spinifera ); and also a strongly serrated distal promarginal curved seta, abdomen with a whitish ventral ring around the spinnerets, trichobothria on tibia III and IV of medium length.

Figures

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 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 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 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 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 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 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 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 45. MYSM-019-MAD (Mysmeninae, Mysmenidae) from Toliara, Madagascar. A–C, male right palp, inverted; A, dorsal view; B, retrolateral view; C, prolateral view. D, E, male left palp, expanded; D, prolateral–ventral view; E, same, detail of embolus and pars pendula; F, female posterior spinnerets; G–I, male left leg I, prolateral view; G, femur and patella; H, metatarsus; I, tarsus. See Appendix 3 for the list of abbreviations.

Figure 47. MYSM-020-MAD (Mysmeninae, Mysmenidae) from Toamasina, Madagascar, male. A–E, left palp; A, retrolateral view; B, prolateral–ventral view; C, same, detail of tip of embolus and cymbial conductors; D, same, ventral–retrolateral view; E, same, prolateral view. F, digested abdomen, tracheal system, dorsal view; G, same, detail of posterior tracheal system. See Appendix 3 for the list of abbreviations.

Figure 48. MYSM-020-MAD (Mysmeninae, Mysmenidae) from Toamasina, Madagascar, male: A, epiandrous spigots; B, cheliceral teeth and fang; C, D, right tibia I, detail of ventral paired setae with protruded bases, prolateral view; E, right tibia I, ventral view; F, right tarsus I, detail of tarsal organ, retrolateral view; G, right leg IV, claws.

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 50. MYSM-023-MAD (Mysmeninae, Mysmenidae) from Antananarivo, Madagascar: A, D, E, female; B, C, F–H, male. A, B, prosoma, lateral view; C, left chelicera, retromarginal view. D–H, right leg I, prolateral view; D, tarsus– metatarsus junction, prolateral–dorsal view; E, F, tarsus; G, tibia; H, metatarsus.

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 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 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 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 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.