Cladistics Cladistics 30 (2014) 67–106 10.1111/cla.12025 Phantoms of Gondwana?—phylogeny of the spider subfamily Mynogleninae (Araneae: Linyphiidae) Holger Fricka,b,* and Nikolaj Scharffb,c aDepartment of Invertebrates, Natural History Museum of Bern, Bernastrasse 15, 3005 Bern, Switzerland; bDepartment of Entomology, Natural History Museum of Denmark, Zoological Museum, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen, Denmark; cCenter of Macroecology, Evolution and Climate, Biological Institute, Universitetsparken 15, 2100 Copenhagen, Denmark Accepted 25 February 2013 Abstract This is the first genus-level phylogeny of the subfamily Mynogleninae. It is based on 190 morphological characters scored for 44 taxa: 37 mynoglenine taxa (ingroup) representing 15 of the 17 known genera and seven outgroup taxa representing the sub- families Stemonyphantinae, Linyphiinae (Linyphiini and Micronetini), and Erigoninae, and a representative of the family Pimoi- dae, the sister-group to Linyphiidae. No fewer than 147 of the morphological characters used in this study are new and defined for this study, and come mainly from male and female genitalia. Parsimony analysis with equal weights resulted in three most parsimonious trees of length 871. The monophyly of the subfamily Mynogleninae and the genera Novafroneta, Parafroneta, Laminafroneta, Afroneta, Promynoglenes, Metamynoglenes, and Haplinis are supported, whereas Pseudafroneta is paraphyletic. The remaining seven mynoglenine genera are either monotypic or represented by only one taxon. Diagnoses are given for all genera included in the analysis. The evolution of morphological traits is discussed and we summarize the diversity and distribu- tion patterns of the 124 known species of mynoglenines. The preferred topology suggests a single origin of mynoglenines in New Zealand with two dispersal events to Africa, and does not support Gondwana origin. © The Willi Hennig Society 2013. Introduction The family represents the second most diverse family of spiders, with 4429 described species (Platnick, 2013) Linyphiidae are the most species-rich spider family in 590 genera, surpassed only by the family Salticidae, within the superfamily Araneoidea, a large clade that with 5570 species in 591 genera. includes one-third of all described spider species The spider family Linyphiidae has been divided into (Blackledge et al., 2009). Linyphiidae have a world- a number of different subfamilies, most of which are wide distribution but are most diverse in colder not monophyletic according to modern phylogenetic regions. At higher latitudes, this family dominates the studies. Currently used subfamilies of Linyphiidae are spider faunas. For instance, 41% of the Danish spider Stemonyphantinae, Mynogleninae, Erigoninae, and fauna (216 of the 523 species) are linyphiids (Scharff Linyphiinae (Micronetini plus Linyphiini; Arnedo and Gudik-Sørensen, 2006) and most species are et al., 2009), of which Mynogleninae with 17 genera widely distributed in northern Europe. At lower lati- and 124 species are one of the smallest and most tudes, the family is well represented but constitutes a recently established. Most species have been described much smaller fraction of the total spider fauna within the past 35 years. The subfamily Mynogleninae (Scharff, 1992, 1993; Sørensen et al., 2002). was established by Lehtinen (1967: 250) for Mynog- lenes insolens Simon, 1905 (now Haplinis rufocephala *Corresponding author. (Urquhart, 1888)) and is “mainly characterized by the E-mail address: [email protected] presence of numerous metatarsal trichobothria and the © The Willi Hennig Society 2013 68 H. Frick and N. Scharff / Cladistics 30 (2014) 67–106 exceptional type of modification of the carapace”. The first mynoglenine was described by Urquhart Unfortunately, Lehtinen (1967) did not specify what (1886) and placed in the well known European genus kind of “exceptional modifications of the carapace” he Linyphia. Simon described the genera Haplinis (1894) used to define the subfamily. A quick look at the cara- and Mynoglenes (1905), but neither Simon nor Urqu- pace of any known mynoglenine reveals that it cannot hart mentions the special cephalic pits in their descrip- be anything but the very characteristic cephalic pits tions. Today we know that most members of this with which all mynoglenines are equipped—males as subfamily occur in the Southern Hemisphere, and that well as females, and juveniles—just below the anterior they have a disjunct distribution pattern with centres lateral eyes (subocular sulci; Figs 14b and 15c: SOS). of diversity in tropical Africa (33 species) and New This is still the best morphological character to distin- Zealand (90 species; including the South Pacific guish between mynoglenines and members of all other islands) and in Australia (one species; Tasmania). linyphiid subfamilies (Blest, 1979: 96). Members of the More species undoubtedly remain to be discovered, subfamily are otherwise rather indistinct, being small especially in Africa where the mynoglenine fauna is to medium in size (2–10 mm) and without any other still very poorly surveyed, but even if more species cephalic modifications or characteristic abdominal should turn up in Africa, the subfamily is still small markings (Fig. 13e and 15a). compared to the Erigoninae, another well defined sub- Pimoa altioculata Stemonyphantes lineata Linyphia triangularis 1 3 >20/100 1/- Bolyphantes luteolus 2 7/93 Australolinyphia remota 4 Diplostyla concolor 3/70 5 Hilaira excisa 5/87 Cassafroneta forsteri 6 5/79 Novafroneta gladiatrix 9 7 11/100 Novafroneta vulgaris 5/84 Hyperafroneta obscura 11 6/86 Metafroneta sinuosa 8 5/72 Protoerigone otagoa 13 1/- Pseudafroneta perplexa Mynogleninae Pseudafroneta pallida 10 1/- 16 Parafroneta minuta 1/- 17 Parafroneta haurokoae 1/- 18 Parafroneta demota clade 15 1/- 19 Parafroneta confusa 12 15 1/- 1/- 20 Parafroneta marrineri 1/- 1/- 21 1/- Parafroneta westlandica Trachyneta jocquei 22 Gibbafroneta gibbosa 1/- Laminafroneta bidentata 23 1/- 25 14 8/99 Laminafroneta brevistyla 1/- 24 1/- Afroneta bamilekei 26 Afroneta subfusca 1/- 27 Afroneta guttata 1/- 28 Afroneta lobeliae 1/- 29 7/99 Afroneta tenuivulva Afromynoglenes parkeri Promynoglenes silvestris 32 30 9/99 Promynoglenes nobilis 2/- 33 5/82 Promynoglenes grandis clade 30 31 Metamynoglenes helicoides 6/95 35 Metamynoglenes gracilis 5/97 36 2/- Metamynoglenes incurvata 34 2/- Haplinis insignis Haplinis abbreviata 37 7/83 Haplinis diloris 40 38 1/- 3/76 Haplinis horningi Haplinis subdola 41 Haplinis tegulata 1/- 42 7/99 Haplinis titan Fig. 1. Strict consensus tree of three most parsimonious trees found using equal weights (L = 873, CI = 0.28, RI = 0.62) including support val- ues (Bremer before and Jackknife after the slash). Node numbers in circles (monophyletic genera in black). The recent distribution is indicated for Africa (dark grey) and Oceania (light grey). Note that node 39 collapses and is not depicted in the consensus tree. H. Frick and N. Scharff / Cladistics 30 (2014) 67–106 69 ra lexa rp pe obscu a ta a et Figure 2b Figure 4 Figure 3 one a r tago o 6 rafron 15 30 udaf pe se Hy P gone i r 144 0 190 0 e 11 6 0 adiatrix 188 1 l lgaris 87 1 178 2 to g 168 0 u 190 1 86 2 ta 165 0 163 0 185 2 85 1 a o 1591 157 0 t 183 1 84 1 Pro 156 0 e 158 1 m 165 1 83 1 142 0 155 0 Metafroneta sinuos Metafroneta e 157 1 79 0 137 0 154 0 153 1 78 1 121 0 90 1 187 1 143 1 169 0 72 1 ia r 120 0 67 2 162 2 100 2 163 0 70 1 11 5 1 60 1 184 1 158 2 87 1 159 1 65 1 ph 104 2 32 1 95 1 151 1 85 1 158 2 56 1 y 101 1 21 0 74 1 Novafron 133 1 81 1 151 1 50 0 Diplostyla concolor Diplostyla 87 0 19 1 39 0 Novafroneta v 121 1 79 0 133 1 45 1 86 0 4 1 25 0 94 1 72 1 104 0 22 1 81 0 1 2 81 91 1 38 1 67 0 17 1 164 2 65 1 176 1 11 39 0 25 0 14 141 1 40 0 174 2 164 2 Linyphia triangularis Linyphia 12 20 1 168 1 149 0 187 1 ustralolin 84 1 31 1 11 7 0 Bolyphantes luteolus 13 83 1 12 0 121 1 142 0 149 0 11 4 0 A 5 67 0 140 0 146 0 189 1 105 1 187 1 62 0 100 2 145 1 184 1 98 2 186 1 178 1 183 0 38 1 61 0 131 1 11 8 1 59 1 184 2 172 0 179 1 177 0 25 0 39 0 127 1 91 35 0 183 2 170 1 171 1 175 0 9 91 1 81 31 0 182 2 169 1 149 1 174 0 89 0 12 148 1 172 0 165 1 86 4 173 2 165 0 i 168 2 159 1 146 0 170 0 144 1 84 1 148 1 167 2 158 1 134 1 169 0 143 1 57 1 147 1 164 2 141 1 133 1 159 1 11 7 0 56 1 138 0 161 1 138 1 11 0 1 158 2 11 6 0 50 2 134 1 147 1 127 1 100 2 152 0 95 1 49 1 11 2 1 145 0 126 1 98 2 150 0 forster 82 1 36 1 43 1 144 2 121 0 90 1 145 0 a 74 1 10 139 0 107 1 88 1 144 2 68 1 137 0 103 0 85 1 143 0 44 1 120 1 125 0 97 1 74 1 138 1 42 1 86 1 ronet 41 0 59 0 122 1 91 0 69 2 11 6 0 f 11 9 1 87 0 63 1 107 1 31 0 48 1 11 5 1 84 2 55 1 99 0 28 1 47 1 11 4 1 66 0 11 1 27 2 20 1 35 1 70 1 35 1 4 71 2 1 31 1 1 9 1 27 2 Cassa 6 1 23 1 184 0 8 3 178 0 176 0 182 2 185 1 11 7 0 190 1 178 1 106 1 105 2 167 0 177 1 65 1 104 1 154 0 174 1 antes lineata antes 60 1 81 1 151 0 169 1 45 1 75 0 141 1 163 1 ph 22 0 40 1 130 0 162 1 y 17 1 28 2 128 1 159 2 12 10 127 1 158 3 on 0 1 2 126 1 142 1 11 5 0 138 2 ata l 189 0 102 0 137 1 99 1 11 4 1 Stem 188 0 cu 186 0 98 2 11 1 1 o 185 0 95 2 11 0 1 ti l 183 1 89 0 101 0 187 1 180 0 62 0 91 0 182 2 171 0 57 1 65 0 173 2 140 1 56 1 22 0 149 1 121 1 28 0 21 1 142 0 120 1 12 19 2 138 2 11 4 0 Pimoa a ira excisa 7 133 1 96 1 a 125 0 64 1 182 1 11 9 1 62 1 149 1 Hil 184 2 103 0 61 1 11 7 1 183 2 84 2 47 1 11 6 1 181 1 83 1 34 0 11 3 1 179 2 23 1 29 0 164 0 108 1 176 2 11 4 0 158 1 106 1 174 2 1 155 0 92 1 168 2 140 0 79 1 167 2 177 1 125 0 77 0 164 2 175 1 78 1 69 1 163 2 157 1 66 0 67 1 162 2 143 1 41 0 63 1 158 4 11 7 1 35 1 55 1 153 1 76 1 34 1 47 0 84 1 75 1 32 1 39 1 41 2 54 1 29 1 27 1 26 1 27 1 28 3 25 1 91 22 1 10 0 19 1 81 12 1 (a) 5 1 61 (b) 6 Fig.