Smeringopodeae Smeringopus Smeringopinae The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae) Huber, Bernhard A. Eberle, Jonas Dimitrov, Dimitar ZooKeys 2018 789 51 101 Simon, 1893 Simon 1893 Arachnida Pholcidae CoL Animalia Araneae 9 60 Arthropoda subFamily Smeringopinae    SmeringopodeaeSimon, 1893: 474. Type genus SmeringopusSimon, 1890, by subsequent designation ( Huber 2011b).  SmeringopinaeSimon; Huber 2011b: 217.  Remarks.  Smeringopinaeis a relatively homogeneous subfamily (with respect to body shapes, colour, webs, and microhabitats), and in this sense similar to Ninetinaeand Arteminaebut very unlike Modisiminaeand Pholcinae. Most of the 125 known species of Smeringopinaeare medium-size to large, have long legs, elongated to cylindrical abdomens, and all have eight eyes. Another similarity to Ninetinaeand Arteminaeis that Smeringopinaeare often found in rather arid regions. The most obvious exception is the largely humid tropical genus SmeringopinaKraus, 1957, which is also the genus with the smallest and largest representatives in the subfamily (with body lengths ranging from 2.5-10 mm) and with the widest range of microhabitats used (leaf litter to large sheltered spaces) ( Huber 2013). The original distribution of the subfamily is Africa, the Mediterranean, and the Middle East. Three species have attained much wider distributions, resulting from human-mediated dispersal ( Huber 2011b). As in previous molecular analyses (  Bruvo-Mađaricet al. 2005, Astrin et al. 2007, Dimitrov et al. 2013), Smeringopinaeis sister to Pholcinae(Figure 1) with reasonable to high support. This relationship is also supported by morphology: the two taxa share tarsus IV comb-hairs spread over the entire length of the tarsus ( Huber and Fleckenstein 2008). The monophyly of Smeringopinaereceives reasonable to high support in all our analyses. Previous molecular analyses have partly supported Smeringopinae, but also suggested rather obscure relationships [e.g., the position of Holocnemus pluchei(Scopoli, 1763) among Ninetinaein Astrin et al. 2007]. Holocnemus plucheiwas included in preliminary analyses of the present data but its position was drastically unstable, so we decided to exclude it from the final analyses. Smeringopinaemonophyly is rather weakly supported by morphology, i.e., by the presence of a large thoracic pit on the carapace (rather than a narrow furrow or an evenly domed carapace; cf. Huber 2011b).  Within Smeringopinae, our data strongly support a basal split between a northern clade (Mediterranean, northern Africa, Middle East, Central Asia) and a southern clade (Sub-Sahara) (Figure 6). This basal split was also recovered in a morphological cladistic analysis ( Huber 2012). Within the northern clade, HoplopholcusKulczynski, 1908 is sister to all other genera and not close to StygopholcusKratochvil, 1932 as repeatedly claimed by Brignoli (1971, 1976, 1979) but contested by Senglet (1971, 2001). The genera Hoplopholcus, Stygopholcus, and CrossoprizaSimon, 1893 all receive full support, but the small Mediterranean genus HolocnemusSimon, 1873 (only three described species) continues to be problematic even after the exclusion of H. pluchei. The two species of Holocnemusincluded in our analyses never group together, and no morphological synapomorphy is known to suggest their sister-group relationship (in fact, Holocnemushas never been revised).  Figure 6. Smeringopinae a Hoplopholcussp. n. "Mar66"(Turkey) b Stygopholcus absoloni? (Bosnia and Herzegovina) c Crossoprizasp. n. "Om11"(Oman) d Smeringopus pallidus(Philippines) e Smeringopina pulchra(Ghana) f Smeringopina ankasa(Ghana).  Thesouthern (Sub-Saharan) clade includes SmeringopusSimon, 1890 and Smeringopina, and is also supported by a unique number of epiandrous spigots (two) ( Huber 2012). The paraphyly of Smeringopushas been suggested before ( Dimitrov et al. 2013), and our larger data set supports this view, but with low support values. Two of the species groups of Smeringopusproposed in Huber (2012)appear closer to Smeringopinathan to other Smeringopus: the chogoriagroup and the rubrotinctus group. Morphological data do not support this view but they neither strongly contradict it: the two species groups lack the distinctive arrangement of pores on the pore plates (in groups or 'islands') and the retrolateral furrow on the male palpal femur present in all other species of Smeringopus( Huber 2012). Remarkably, Smeringopusand Smeringopinaare largely separated geographically, with Smeringopusbeing most diverse in southern and eastern Africa, and Smeringopinain western and central Africa ( Huber 2012, 2013). The chogoriaand rubrotinctus groups are geographically restricted to an area where Central Africa (the Guineo-Congolian center of endemism) meets East Africa ( Huber 2012). Other than that, our sampling in Smeringopusis not dense enough to test the species groups proposed in Huber (2012). Remarkably, though, the isolated 'basal'position of S. ngangaoHuber, 2012 is supported by the present analyses. Our analyses include 30 of the 44 described species of Smeringopina(68%), and all species groups proposed in Huber (2013)except two monotypic 'groups'( S. fonHuber, 2013; S. ngunguHuber, 2013). Even though for some species only one gene (CO1) was sequenced, our analyses support several species groups and deeper relationships proposed previously ( Huber 2013), based on cladistic analysis of morphological characters. Morphology placed the West African guineensis group as sister to all other Smeringopina; all our analyses support both the monophyly of the guineensis group and its sister-group relationship with all other congeners. The next two branches are composed of representatives of the lekoni group, which is thus here considered paraphyletic rather than monophyletic. The ankasaand cornigera groups are both supported, as is their sister group relationship to each other. The attuleh group is supported, but not as sister to the ankasa+ cornigera groups but as sister to the following group. The last clade is composed of representatives of the simplex and beninensis groups, but the clear dichotomy in the molecular trees is not equivalent to these groups. Instead, the simplex group includes all 'basal'representatives originally assigned to the beninensis group; the beninensis group includes only those species that have a light transversal element ventrally on the abdomen (character 9 in Huber 2013, which is thus less homoplastic than previously thought).