Smeringopinae Simon, 1893

Huber, Bernhard A., Eberle, Jonas & Dimitrov, Dimitar, 2018, The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae), ZooKeys 789, pp. 51-101: 60-63

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Smeringopinae Simon, 1893


Subfamily Smeringopinae Simon, 1893  Figure 6

Smeringopodeae  Simon, 1893: 474. Type genus Smeringopus  Simon, 1890, by subsequent designation ( Huber 2011b).

Smeringopinae  Simon; Huber 2011b: 217.


Smeringopinae  is a relatively homogeneous subfamily (with respect to body shapes, colour, webs, and microhabitats), and in this sense similar to Ninetinae  and Arteminae  but very unlike Modisiminae  and Pholcinae  . Most of the 125 known species of Smeringopinae  are medium-size to large, have long legs, elongated to cylindrical abdomens, and all have eight eyes. Another similarity to Ninetinae  and Arteminae  is that Smeringopinae  are often found in rather arid regions. The most obvious exception is the largely humid tropical genus Smeringopina  Kraus, 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-Mađarić et al. 2005, Astrin et al. 2007, Dimitrov et al. 2013), Smeringopinae  is 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 Smeringopinae  receives 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 Ninetinae  in Astrin et al. 2007]. Holocnemus pluchei  was included in preliminary analyses of the present data but its position was drastically unstable, so we decided to exclude it from the final analyses. Smeringopinae  monophyly 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, Hoplopholcus  Kulczynski, 1908 is sister to all other genera and not close to Stygopholcus  Kratochvil, 1932 as repeatedly claimed by Brignoli (1971, 1976, 1979) but contested by Senglet (1971, 2001). The genera Hoplopholcus  , Stygopholcus  , and Crossopriza  Simon, 1893 all receive full support, but the small Mediterranean genus Holocnemus  Simon, 1873 (only three described species) continues to be problematic even after the exclusion of H. pluchei  . The two species of Holocnemus  included in our analyses never group together, and no morphological synapomorphy is known to suggest their sister-group relationship (in fact, Holocnemus  has never been revised).

The southern (Sub-Saharan) clade includes Smeringopus  Simon, 1890 and Smeringopina  , and is also supported by a unique number of epiandrous spigots (two) ( Huber 2012). The paraphyly of Smeringopus  has 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 Smeringopus  proposed in Huber (2012) appear closer to Smeringopina  than to other Smeringopus  : the chogoria  group 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, Smeringopus  and Smeringopina  are largely separated geographically, with Smeringopus  being most diverse in southern and eastern Africa, and Smeringopina  in western and central Africa ( Huber 2012, 2013). The chogoria  and 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 Smeringopus  is not dense enough to test the species groups proposed in Huber (2012). Remarkably, though, the isolated ‘basal’ position of S. ngangao  Huber, 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. fon  Huber, 2013; S. ngungu  Huber, 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 ankasa  and 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).