Arteminae 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: 54-56

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Arteminae Simon, 1893


Subfamily Arteminae Simon, 1893  Figure 2

Artemeae  Simon, 1893: 463. Type genus Artema  Walckenaer, 1837, by monotypy.

Arteminae  Simon; Huber 2011b: 212.


All our analyses exclude the name-giving genus Artema  from the clade containing all other Arteminae  and invariably place Artema  as sister to Ninetinae  (Figure 2), formally precluding the use of the name Arteminae  for this clade. We do not propose a new subfamily name for this clade but treat it as 'other Arteminae  ' because we consider the position of Artema  dubious. Artema  shares with 'other Arteminae  ' a unique pair of structures on the procursus: a ventral pocket and a dorsal apophysis. These structures are associated with asymmetric palp insertion in both species studied with respect to this detail [ Physocyclus globosus  (Taczanowski, 1874), Artema nephilit  Aharon et al., 2017: Huber and Eberhard 1997, Aharon et al. 2017]. The structures are present in all Arteminae  , even in taxa that were previously thought to be representatives of other subfamilies, such as Chisosa  and Nita  (previously in Ninetinae  ; see above), and Wugigarra  Huber, 2001 (previously in Modisiminae  ; see below) ( Huber 2000, 2001, Huber and El Hennawy 2007). By contrast, these structures are apparently absent in all other Pholcidae  . Curiously and unexplainable to us, previous molecular analyses have supported a position of Artema  among 'other Arteminae  ' ( Astrin et al. 2007: fig. 1, Dimitrov et al. 2013).

Some of the 99 currently known species of Arteminae  are relatively large spiders with long, strong legs and high globose abdomens. The genus Artema  , in particular, includes probably the largest pholcids in terms of body mass ( Aharon et al. 2017). However, tiny species that were previously assigned to Ninetinae  partly because of their size ( Chisosa  , Nita  ) are now included in Arteminae  , and their ‘basal’ position in the cladogram suggests that ancestral Arteminae  may in fact have been tiny. Just like Ninetinae  , Arteminae  often occur in rather dry regions, sometimes even in deserts like the Australian Trichocyclus  Simon, 1908. They have a wide distribution, but are apparently absent from Sub-Saharan Africa and from South America (except for “Geneve59”, a tiny undescribed species representing a new undescribed genus on Curaçao and Aruba).

The monophyly of 'other Arteminae  ' is supported in all our analysis, even though with low support (possibly because of the dubious position of Artema  , see above). Similar to our previous analysis (i.e. except for the position of Artema  ; Dimitrov et al. 2013), 'other Arteminae  ' is sister to Modisiminae  , with variable support (reasonable support only in the RogueNaRok tree; in other trees, bootstrap support is low but SH values range from 82 to 99). This sister group relationship is weakly supported by morphology: 'other Arteminae  ' and Modisiminae  lack epiandrous spigots. However, epiandrous spigots have been lost several times convergently in Pholcidae  ( Huber 2000, BA Huber, unpubl. data).

Internal relationships in 'other Arteminae  ' are partly resolved with reasonable support. The data suggest a large Indomalayan-Australasian clade, including the genera Trichocyclus  and Wugigarra  (Australia), Holocneminus  Berland, 1942 (SE Asia and Pacific; excluding the misplaced and highly isolated H. huangdi  Tong & Li, 2009), and a new undescribed genus (without any described species; ranging from Eastern Indonesia to the Pacific). Sister to this clade is either the New World genus Physocyclus  Simon, 1893 alone or Physocyclus  together with the Middle-Eastern monotypic Nita  . However, support values for any of these options are low and morphological data do not favour (nor contradict) any of them. Finally, the ‘basal’ branches, i.e., those leading to the taxa outside the Indomalayan-Australasian clade and Physocyclus  (and Nita  in the case of the IQ-TREE analysis) lead to a group of North American and Caribbean taxa (the North American genus Chisosa  being sister to a tiny undescribed species representing a new undescribed genus on Curaçao and Aruba: “Geneve59”), and to the SE-Asian Holocneminus huangdi  , an isolated species that appears misplaced also by morphological criteria (A Valdez-Mondragón, pers. comm., Nov. 2015).