Soesiladeepakius arthrostylus, Ruiz & Maddison, 2012

Ruiz, Gustavo R. S. & Maddison, Wayne P., 2012, DNA sequences corroborate Soesiladeepakius as a non-salticoid genus of jumping spiders: placement with lapsiines, phylogeny, and description of six new species (Araneae, Salticidae), Zoological Journal of the Linnean Society 165 (2), pp. 274-295 : 286-291

SOESILADEEPAKIUS ARTHROSTYLUS SP. NOV.

Type: Male holotype from Porto Urucu , Rio Urucu, Coari, Amazonas, Brazil, -4.895139, -65.188389, 11-20.vii.2003, A. B. Bonaldo et al., deposited in MPEG 11384 .

Etymology: The specific epithet is a Latin combination and refers to the apparently articulated embolus in the male palp. Noun in apposition.

Diagnosis: Males of this species can be easily distinguished from those of the other species by the apparently articulated embolus and the presence of a conductor.

Description: Male. Total length: 2.51. Carapace light brown, 1.23 long, 0.89 wide, 0.45 high, with darker posterior borders. Ocular quadrangle 0.61 long. Anterior eye row 0.95 wide, posterior 0.84 wide. Clypeus densely covered with white long scales. Chelicera yellow, prolaterally dark brown. Endite, labium, and sternum yellow. Palpal femur dorsally covered by a fringe of long white scales, ventrally with a retrolateral keel ( Fig. 49 ) and retrolaterally with a short, rectangular apophysis ( Fig. 44 ). Palpal tibia with a ventral round projection and a RTA reaching the proximal tegulum; cymbium with a prolateral proximal apophysis; tegulum round with an apophysis on the retrolateral portion (median apophysis?) and another on the prolateral proximal portion; embolus straight, articulated with tegulum and a triangular conductor-like structure (true conductor?) on the distal retrolateral portion ( Figs 41–44 ). Legs as in S. gasnieri sp. nov. Abdomen yellow, with three sparse, dark brown transverse stripes, anterior two medially interrupted; with a dorsal scutum covering all its length; two pairs of tufts of white scales on dorsal surface and another above the anal tubercle; ventrally with longitudinal, wide dark brown stripe. Scales as in S. gasnieri sp. nov. Spinnerets yellow, variegated with dark brown.

Female. Unknown.

PHYLOGENY OF SOESILADEEPAKIUS

The results of the phylogenetic analyses are presented in Figures 54–59 . Under parsimony ( Figs 54– 55 ), the monophyly of family Salticidae was recovered only in the combination 28S + Actin + 16S. In all the other gene combinations or analyses of single genes, Salticidae was nonmonophyletic. Hisponinae appeared as monophyletic in the analyses of 28S, 28S + Actin, 16S, 28S + Actin + 16S, and in the allgene matrix, whereas Lyssomaninae , as presently defined, was never recovered as monophyletic. Spartaeinae appeared as paraphyletic in the 28S-singlegene analysis, but its monophyly was recovered in the Actin, 28S + Actin, 28S + Actin + 16S, ND1 + 16S, and in the all-gene analyses. Salticoids were recovered as monophyletic in individual analyses of 28S and Actin, in 28S + Actin, 28S + Actin + 16S and in all-gene combinations, never including Soesiladeepakius . This genus, in turn, always appeared related to the lapsiine Galianora , although the monophyly of the lapsiines was recovered only in analyses with Actin-single gene, in the combinations ND1 + 16S, Actin + 28S and in the all-gene matrix.

Basic patterns seen in likelihood analyses coincide with those seen under parsimony ( Figs 56–58 ). For the 28S-single gene analysis, unlike parsimony, Hisponinae, Salticoida , and Spartaeinae are recovered as monophyletic, whereas Thrandina appears as the sister group of Lyssomanes , as observed under parsimony, failing in recovering the monophyly of lapsiines. The results found in the Actin-single gene analysis are very similar to those found under parsimony. The analysis of the combined ND1 + 16S corroborates the monophyly of Salticoida, Hisponinae , lapsiines, and Spartaeinae . Only lapsiines and Spartaeinae were recovered using the same fragments under parsimony, showing the ability of partitions/ models in helping reconstruct trees using only fastevolving genes. The all-gene analysis recovered Spartaeinae, Salticoida , and Hisponinae as monophyletic, but failed in recovering the monophyly of Salticidae , Lyssomaninae , and lapsiines. Nevertheless, Soesiladeepakius is strongly confirmed as closely related to Galianora .

Based on these results, we restricted our sampling for morphological studies. The morphological matrix scored comprised 12 taxa and 24 characters (see Material and methods and List of characters). The cladistic analysis resulted in two equally parsimonious trees of 29 steps ( CI = 86, RI = 93). The difference between the two relies on the affinities of Galianora ; the genus is the sister group of Soesiladeepakius in one tree whereas it forms a polytomy with Thrandina and Lapsias in the other. As the sister-group relationship is supported by DNA analyses, the former tree is preferred and it is used to discuss relationships and character evolution within Soesiladeepakius .

The cladistic analysis of morphological features suggests that the reduction of teeth number to a pair of retrolateral teeth on the chelicerae of both sexes could be a synapomorphy for lapsiines, although this would only be corroborated after more inclusive analyses.

The monophyly of Soesiladeepakius is supported by several apomorphic states. Synapomorphies of the genus are the male abdominal scutum (char. 3), abdominal clusters of white scales (char. 5), elongate ventral spines on front legs (char. 7), fringe of white scales on dorsal male palpal femur (char. 8), male palpal femur with retroventral keel (char. 9), a proximal tegular apophysis (char. 19), the loss of the flexible median apophysis (char. 20), the anterior epigynal pocket (char. 23), and the relocation of copulation openings on the female abdomen (char. 24). The shape of the fundus (char. 16) might also be a synapomorphy of Soesiladeepakius , but further studies are needed. Within the genus, a deep cladogenesis separated the lineage of the species with bare femora from the lineage of the ones with femoral apophyses. The group with no femoral apophyses is supported by apomorphic characters concerning the RTA (chars 13–14) and tegulum (char. 18). The group of species with femoral apophyses, supported by the femoral apophysis itself (char. 10) and eye row width (char. 1), is further divided into two, separating the strange S. arthrostylus from the remaining species, which have more complex and developed femoral structures (chars 11–12) and longer emboli (char. 22), besides the short RTA (char. 13) and the prolateral tegular flap (char. 15).

Figures

Figure 54. Phylogenetic reconstructions of salticid lineages under parsimony: majority consensus (0.5) using one (28S: 13 trees; Actin: two trees; 16S: six trees), two (ND1 + 16S: three trees; 28S + Actin: one tree), and three gene matrices (28S + Actin + 16S: six trees). Numbers on branches show group frequency. Names in bold letters and grey boxes represent salticid taxa.

Figure 55. Phylogenetic reconstruction of salticid lineages under parsimony: single most parsimonious tree using the all-gene matrix. Names in bold letters and grey boxes represent salticids, whereas names on the right of the vertical bars represent major salticid taxa.

Figure 56. Phylogenetic reconstructions of salticid lineages under maximum likelihood: best trees using single gene regions. Branch length is proportional to substitution rate per site. Names in bold letters represent salticid taxa.

Figure 57. Best tree of phylogenetic reconstruction analyses of salticid lineages using maximum likelihood and the all-gene matrix. Branch length is proportional to substitution rate per site. Names in bold letters represent salticid taxa.

Figure 58. Consensus and bootstrap values (500 pseudoreplicates) found by maximum likelihood of the all-gene matrix. Names in bold letters represent salticids, whereas names on the right of the vertical bars represent major salticid taxa.

Figure 59. Chosen best tree of phylogenetic reconstruction of the genus Soesiladeepakius using morphological data. White and black circles on branches represent, respectively, homoplasious and nonhomoplasious transformations. Numbers over branches are Bremer support values. Numbers under circles indicate number of listed characters. For discussion on question marks under these numbers, see List of characters.

Figure 47–52. Soesiladeepakius spp., left male palpal femora, ventral view. 47, Soesiladeepakius lyra sp. nov. 48, Soesiladeepakius retroversus sp. nov. 49, Soesiladeepakius arthrostylus sp. nov. 50, Soesiladeepakius biarmatus sp. nov. 51, Soesiladeepakius gasnieri sp. nov. 52, Soesiladeepakius uncinatus sp. nov.

Figure 41–46. Soesiladeepakius arthrostylus sp. nov. 41, left male palp, prolateral view. 42, ventral view. 43, retrolateral view. 44, ventroretrolateral view. 45, bulb, clarified, ventral view. 46, posterior view. co?, putative true conductor; ma?, putative median apophysis.