Research Article |
Corresponding author: Wayne P. Maddison ( wmaddisn@mail.ubc.ca ) Academic editor: Jeremy Miller
© 2016 Wayne P. Maddison.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Maddison WP (2016) Sumakuru, a deeply-diverging new genus of lyssomanine jumping spiders from Ecuador (Araneae: Salticidae). ZooKeys 614: 87-96. https://doi.org/10.3897/zookeys.614.9368
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The lyssomanine jumping spider genus Sumakuru gen. n. is here described for Sumakuru bigal sp. n., from the Bigal River Biological Reserve in Ecuador. Known from a single male, the embolus of the palp takes the form of a smoothly arching curve, and appears fully mobile, being connected to the tegulum by a thin sclerite and a twisted hematodocha. Data from four gene regions (28S, 16SND1, CO1, wingless) indicate that Sumakuru is the sister group to all other sampled lyssomanines, diverging deeply on the stem lineage of the clade of other known lyssomanines. Unlike previous molecular results, the sampled species of Lyssomanes Hentz, 1845 are supported as monophyletic, with Chinoscopus Simon, 1900 as the sister to Lyssomanes.
Jumping spider, Lyssomaninae , molecular phylogeny, new genus, new species
The distinctive lyssomanine jumping spiders include two described genera, Lyssomanes Hentz, 1845 and Chinoscopus Simon, 1900, both neotropical (
A single male of a new species of lyssomanine from the Bigal River Biological Reserve in Ecuador bears a distinctive palp, but it appears to be within the range of morphological diversity in Lyssomanes (
The preserved specimen was examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope.
Terminology is standard for Araneae. All measurements are given in millimeters. Descriptions of color pattern are based on the alcohol-preserved specimen. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; PLE, posterior lateral eyes; PME, posterior median eyes (the “small eyes”).
DNA sequences of the genes or gene regions 28S, wingless, CO1 and 16SND1 were obtained from the holotype of Sumakuru bigal using the protocols of
Prior to phylogenetic analysis, multiple sequence alignment was done for 28S and the noncoding portion of 16SND1 with MAFFT (
Maximum likelihood phylogenetic analyses were run using RAxML version 8.2.8 (
To choose a partitioning scheme for the RAxML analyses, PartitionFinder 1.1.1 (
Alignments and trees are deposited in the Dryad data repository (https://doi.org/10.5061/dryad.2g8j2).
Sumakuru bigal Maddison, sp. n.
From the Quechua sumak, “great, marvellous” and uru, “spider”. The same root sumak is the source of the name of the volcano Sumaco, from whose southeastern slopes the type species is known. Sumakuru is to be treated as grammatically masculine.
Delicate, pale, and long legged as in other lyssomanines, but with a distinctive palp in which the smoothly arching embolus is connected to the tegulum by a thin sclerite and twisted hematodocha (Fig.
Sumakuru bigal sp. n., holotype, except 7 (Lyssomanes for comparison). 1–6 Left palp. 1 Prolateral view 2 Oblique prolateral-ventral view (scale bar 0.1 mm) 3 Ventral view 4 Retrolateral view 5 Oblique prolateral-ventral view showing path of spermophore (as seen by clove oil clearing) and interpretation of parts 6 Same, ventral view 7 Trypsin-cleared left palp of Lyssomanes viridis, ventral view, showing path of spermophore 8 Carapace, dorsal view 9 face (scale bar 1.0 mm) 10 Carapace, lateral view 11 Photograph of holotype, with left palp and left second leg separated. Abbreviations: e = embolus; c = conductor; ma = median apophysis; t = tegulum; st = subtegulum.
Holotype: male, ECUADOR: Orellana: Río Bigal Reserve, Mirador Trail. S 0.5282 W 77.4195. 950 m elev. 2–4 November 2010. W & D Maddison, M Vega, M Reyes. WPM#10-043. DNA voucher d448. The specimen pertains to the
Based on the type locality.
The distinct arching spiral of the embolus (Fig.
The single male was found by beating understory vegetation in a relatively open tropical rainforest along a ridge. It landed injured on the beating sheet, having lost most of its legs. The preserved specimen now has both palpi, but just 3 legs: the second legs on both sides, and the third leg on the right side.
Male (holotype, DNA voucher d448). Carapace length 1.7; abdomen length 2.8. Chelicera (Fig.
Sequences of the holotype of Sumakuru bigal were obtained for 28S (GenBank accession number KX578224), 16SND1 (KX578225), CO1 (KX578226) and wingless (KX578227). Alignment by MAFFT appeared reasonable except for a few obvious shifts of 3 to 7 nucleotides near the starts or ends of some outgroup sequences, which were corrected by hand (28S Aelurillus cf. ater, Cucudeta zabkai, Onomastus nigrimaculatus, Cocalodes longicornis; 16SND1 Afromarengo sp., Naphrys pulex).
PartitionFinder determined the best partitioning scheme was to keep all partitions separate except to group the noncoding portion of 16SND1 with the first codon position of ND1, and to group first and second codon positions of wingless. For all of these GTR+I+G was determined as the best model, except for ND1 second position (GTR+G).
Phylogenetic trees inferred are shown in Figs
Maximum likelihood phylogenetic trees from RAxML analyses. Appended to taxon names are the identification codes of voucher specimens used 12 Phylogeny from all 4 genes concatenated; bootstrap percentages shown for lyssomanines only 13 Lyssomanine portion of phylogeny from 28S alone 14 Lyssomanine portion of phylogeny from three gene regions concatenated, 16SND1+CO1+wingless.
Sumakuru is placed with strong support as a lyssomanine and as the sister group to Chinoscopus + Lyssomanes. The monophyly of the Lyssomaninae is supported in 100% of the bootstrap replicates in the All Genes analysis (Fig.
The molecular phylogeny provides sufficient reason to recognize Sumakuru as a distinct genus, given the study’s inclusion of the type species of Lyssomanes (L. viridis) and the close similarity between the species of Chinoscopus studied and the type species C. gracilis (Taczanowski, 1872). However, the molecular phylogeny does not provide much support for the full monophyly of Lyssomanes as currently composed, as we lack molecular data for many species now placed in Lyssomanes, and there has been little phylogenetic work using morphology. The delicate body and unusual genitalia of Chinoscopus can be interpreted as synapomorphies for that genus, but no such synapomorphies are known for Lyssomanes as a whole. Although many excellent figures of male palps of Lyssomanes species have been published (e.g.
That Lyssomanes could be monophyletic receives some support from the fact that the ten Lyssomanes species in the molecular phylogenetic analysis cover a broad spectrum of the genus, including representatives of
Among the Lyssomanes species not studied in the molecular phylogeny are some whose similarities to S. bigal are likely convergent. For instance, Lyssomanes spiralis F.O. Pickard-Cambridge, 1900 also has a spiraled embolus (
Some species currently placed in Lyssomanes stand out as unusual, therefore possibly falling outside of Lyssomanes, either as relatives of Sumakuru bigal, or as separate deeply-branching lineages. Lyssomanes tarmae Galiano, 1980 has a distinctive palp with an arching embolus that resembles that of Sumakuru bigal, though more robust. L. elongatus Galiano, 1980, known only from the female, has a carapace shape similar to S. bigal. Although not obviously similar to Sumakuru bigal, the species Lyssomanes romani Logunov, 2000 may have a special phylogenetic position, based on the median apophysis which is apparently articulated (
Sumakuru appears to have diverged relatively long ago from the lineage leading to Lyssomanes and Chinoscopus, given the length of the branch below the clade of Lyssomanes + Chinoscopus (Fig.
I thank Thierry Garcia and Marion Hiruois of the Fundación Ecológica Sumac Muyu for their work to preserve the Río Bigal area, and for their facilitating access to the habitat. Mauricio Vega provided assistance in arranging the expedition to Ecuador. David Maddison, Mauricio Vega, and Marco Reyes assisted with collecting. The Ecuadorian Ministry of the Environment and the Museum of Zoology of the Pontificia Universidad Católica de Ecuador assisted with permits. Geneviève Leduc-Robert and Edyta Piascik performed DNA extractions and PCR for sequencing. Dmitri Logunov provided assistance in identifying some of the Lyssomanes species. Dmitri Logunov, Gustavo Ruiz and Tamas Szűts provided constructive suggestions on the manuscript. D. Logunov suggested the possible relationship between S. bigal and L. tarmae. Funding was provided primarily from an NSERC Discovery grant to the author, with some funding for the 2010 field work supplied by David Maddison from the Harold E. and Leona M. Rice Endowment at Oregon State University.